/ /? ^ t- o HARVARD UNIVERSITY Library of the Museum of Comparative Zoology Tlie Great Basin Naturalist VOLUME 40, 1980 Editor: Stephen L. Wood Published at Brigham Young University, by Brigham Young University "b 0) n. TABLE OF CONTENTS Volume 40 Number 1 - March 31, 1980 Some aspects of succession in the spnice-fir forest zone of northern Utah. David J. Schimpf, Jan A. Henderson, and James A. MacMahon 1 Utah flora: Malvaceae. Stanley L. Welsh 27 I'tah flora: miscellaneous families. Stanley L. Welsh 38 The taxononiic status of the rosy boa LicJianura roseofiisca (Serpentes: Boidae). John R. Ottley, Robert W. Murphy, and Geoffrey V. Smith 59 Hesperoperla hoguei, a new species of stonefly from California (Plecoptera: Perlidae). Richard W. Baumann and Bill P. Stark ...! 63 Reproduction in three sympatric lizard species from west-central Utah. John B. Andre and James A. MacMahon 68 Haplopappus aJpinus (Asteraceae): a new species from Nevada. Loran C. Anderson 73 Miscellaneous plant novelties from Alaska, Nevada, and Utah. Stanley L. Welsh and Sherel Goodrich 78 New genera and new generic synonymy in Scolytidae (Coleoptera). Stephen L. Wood .... 89 The bacterium Thioploca ingrica on wet walls in Zion National Park, Utah. Samuel R. Rushforth, Sheril D. Burton, Jeffrey R. Johansen, and Judith A. Grimes 98 Number 2 - June 30, 1980 Feeding ecology of Gilu l)or(ixo})itis (Osteichthyes: Cyprinidae) endemic to a thermal lake in southeastern Oregon. Jack E. Williams and Cynthia D. Williams 101 First record of the pallid bat {Antwzuiis pallidus) from Montana. Jeff Shrver and Dennis L. Flath ' 115 A CJuracanthiimi spider bite. Dorald M. Allred 116 Identity of narrow-leaved Chnjsothamnus viscidiflorus (Asteraceae). Loran C. Anderson 117 Ribulose diphosphate carboxylase activities in cold-resistant common mallow, Malva ne- glecta Wallr. and a cold-sensitive tomato, Lycopersicon esculentum L., Ace 55 var. William R. Andersen and Jack D. Brotherson 121 Recovery of Gambel oak after fire in central Utah. L. M. Kunzler and K. T. Harper 127 Relationships among total dissolved solids, conductivity, and osmosity for five ArtemUi habitats (Anostraca: Artemiidae). Nicholas C. Collins and Gray Stirling 131 Spawning of the least chub {lotichtlufs pJilegethontis). Thomas M. Baugh 139 Transferrin polymorphism in bighorn sheep, Ovi.s canadensis, in Colorado. Patrick W. Roberts, Donald J. Nash, and Robert E. Keiss 141 The genus Eriogonum Michx. (Polygonaceae) and Michel Gandoger. James L. Reveal 143 Parasites from two species of suckers (Catostomidae) from southern Utah. J. Craig Brein- holt and Richard A. Heckmann 149 Soil water withdrawal and root distribution under gnibbed, sprayed, and undisturbed big sagebnish vegetation. David L. Sturges 157 Swarming of the western harvester ant, Pogononiywiex occidentalis. Dorald M. Allred ... 165 Relationship between environmental and vegetational parameters for understory and open-area communities. William E. Evenson, Jack D. Brotherson, and Richard B. Wilcox 167 Seasonal activity pattern ot Columbian ground squirrels in the Idaho primitive area. Charles L.' Elliott and Jerran T. Flinders I'^S Habitat and plant distributions in hanging gardens of the Narrows, Zion National Park, Utah. Ceorge P. Malanson 1^^ Short-term effects of logging on red-backed voles and deer mice. Thomas M. Campbell III and Tim W. Clark 183 Terminal bud formation in limber pine. Ronald M. Lanner and James A. Bryan 190 Stinger utilization and predation in the scorpion Paniwctonus horeus. Bmce S. Gushing and Anne Matherne 1^'^ Number 3 - September 30, 1980 Spatiotemporal variation in phenology and abundance of floral resources on shortgrass prairie. V. J. Tepedino and N. L. Stanton 197 Doe owners and hvdatid disease in Sanpete County, Utah. Peter M. Schantz and Ferron L. Andersen -i" New grass distribution records for Arizona, New Mexico, and Texas. Stephan L. Hatch .. 221 A comparison of epiphytic diatom assemblages on living and dead stems of the common grass Phr(iW 4a2N 4j<^JV ii 1 1 1 1 1 1 1 1 1 1 1 1 1 rti Fig. L Map of northern Utah, locating th. .School Forest (SF«). Garden Citv Su„n„il .CC ^ ' and the Inuus ot- unshacled lexeept Hear l.akei and honndaries at I.5()(), 2 KM), and 2700 m. March 1980 SCHIMPF ET AL.: SpRl CE-FiR SUCCESSION onlv a few widely spaced spruce older than 275 vears. The oldest known trees are 367 vears (spruce) and 278 years (fir) (T. W. Dan- iel, pers. conmi.). Scattered individuals or small groups of lodgepole or aspen occur in the spmce-fir stands. Within this broad for- ested area are small meadows fringed with aspen clones on various fractions of their per- imeters (Fig. 2). Young spruce and fir are fre- quently observed in these aspen stands but seldom in the meadows, except on the north margins of conifer stands, including the occa- sional small clumps of mature spruce or fir within the meadows. No significant logging has taken place on the site. Cattle and sheep have grazed the site since around 1900. The zonal climax vegetation of the studv area be- longs to the Abies lasiocdipa / Pcdicularis mcemosa climax comniunit\ tvpe and habitat type (Henderson et al. 1976). The life-form composition of the vegeta- tion is rather simple, consisting mainlv of herbs and, except in the meadows, trees. Indi- viduals of shrubbv species are uncommon; shrub biomass is less than 1 percent that of herbaceous biomass in meadows and under aspen, and less than 5 percent under spruce- fir. Small tree species and lianas are absent. These features distinguish the sere under study from those on most other forest succes- sion research sites, especially those in the de- ciduous forest biome. Cryptogams occur, but are not a conspicuous element of anv stratimi of the vegetation, except for periodic emer- gence of basidiocarps in the forests. The gen- eral aspect of the several categories of vege- tation is illustrated in Figure 3. For a discussion of the compositional and environ- mental relationships of this forest to other lo- cal forests, see Henderson et al. (1976) and Lawton (1979). A preliminary stud) of the structure of over 100 stands on environmentally equiva- lent sites in the School Forest (Sperger and Henderson, unpubl.) indicated that there are probably four major pathways of succession leading to spruce-fir forests (Fig. 4). Pathway "1" represents succession following the de- struction by fire of a forest containing signifi- cant aspen root biomass. The aspens sucker within a short period of time to produce an aspen-dominated stand. Spruce and fir sub- secjuently invade, and eventuallv outlive and replace the aspen. Pathway "2" occurs in the lower elevation- al range of the habitat type around the edge ot the School Forest area. If aspen is not lo- cally abundant, then lodgepole is the postfire pioneer, provided that a local .seed source ex- ists. Lodgepole cones are not serotinous in this area, so seeding is from adjacent stands. Spruce and fir establish and grow more slow- ly than lodgepole; thus, a pine-dominated ecosystem exists for some time prior to spmce-fir stand recovery. Spruce and fir may establish soon after fire, without site amelio- ration by aspen or lodgepole, if a significant quantity of unburned woody material re- mains as protection. This is pathwav "3," where the climax species establish without preclimax tree species. The climax stand structure typically takes less time to develop than through pathways "1" and "2." Pathway "4," the most common, and that which we studied intensively, begins with long-persistent meadows, probably not of fire origin. These are eventually invaded by a.s- pen suckers followed by spruce and fir. Typi- cally the fringe of aspen clones is dis- continuous around the meadow, yielding iiregular patterns of meadow invasion (Fig. 2). Once aspen is present, the time frame of events resembles that of the other three path- ways. However, the sere as a whole mav take hundreds of years longer, because of the long-persistent meadow stage. Pathway "4" was studied by sinmltaneous investigation of plots characterized as mead- ow, aspen, fir, and spruce-fir. The fir-domi- nated plots represent a stage, containing some spruce, which sometimes occurs be- tween occupancy by aspen and the t\pical spruce-fir mixture. The inference of these plots as a chronosequence seems reasonable, based on the minimal relief of the studv site and its uniform soil parent material. Because the time since the meadows were last forest- ed could not be determined, the stands can- not be positioned on an absolute time axis. Common herbaceous species in each stage are listed in Table I. These represent samples from one series of stands at Big Meadow and should not be interpreted as ranks for the study site in general. Eriogonum is semi- shnibbv rather than wholh' herbaceous. A Great Basin Naturalist Vol. 40, No. 1 Fii;. 2. Aoiial pliolo iiicliicliii<4 Ihc Ikul'^i-r station in the Stliool Forest. Lower dianrani inclieates type ol forest various sections of tlie photo. .Xrrow jxjints to tlie weather station. March 1980 SCHIMPF ET AL.: SpRUCE-FiR SUCCESSION substantial change in species composition oc- curs along the siiccessional secjuence, with no species being abundant in more than two stages. The niunber of species is lowest in the meadow and highest in the preclimax aspen and fir stages, based on equal sampling effort. The proportion of short-lived species in the species list declines onlv slightly along the sere. The semishrub Vacciniuin scopdiiuin (Leiberg), perhaps the most abundant under- storv species in the Rocky Mountain sub- alpine /.one (Daubenmire 1978), was not found on the site. Mannnal species of the four stages are list- ed in Table 2. The Northern Pocket Gopher {Tliomoniy.s falpoidcs) is abundant in the meadows, producing considerable distur- bance of the upper portions of the soil hori- Fig. 3. Photographs showing general physiognomy of four stages of succession: meadow, aspen, fir, spruce-fir. Pho- tos courtesy of D. .\ndersen. 8 Great Basin Naturalist Vol. 40, No. 1 zons. The amount of bare ground between meadow plants is substantial and largely due to gopher activity. Gophers are less abundant under aspen and rare imder conifers. Their effect on conifer invasion rate is not well un- derstood, for though they can directly de- stroy young spruce or fir, their activities may also reduce herbaceous species competition with tree seedlings. Gophers may also create more favorable .seedbeds for conifers by ex- posing mineral soil. The dominant mammals in the coniferous stages are the Snowshoe Hare {Lepus ameri- canus) and the Red Squirrel (Tamiasciiirus hudsonicus). Fall and winter browsing by hares may retard conifer stand development (Baker et al. 1921); an individual may con- sume some 300 g (fresh weight) of woody stems 4 mm or less in diameter daily in win- ter (Pea.se et al. 1979). Hare population den- sities in this area fluctuate little by com- pari.son with their boreal counterparts (Dolbeer and Clark 1975). The Red Scjuirrel harvests spruce and fir DISTURBANCE (fire) GRASS -FORB MEADOW ASPEN LONG PERSISTING MEADOW LODGE POLE PINE ASPEN FIRS SPRUCE SPRUCES FIR SPRUCE / FIR SPRUCE DOMINATED CLIMAX WITH FIR AS A SUBORDINATE. SPARSE UNDERSTORY OF Pedicularis racemosa Fi^. 4. Summarv ol types of succfssioiial patli\\a\> tli()in;lil to \w operative on liie Scliool Forest. cones before the seeds are shed, transporting them to large caches at shady bases of trees. Finlev (1969) concluded that Red Squirrels harvest almost all the cones produced in vears of poor to average seed output, and that only in high seed production years are enough .seeds dispersed that significant tree recruitment is possible. Further information on the mammalian component of the ecosys- tems is in Andersen et al. (1980). Bird .species of the four stages are listed in Table 3. Only the spruce-fir data are from the site; preceding stages were inventoried in more extensive stands nearby. Meadows fea- tiue low numbers of both species and individ- uals. Conspicuously absent from the co- niferous stands is the Gray (or Canada) Jay (Perisoreus canadensis), a common per- manent resident throughout most of the Rocky Mountain .subalpine zone. Similarity in composition of the vertebrate species among stages (Jaccard coefficient of commu- nitv) and its variation between years in the case of the avifauna are shown in Figure 5. Further avian information is available in Smith and MacMahon (submitted). The comparisons of species similarities for all vertebrates (Fig. 5) across the .sere show that the most mature stage, spiTice, is most different from the least mature stage, mead- ow. The most similar stages are the two con- ifers, spruce and fir. All pair-wise serai stage comparisons of mammals, though showing these patterns, are more similar than those for birds. The implication is that the birds, as we would expect, respond more dramatically to the physiognomic changes attendant to de- veloping from meadow to a deciduous forest and finally to a coniferous forest. Mammals, on the other hand, respond to the presence or absence of trees, but do not vary as much with tree species or leaf habit. The verte- brate species count of 13-16 mammal .species and 20-30 bird species for each of oiu' serai stages correspond to mammal and bird spe- cies counts from similar communities through North America and also northern Europe (e.g., Erskine 1977. Han.sson 1974, Jiirvinen and Viiisanen 1976, Sabo and Whittaker 1979). The most abundant insects in the meadows are .species of aphidids and thripids. Follow- ing a winter drought, the aphids declined March 1980 Sc:himpf et al.: Spruce-Fir Succession 9 precipitously in 1977, when two species of cicadelhds were the most abundant. A cica- delHd is the most abundant in the aspen un- derstorv, followed bv three species of thrip- ids. In the aspen canopv the most common species is a serpentine leaf miner (Gracila- riidae), followed bv an aphidid, a cicadellid, and a blotch mining gracilariid. In the con- ifer understory the most common species is an aphidid, followed by two thripids and two more aphidids. During 1977 the aphids were scarce and a cicadellid was the most abun- dant. In the conifer canopy, the most abun- dant species are an encyrtid, a mirid, a thrip- id, and an eriosomatid. The insects on spruce were observed to be verv similar to those on fir. Among soil and litter metazoan in- vertebrates, numbers increase markedly along the serai sequence. The relative abun- dance of Collembola and the plant-feeding nematodes (Tylenchida, Dorylaimida) does not change much. The forested stages harbor increased proportions of bacterivorous nema- todes (Rhabditida), detritivorous mites (Ori- batei), and predaceous mites (Mesostigmata). Oligochaetes are essentially absent, a charac- teristic of the region (Gates 1967), and gas- tropods are also rare on the site. Populations of protozoans were not estimated. Soil micro- organisms also exhibit large absolute increas- es in number through successional time and are generally highest in density in the conifer litter. Estimates of abundance for several Table 1. Herbaceous species comprising 5 percent or more of the mean daily herbaceous bioinass in samples at the Big Meadow succession stages. 1977. Numbers following a taxon correspond to its biomass rank, with percent of the herbaceous biomass indicated for top-ranked species. Number of various categories of species samples and her- baceous biomass in relation to tree leaf biomass are also included. Only herbaceous tissue of the semishrub Eriogo- nititi is included. Meadow .^spen Fir Spruce-Fir Achillea miUefoUiim L. ssp. kinulosa (Nutt.) Piper Agropyron trachijcaiilum (Link) Malte var. lati^himc (Scribn. & .Smith) A. \. Beetle Agropijron tracJiijcauhnn (Link) Malte var. ghiiicuiu (Pease & Moore) Malte Aster engelmannii (Eat.) Grav Bwmiis cciriudttts Hook & .\rn. Dcscurainia nchdidsonii (Sweet) Schidz var. sonnet (Robins.) C. L. Hitchc. Erigeron speciosus (Lindl.) D. C. var. niacninthiis (Nutt.) Croncj. Eriogonum herarleoidcs Nutt. Cilia aggregata (Pursh) Spreng. Ligustiriim filicinitni Wats. Lnpinits argenteus Pursh var. ruhricanlis (Greene) Welsh Pediciilahs raceinosa Dugl. var. alba (Pennell) Cronq. Poa nervosa (Hook.) Vasey var. wheeleri (Vasev) C. L. Hitchc. Potentilld argiita Pursh var. convallaria (Hvdb.) Tli. Wolf Riidheekia ocridenlidis Nutt. var. ocridentcdis '^eneeio crasstiliis (Jrav Seneeio serra Hook. Stellaria jamesiana Torr. Trisetum spicatum (L.) Richter Total number of species sampled Total number of annual species sampled Total number biennial/short-lived perennial species sampled Herbaceous standing crop H.S.C. -(- tree leaf standing crop 1 (29%) 25%) Rank 3 1 (25%) 1 (26%) 7 3 6 2 6 41 62 58 49 8 10 7 8 2 4 3 2 100% 2% 0.1% 0.06% 10 Great Basin Naturalist Vol. 40, No. 1 phylogenetic and functional groups are sum- marized in Table 4. Methods of Site Analysis Field studies were conducted during 1976, 1977, and 1978. From 1970 through 1976 air temperatures and precipitation had been re- corded at a station (hereafter the Badger sta- tion) near the edge of a small meadow on the study site at 2650 m elevation (Fig. 2). Mean monthly values recorded in Lomas (1977) were used as dependent variables in multiple linear regression models that employed as in- dependent variables the values for the same months at nearby lower elevation stations, for which long-term means are available. For both precipitation and temperature models these stations are Logan, USU (elevation 1458 m, 27 km SW of the site) and Laketown (1825 m, 16 km E-SE); data from Richmond (1426 m, 25 km W-NW) were also used in the precipitation models (see Fig. 1). These data are published in the corresponding years of Climatological Data, Utah (U.S. Weather Bureau, U.S. Department of Commerce, Washington, D.C.). The published mean val- ues for these stations for the 1941-1970 peri- od were entered into the appropriate regres- sion models to produce an estimated monthly mean temperature or precipitation total for MAMMALS 1976-77 BIRDS 1976 BIRDS 1977 Fig. 5. Diagrams showing degree of affinity among vertebrate assemblages in the four serai stages. Number between stages is the Jaccard coefficient of community. M = meadow, A = aspen, F = fir. S = spruce-fir. T.\BLE 2. .Mammal species observed or trapped in various examples of the serai stages at the School Forest from 1976 through 1978. Meadow .\spen Fir Spruce-Fir Cervus canadensis (Elk) Cletliriotwinys ^apperi (Boreal Redback Vole) Erctliizon dorsatuin (Porcupine) Eutamki.s niiniiutt.s (Least Chipmunk) Eiitaiiiias tind)rinus (Uinta Chipmunk) Glaucomijs sabrinus (Northern Flying Scjuirrel) Lagurus curtatus (Sagebrush Vole) Lepus umericanus (Snowshoe Hare) Mu.stela crminea (Shorttail Weasel) Mustela frenata (Longtail Weasel) Neotoma cinereu (Bushytaii Woodrat) Odocoileus heiniontts (Mule Deer) Pcromijsnts manicttlatus (Deer Mouse) .Sorc.v sp. (Shrew) Spcnnophilus latemlis (Colden-maiitled Stjuirrei) Tainiascittrus hudsoninis (Red Sfjuirrel) Ttiomomys talpoidcs (Northern Pocket Copher) Zapus princeps (Western Jumping .Mouse) Total X X X X X X X X X X X X X X X X X X X X X X ■p X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 13 16 14 1.3 March 1980 SCHIMPF ET AL.: SpRUCE-FiR SUCCESSION 11 the Badger station for the same 30-vear peri- od. Air temperature, relative humidity, and precipitation were recorded and pan evapo- ration measured during the summers of 1976, 1977, and 1978 at two other stations (eleva- tion 2560 m) 1.5 km \-NE of the Badger sta- tion. One station was located in a meadow about 20 m from the edge of an aspen stand; an identical one was situated about 80 m into the forest from the first, under a well-devel- oped spnice-fir canopy. These are referred to as Big Meadow (meadow) and Big Meadow (conifer), respectively. Instruments used were tipping bucket recording precipitation gauges (unshielded), recording hvgrothermo- graphs in standard shelters, and U.S. Depart- ment of Agriculture Class A evaporation pans. Pan evaporation was measured at ir- regular intervals which averaged about 7 days. The instruments were mounted on a platform about 2.5 m above ground, higher than normal because of the deep snows which cover the area. Snowpack dynamics were inferred from periodic observations at the site, in conjunction with measurements made by the U.S. Department of Agriculture, Soil Con.servation Service, at Garden City Summit, 8 km north of the site (elevation 2400 m) (Fig. 1). Stem xylem water potential (predawn) of conifers along a 10 X 50 m transect from meadow through aspen to conifer was mea- sured with a FMS pressure bomb (Waring and Cleary 1967) on 3 August 1977. At least three different branches of each spruce or fir were measured; these trees were all about 1 m tall. Tree species population structure along this same transect was assessed by aging stems and estimating their heights. Ages were determined by counting terminal bud scale scars or annual growth rings in xy- lem cores. In the latter case, the age at the coring height (1.37 m) was corrected to total age by adding 5 years for aspen and 20 years for fir or spruce, values based on average number of bud scale scars at that height. Soil pits were dug to 1 m depth in two ex- amples of each stage of the sequence in the same stands being sampled for other ecosys- tem attributes. Each soil horizon was identi- fied, measured, and sampled prior to phvsical and chemical analvsis in the laboratory. Per- cent coarse fragments was estimated bv eye in the field. Composition of the fine particle fraction was determined by the hydrometer method and with sieves. The following chem- ical parameters were measured: pH (saturat- ed paste), organic carbon (Walkley-Black procedure), cation exchange capacity (sodium acetate method), total cations (ammonium acetate), potassium and phosphorous (sodium bicarbonate), iron (DTPA extraction), and to- tal nitrogen (Kjeldahl). Each of the eight pe- dons studied was classified to the Great Group level using the system of USDA Soil Conservation Service (Anonymous 1975). Soil moisture was monitored in 1977 and 1978 at the Badger station as a continuation of previous work (Lomas 1977). Volume per- cent moisture was measured with a Troxler neutron probe in five tubes in a meadow and ten tubes in a spruce-fir stand. Values were recorded for each 30.5 cm increment of a 122 cm deep profile. Results Estimated 1941-1970 monthly mean tem- peratures and precipitation totals for the Badger station are reported in Table 5, along with the multiple coefficient of determina- tion for the regression model by which each estimate was derived. These estimates are plotted against the monthly means (1941-1970) for three weather stations in the Rocky Mountain subalpine zone with long- term records (Figs. 6, 7). Most of the precipi- tation at Badger is received as snow, with nearly 30 percent of the annual precipitation falling in December and January. Rainfall de- creases to very low amounts in Julv, rising somewhat in August. Long-term snowpack records for Garden City Summit are present- ed in Figure 8. Meadow peak snowpack depths at the study site averaged about 40-50 cm greater than those at Garden City Sum- mit during the corresponding winters. Selected temperature, humidity, and pre- cipitation data from the Big Meadow station for the smnmers of intensive studv are pre- sented in Table 6. Pan evaporation for these summers is plotted in Figure 9 against days after snowmelt in the meadows; these can be converted to calendar dates from the snow- melt dates in Table 6. The evaporation 12 Great Basin Naturalist Vol. 40, No. 1 curves were integrated planimetrically over various time intervals (Table 7). Over the in- terval from 43 to 73 days after snowmelt, evaporation in the conifer forest averaged 38 percent (1976), 46 percent (1977), and 46 percent (1978) as great as that in the mead- ow. Soil moisture trends at the Badger station during the three summers of intensive study are represented in Figure 10. From these data we can generally charac- terize the three summers during which eco- system properties were analyzed. During 1976 perhaps the most favorable conditions for plant growth and development occurred, with above average July rain, low evapo- ration, and a moderately long frost-free peri- od. The summer of 1977 followed an extraor- dinarily dry winter, resulting in low soil moisture content. The growing season began very early and was much longer than for the other two summers. June and August were warmer than in 1976 and 1978, but not July. Pan evaporation was very high, perhaps in part because it was measured during the longer days of early summer, since the satura- tion deficits were not especially great. Ex- tremely high August rainfall resulted from a rare deep continental intrusion of a tropical storm. The 1978 frost-free season began late and ended early. Temperatures and evapo- ration rates were intermediate, and satura- tion deficits were relatively low in June and July but high in August. Less rain fell than during the other two summers. Tree water potential and forest height structure along the meadow-aspen-conifer Table 3. Status of avian species observed on study areas in 1976 and 1977. P = permanent resident; B = sum- mer breeder; F = feeder in serai stage, but not breeder; V = mitirating or wandering visitor; W = winter resident. Species Meadow Aspen Fir Spruce-Fir Turkey Vulture Goshawk Cooper's Hawk Sharp-shinned Hawk Marsh Hawk Red-tailed Hawk Golden Eagle Sparrow Hawk Blue Grouse Ruffed Grouse Mourning Dove CJreat Horned Owl Pygmy Owl Common Nighthawk Poor-will Broad-tailed Hununingl)ird Rufous Hiuiimingbird Red-shafted Flicker Yellow-bellied Sapsucker WilHamson's Sapsucker Hairy Woodpecker Downv Woodpecker Northern Three-toed Woodpecker Lewis" Woodpecker Dusky Flycatcher Western Wood Pewee Olive-sided Flycatcher Violet-green Swallow Tree Swallow Steller's Jay Black-billed Magpie Clark's Nutcracker Common Raven Black-capped C^hickadee Monntairi (Chickadee rare permanent resident rare permanent resident rare permanent resident V V V V B V rare permanent resident V V W W W P W W F V V V p p p V V V V F B V V F F V V F B B B V B B P P P B V P V V B B B? B B B B V F B P P V V V V V V V V V V B P P March 1980 SCHIMPF ET AL.: SpRUCE-FiR SUCCESSION 13 transect is plotted in Figure 11, and age structure in Figure 12. For tabulated age data, see Daniel et al. (1979:286). It can be seen that the water stress of conifers one to two meters tall increases as the aspen is re- placed bv niatiue spruce and fir. The eight pedons sampled are character- ized in Table 8. Their taxonomic assignments may be interpreted as follows: Cryochrepts are soils with cold mean annual temperatures showing little development. Crvoborolls have dark surfaces, cold mean annual temper- atures, and little development of the profile. Paleboralfs are similarlv cold, but have an ar- gillic horizon (accumulation of clav) deep in the profile. This extensive development is in- ferred to be the product of a different cli- mate in the past. Cone crops on the School Forest have been estimated annually since 1947 (T. W. Daniel, Table 3 coutiuuccl. pers. comm.). In 1976 fir and spruce had very low cone abundance, but in 1977 both bore heavy crops. In 1978 the spruce crop was very low, and fir bore a moderate crop. Discussion The greater similarity of summer air tem- peratures than of precipitation patterns among the geographically disparate spruce- fir sites in Figures 6 and 7 lends support to the assertion by Daubenmire (1956) that lower growing season temperatures dis- tinguish this zone from other Rocky Moun- tain forests. A variety of precipitation re- gimes permits the existence of spruce-fir forests, from the summer-dominated precipi- tation south and east in the Cordillera to the winter-dominated patterns north and west. Few other data on humidity or pan evapo- Species Meadow .\spen Fir Spruce-Fir White-breasted Nuthatch Red-breasted Nuthatch Brown Creeper House Wren .\merican Robin Townsend's Sohtaire Hermit Thrush Mountain Bluebird Colden-crowned Kinglet Ruby-crowned Kinglet Northern Shrike Warbling \'ireo Orauge-Crowued Warbler Audubon's Warbler MacGillivray's Warbler Wilson's Warbler Townsend's Warbler Western Meadowlark Brewer's Blackbird Western Tanager Black-headed Grosbeak lazuli Bunting Cassin's Finch .\merican Goldfinch Pine Grosbeak Gray-crowned Rosv Finch Pine Siskin Red Crossbill White-winged Crossbill Green-tailed Towhee \ esper Sparrow IDark-eyed Junco (3 races) Chipping Sparrow Brewer's Sparrow White-crowned Sparrow Lincoln's Sparrow B B P P B P B B B B V B B B W P V B V B B B B V V B B V B V V V B V B B B V B B V w P P V B B B B B V B V V B F B B B F B B B B V B B V V 14 Great Basin Naturalist Vol. 40, No. 1 ration in this zone have been pubHshed. The three-year monthly averages for saturation deficit at Big Meadow (Table 6) are 9 per- cent greater (June), 53 percent greater (July), and 28 percent greater (August) than those we computed from one year of temperature and humidity records for a spnice-fir site in the Front Range in Colorado (Marr 1967). Pan evaporation rates in 1918 and 1919 in the subalpine of Arizona (Pearson 1931) were far less than ours (Table 7) in both forested and exposed sites. The Arizona site receives large amounts of summer rain, and may expe- rience lower saturation deficits. Summer davs o o UJ a: UJ Q. u — BRIGHTON --B-- BADGER LEADVILLE — -SUMMIT Fig. zones: mates; JFMAMJJASOND 6. Mean niontlilv temperatures during 1941-1970 period for four sites in the Rocky .Mountain spruce-fir Badger station, Utah; Brighton, Utah; Leadville, Colorado; Summit, Montana. Means for Badger are esti- those for the other sites are measured parameters. Table 4. Relative abundance of phylogenetic and fimctional groups of soil microorganisms. Total number of or- ganisms in the phylogenetic and functional group coimts do not agree because different methods were used; func- tional groups are also not nuitually exclusive. Soil samples are from topmost 5 cm. l(P organisms g"^ dry substrate Percent of total Aerobic bacteria Streptonivcetes Fungi Anaerobic bacteria Sample type Summer/Fall Summer/Fall Summer/Fall Summer/Fall Summer/Fall Meadow soil 2.9/5.6 66/64 32/34 2/1 1/1 -\spen soil 4.9/9..3 68/68 29/29 3/2 0.2/0.2 Fir soil 7.7/14.7 77/78 20/20 3/2 0.2/ < 0.1 Spruce-fir soil 7.5/10.7 77/70 20/26 3/4 <0.1/<0.1 Fir litter .39.6/28.0 86/84 13/14 1/2 <0.1/<0.1 Spnice-fir litter 15.0/22.0 77/84 21/13 2/2 <0.1/<0.1 March 1980 SCHIMPF ET AL.: SpRUCE-FiR SUCCESSION 15 15 1 BRIGHTON o lOH I- < ^ 5- a: a. 0- LEADVILLE F M A M A S N D Fig zone ^.7. Mean monthly precipitation totals during 1941-1970 period for four site.s in the Rockv Mountain spruce-fi Badger station, Utah; Brighton. Utah; Leadville, Colorado; Summit, Montana. Means for Badger are estimates those tor the other sites are measured parameters. Table 4 continued. Proteolytic Summer/Fall l(fi organisms g-^ dry substrate Hemicellulolytic Summer/Fall Chitinolytic Summer/Fall Lipolytic Summer/Fall Cellulolytic Summer/Fall 0.8/1.3 0.8/.3.0 1.5/1.8 1.2/1.4 3.6/3.7 1.8/2.0 0.1/0.02 0.3/0.08 0.9/0.4 1.0/0.4 2.6/1.3 1.2/1.4 0.3/0.6 0..3/0.7 0.4/0.3 0.4/0.3 0.6/0.5 0.6/0.4 0.08-0.04 0.2/0.07 0.3/0.1 0.3/0.1 0.4/0.8 0.4/0.2 0.1/0.01 ().()6/().()3 0.07/0.09 0.09/0.06 0.2/0.2 0.3/0.2 16 Great Basin Naturalist Vol. 40, No. 1 80 ^^ E e 60 5 40- o Q. < u 20 < a. 40 —J— 50 60 70 80 90 00 DAYS AFTER SNOWMELT Fig. 8. Pail evaporation at the Big Meadow conifer (C) and meadow (M) stations during summers 1976, 1977, 1978. Time is expressed as days after snowmelt in the meadow. Evaporation is plotted as mm wlr* at the midpoint of the measuring period, with curves fitted bv eve. Ciuves not labeled bv vear are three-vear averages. a. LU Q O <: Q_ O C/) 200 1 1 50 - 100 - 50 DEC JAN FEB MAR APR MAY Fig. 9. Time course of snowpack depths at ('.arden City Summit. Broken lines indicate records from the 196()s, and solid lines indicate records from the 197()s. Years labeled are those in which the particular snowpack melted. March 1980 SCHIMPF ET AL.: SpRUCE-FiR SUCCESSION 17 would also be shorter there. These com- parisons, coupled with the paucity of summer rain at our site (Fig. 7), lead us to venture that ours is a comparatively dry example of spruce-fir forest. Our pan evaporation statistics indicate that animals and plant shoots in the lower synusia of the forest should have lower rates of water loss than those in the meadows (Table 7, Fig. 8). Because saturation deficits were nearly the same at both Big Meadow stations, the differences nnist be largely due to differences in net radiation or mean wind velocity. The evaporation rates measiued in the meadow compare favorably with those estimated for tlie Badger station in 1970 bv use of Pen- mans combination method (Eaton 1971). Conifers of comparable small size exliibit greater water stress in conifer-dominated stands than in aspen stands (Fig. 11). We in- terpret this to mean that competition for soil water from large conifers is more severe than from aspen or small conifers. From the pan evaporation data (Fig. 8) we would expect greater evaporative stress under aspen rather than conifers, so the low xylem potentials un- der conifers must be due to inability to ob- tain sufficient soil water. This is probably an important factor limiting recruitment in ma- tiue spruce-fir stands. The tree population age structures along the transect (Fig. 12) indicate that aspen in- vades the meadows at an average of about 19 cm yr', and is followed in about 20 years by successful conifer establishment. The results o o > 50 25 0-305 cm 50-1 - 25- 50 25 A. ^^M77 ^C __M.C76 30.5-61.0 cm ^^ M77 ^^ C76 "I r :)U- ^ M78 61.0-91.5 cm '^?-^^^^^=^'^^^-i'»S3E_,^^^J^J^ M76 25- ^077 C76 0^ T r 1 1 1 JUN JUL AUG SEP OCT Fig. 10. Soil moisture in the meadow and conifer stages at Badger station during the summers of 1976, 1977, and 1978. Volume percent water content is ex- pressed for four .successive 30.5 cm horizons of the soil profile. T.\BLE 5 Estimated mean monthly temperatures and monthly precipitation totals for the Badger station during the period 1941-1970. R^ is the coefficient of determination for the multiple regression model bv which each value was estimated. Month Precipitation, cm R2 13..5 0.74 8.9 0.03 12.7 0.73 10.4 0.39 7.1 0.81 5.2 0.72 1.2 0.61 3.0 0.68 3.4 0.95 6.2 0.68 9.2 0.90 14.3 0.73 Temperature, °C R2 [anuarv Fehniarv March April May June Julv .\ugust September October .November December Annual total or mean 95.1 -10.2 -9.0 -5.3 -0.2 2.9 8.7 14.5 13.4 8.6 3.9 -5.0 -9.7 1.1 0.64 0.55 0..39 0..38 0.69 0.79 0.45 0.96 0.61 0.53 0.62 0.79 18 Great Basin Naturalist Vol. 40, No. 1 presented are in general agreement with oth- er transects on the study site. Altered slopes in the aspen age vs. transect distance rela- tionship suggest that advance of the clones was slowed during the 1930s and accelerated in the past decade. The 1930s were generally warmer and drier than the succeeding period, and the precipitation since 1970 (Lomas 1977) generally exceeded the 1941-1970 esti- mated means. Aspen dominates a plot on the School For- est for 100 to 150 years before spruce or fir assumes dominance (Fig. 12) and hastens the demise of aspen through shading. Persistence of occasional aspen stems in more open por- tions of the spruce-fir canopy is important from the standpoint of regeneration poten- tial. The transition from aspen to spruce-fir occurs over a relatively short time and dis- tance (Fig. 12), explaining why we were i2 -20n < -15- LU O _J X -10- LU ^ -5i spruce Spruce y oFlrV r30 h20 E X LlI hlO ^ 10 To 30^ — I — 40 — I — 50 DISTANCE FROM MEADOW (m) Fit;. 11. Average tree height of aspen, fir, and spruce populations, and water potential of short conifers along a meadow-aspen-conifer transect. Water potential (^) of fir and spruce of about one meter height was measured .3 August 1977. Values are plotted against distance from the edge of a meadow. Table 6. Climatological data from Big Meadow weather stations. Precipitation is average of meadow and conifer station totals. Snowmelt is first date when meadow snow cover v.as estimated to be less than 10 percent. Saturation deficit is computed from weekly mean temperature and mean relative humidity for the meadow station. .\ir tempe ature , °c Snowmelt Mean High Stage/year June July August Meadow 1976 7.8 13.7 9.9 24.7 June 4 1977 10.9 12.7 11.7 23.3 Mav 15 1978 - 12.6 10.9 25.6 June 16 Conifer 1976 8.7 14.6 10.6 27.8 1977 12.0 13.9 12.4 26.7 1978 8.8 13.9 12.0 26.1 March 1980 SCHIMPF ET AL.: SpRUCE-FiR SUCCESSION 19 unable to find mixed aspen-conifer stands sizeable enough to study. Tabulation of the ages of trees by species indicates that the in- vasion of spruce and fir occurs with less tem- poral regularity than aspen invasion of mead- ows (Daniel et'al. 1979:286). The great similarity in soil physical and chemical properties among the four stand types (Table 8) strengthens our view that the stands differ mainly as a consequence of suc- cession and not because of differences in site potential. Most of the edaphic differences ex- isting among stages are manifested near the soil .surface, reflecting the influences of the current resident biota. The absence of char- coal in the upper few cm leads us to con- clude that fire has not been a major force in the meadow ecosystems during the past few centuries. 10 20 30 40 DISTANCE FROM MEADOW (m) Fig. 12. .\ge stnicture of aspen, fir, and spruce populations along a nieadow-aspen-tonifer transect. Mean ages are plotted against distance from the edge of a meadow. Table 6 continued. Last First freeze Precipitation, cm Saturation deficit, g nr3 freeze July August June July August June 26 May .30 June 26 June 27 May 31 Jime 26 Aug. 26 Aug. 28 Aug. 16 Aug. 26 Aug. 27 Aug. 15 4.0 2.3 2.2 12.7 0.5 3.5 5.10 4.42 3.54 7.49 5.65 5.21 4.35 5.13 5.49 20 Great Basin Naturalist Vol. 40, No. 1 We must, then, invoke reasons other than fire for the existence of the meadows we studied. We .suggest that we are .seeing the disappearance of subalpine meadows that were chmatic chmax during a colder period. If this is true, then the sequence we describe is botii allogenic (warmer temperatures al- lowing forest expansion) and autogenic (cli- max forest establishment facilitated by aspen invasion) in character. It is important to de- termine whether these meadows are old or yoimg. Even though phvsiognomy and ener- getics may be similar in pioneer meadows and mature meadows, the life history charac- teristics of the organisms and the structure of the communitv mav well differ due to the length of time a stand has been in existence (MacArthur and Wilson 1967). There is evidence from around the globe for cooling during this earlier period (Brav 1971), though we know of no published evi- dence for colder conditions in the immediate vicinity of our site. A dendrochronological study of a site 375 km SW of ours indicates that temperatures were cooler than at pres- ent from 1541 to 1780 (LaMarche and Stock- ton 1974). This corresponds to the dearth of trees older than 275 years on our site, though their absence could be interpreted as the re- .sult of insect devastation (Miller 1970). Whipple and Dix (1979) also found fewer .spmce between ages of 300 and 400 years in Colorado than they expected. Whether this cooling was sufficient to change the general aspect of our studv site we can only gue.ss. There is general agree- ment that the cooling was greatest at high latitudes and altitudes (Bray 1971). The limit- ing summer air temperatures for normal growth of spruce or fir are not well under- stood; Wardle (1968) presented evidence that the limit lies between 11 and 12 C (July) for spruce in Colorado. A July decline of 2.5 C (Table 5) might have been sufficient, in com- bination with the level topography of our site, to produce an open subalpine parkland of herbaceous meadows containing scattered trees (Billings 1969). The wide dispersion of old conifer individuals on the School Poorest is reminiscent of the tree patterns in such park- lands. Forest expansion since this colder peri- od may have been anomalously rapid; if our site reflects climatic trends in llie Rocky Mountains as a whole, then the years 1870 to 1945 may have been the most warm-moist 75-year period since 1130 (Bradley 1976). In conclusion, we would like to reempha- size the integral role played by preclimax trees in the structuring of the climax forest. The successional change we describe is not merely a consequence of the passage of time and the differential growth rates of preclimax and climax species (Dmry and Nisbet 1973). Species characteristic of the climax are not "present but inconspicuous" early in succes- sion; they establish extremely slowly and only on the north margins of their stands in the absence of aspen or lodgepole. This is illustrated in two discrete subalpine basins we have observed. Birch Creek and Summit Creek South Fork, 17 km W of our study site. Lodgepole is absent throughout this western portion of the mountain range at this latitude. These two basins are unusual in that they are also devoid of aspen in the sub- alpine zone. Aspen may have been locally ex- tirpated during the Pleistocene, when both basins held small alpine glaciers (De Graff 1976). Now aspen is unable to invade from adjacent topographic units because its in- ability to reproduce by seed. Aspen is present at lower elevations in these drainages, but perhaps only as the late-leafing form which is not known to occur at higher elevations (Cot- tam 1954). The spruce-fir stands of these two topographic units, at the same elevations as our study site, are small discrete groves in a Table 7. Estimated pan evaporation rates at the Big Meadow stations, expressed as mm wk"'. Means for vari- ous intervals were obtained by planimetric integration ot the turves in Fisrure 8. Stage Period Rate Meadow 1976, 4.3-73 days after melt 40 1977, 4.3-73 davs after melt .56 1978. 4.3-73 days after melt 43 1976, entire curve 39 1977, entire ciuve 50 Three-year mean, entire cur\e 42 (>oniter 1976. 43-73 da>s after melt 15 1977, 43-73 davs after melt 26 1978, 43-73 days after melt 20 1976, entire curve 16 1977, entire curve 23 Three-year mean, entire curve 17 March 1980 SCHIMPF ET AL.: SpRUCE-FlR SliCC.ESSIOX 21 matrix oi herbaceous veij;etation. We h\ poth- esize that this pattern represents exactly the result we would expect for the allo.'5. Co.N.NEi.i.. J. H.. AM) B. (). Si.ATYKH. 1977. .Mechanisms ot succession in natural conniumities and their role in cominunit\ stabilitx and organization. Amer. \atur. Ill: 1119-1144. (.'oTiAM. W. P. 19.54. Prevernal leafing of aspen in I'tah mountains. |. Arnold Arbor. .35: 239-2.50. Danu:!.. T. W.. J. A. lli;i,\is. am) F. S. Baker. 1979. Principles of sihicullure. 2d ed. McCirau-Hill, \e\v '^'ork. I)\\u 1.. T. \\'.. a.m) J. Sc:iiNnnr. 1972. Lethal and non- lethal effects of the organic horizons of forested soils on the germination of seeds from several as- sociated conifer species of the Rockv Mountains. Can. J. For. Res. 2: 179-184. l)\i luwuHK. R. I". 1943. V'egetational zonation in the Bock\ Mountains. Bol. Rev. 9: .32.5-.393. 19.56. Climate as a determinant of vegetation dis- tribution in eastern Washington and northern Idaho. Ecol. Monogr. 26; 131-1.54. 1978. Plant geography, with special reference to North Xnierica. Academic Press. New York. l>i: ('.RAFF, ). V. 1976. Quaternarv geomorphic features of the Bear River Range, north-central Utah. Un- published thesis. Utah State L'niv.. Logan. Dixon. 11. 19.35. Ecological studies on the high plateaus of Utah. Bot. Caz. 97: 272-320. DoLREER, R. A., A.M) \\ . R. Ci.ARK. 1975. Population ecology of snowshoe hares in the central Rocky Mountains. J. Wildl. Manage. .39: .5.3.5-.549. l>Ri R^. \\ . H.. AM) 1. (]. T. \isBET. 1973. Succession. J. .\rnold Arlior. .54: .331-368. Di NWHJDiE. P. W. 1977. Recent tree invasion of sub- alpine meadows in the Wind River Mountains, Wyoming. Arc. Alp. Res. 9: 393-399. Eaton. F. D. 1971. Soil inoisture depletion, actual and potential e\apotranspiration in an Engelmann spruce-subalpine fir forest. Unpublished thesis, Utah State Uni\ .. Logan. Erskine. .\. J. 1977. lihds in boreal Canada: (Commu- nities, densities and adaptations. Can. Wildl. Serv. Rep. Series No. 41. FiM.E'i . R. B.. Jr. 1969. Cone caches and middens of Ta- niia.'icitints in the Rock\ .Mountain region. Pages 2.3.3-273 iti J. K. Jones, Jr., ed. Contributions in mammalogv. Univ. Kansas Mus. Nat. Hist. .Misc. Pnbl. 51, Lawrence, Kansas. FowEi.i.s. H. A. 1965. Silvics of forest trees of the United States. U.S. Dept. Agric. Agric. Hndbk. No. 271. (^ATFs. C;. E. 1967. On the earthworm fauna of the (ircat American Desert and adjacent areas. Crcal Basin Natur. .37: 142-176. Hansso.n, L. 197 L Small mannnal productivity and con- sumption in spruce forest and reforestation in south Sweden. Oikos 25: 1.53-1.56. Hei.i.mfrs, H., M. K. Centiie, a.nd F. Ronco. 1970. Temperature affects growth and development of Engehnaim spruce. For. Sci. 16: 447-4.52. iliNDiRsoN. |. .\., R. L. Maik. D. L. .\nderso.\. R. KeTC lUK. P. I.AWTON, S. SiMO.V, R. H. Spercer, R. \\ . YoiNt.. AM) .\. YoiNCiBEOon. 1976. Prelimi- narv forest habitat types of northwestern Utah and adjacent Idaho. Dept. Forestry and Outdoor Recreation, Utah State Univ., Logan. I1()H\. H. S. 1971. The adaptive geometr\' of trees. Princeton Univ. Press, Princeton, New Jersey. [vHM.NEN, O., AM) R. \. V.\is.\NE.\'. 1976. Between-vear component of diversit\ in communities of breed- ing land birds. Oikos 27:.34-.39. 22 Great Basin Naturalist Vol. 40, No. 1 Table 8. Physical and chemical characteristics, and classifications of two pedons in each of four stages of succes- sion in the School Forest. Roots: f = few , c = common, m = many. vf = very fine, f = fine, m = mediiun, c = coarse. Cation Percent Percent Percent exchange sand, coarse organic capacity Location, depth, cm silt, clay fragments Texture pH carbon meq/lOOg Big Meadow (Meadow \) 0-9 .30,53,17 30 gravel 10 stones gravelly .silt loam 63 1.9 14.2 9-38 30,53,17 30 gravel 10 stones gravelly silt loam 6.0 1.5 15.7 38-ft3 33,47,20 30 gravel 10 stones gravelly loam 5.8 1.1 13.8 63-98 45,.37,18 40 gravel 5 cobbles gravelly loam 5.5 0.3 8.5 98-100 4633,21 40 grave! 5 cobbles gravelly loam 5.4 0.2 9.5 Doc's Meadow (Meadow B) 0-9 35,45,20 less than 10 loam 6.3 1.7 14.8 9-26 .35,43,22 less than 10 loam .5.8 1.4 14.2 26-40 .35,42,23 10 gravel loam 5.5 0.7 11.5 40-67 45,.33,22 30 gravel gravelly loam 5.2 0.3 10.1 67-100 58,26,16 25 gravel gravelly fine sandy loam 5.3 0.2 62 Big Meadow Fir (Fir A) 0-20 .33,56,11 20 gravel gravelly .silt loam 5.8 1.7 18.7 20-62 47,.38,15 30 gravel 10 cobbles gravelly loam 5.7 0.7 9.1 62-88 .3.3,.5611 30 gravel 5 cobbles gravellv silt loam 5.2 0.5 12.2 88-100 52,31,17 40 gravel gravelly very fine sandy loam 5.3 0.3 7.9 Harts Fir (Fir B) 0-10 43,4.3,14 25 gravel gravelly loam 5.7 2.5 9.9 10-.35 40,44,16 30 gravel gravellv loam .5.8 1.0 8.8 .35-62 40,4.3,17 .30 gravel gravelly loam .5.6 0.8 8.4 62-100 .37,.32,31 30 gravel gravellv clay .5.4 0.3 19.4 Big Meadow Aspen (.\spen A) 4-0 litter 0-6 6-27 organic .34,44,22 organic ,5.6 30 gravel gravelly 5.6 20 stones loam 25.3 1.7 29.4 19.4 March 1980 SCHIMPF ET AL.: SPRUCE-FiR SUCCESSION 23 Table 8 continued. 64 Percent base saturation K, ppm 210 P, ppm 16.3 Fe, ppm Roots Charcoal 24 nnf nif July percent total nitrogen & depth, cm Great group 0.11 top 3 Crvohoroll 54 146 52 69 72 33 72 44 6.5 3.9 3.5 3.1 31 40 26 24 cvf cf cvf cf ff ff 0. 1 1 25 0.05 50 0.03 100 64 128 16.0 .34 cf 0.(W top 3 54 69 7.7 45 ff + 0.06 25 65 51 3.9 55 ff 4- - 42 3.3 47 fvf + 0.05 50 71 22 3.2 37 0.03 100 Cr\oc]irept Paleboralf 68 73 82 87 122 50 42 31 5.4 5.0 2.6 34 44 20 nivf inf.fin fc cvf cf.fm fc cvf cf ff 1..35 litter 0.19 top 3 0.11 25 0.07 50 0.05 100 Paleboralf - 182 42.4 - cfjni fc + 1.. 39 litter 0.19 top 3 64 200 .37.7 56 cf.fm fc + 0.06 25 63 148 32.8 51 ff.fm fc + 0.04 .50 87 161 21.0 32 fm + 0.04 1(K) Crvoboroll 62 400 + 192 .30.9 4.4 34 29 invt mf mill nnf lilt. mill mc 0. 1 1 top 3 0.17 25 24 Great Basin Naturalist Vol. 40, No. 1 Kaufmann, M. R., and a. N. Eckard. 1977. Water po- tential and temperature effects on germination of Engelmann spruce and lodgepole pine seeds. For. Sci. 2.3: 27-33. La.Marche, V. C, Jr., and C. W. Stockton. 1974. Chronologies from temperature-sensitive bristle- cone pines at upper treeline in western United States. Tree-ring Bull. 34: 21-45. Langenheim, J. H. 1962. Vegetation and environmental patterns in the Crested Butte area, Gunnison County, Colorado. Ecol. Monogr. 32: 249-285. Table 8 continued. Lawton, p. M. 1979. An investigation of the environ- mental relationships of selected forest habitat types in northern Utah. Unpublished thesis, Utah State Univ., Logan. LeBarron, R. K., and G. M. Jemison. 1953. Ecology and silviculture of the Engelmann spnice-alpine fir type. J. For. 51: 349-355. Lo.viAS, D. A. 1977. Soil water depletion following clear- cutting small plots in a spnice-fir forest in north- ern Utah. Unpublished thesis. Utah State Univ., Logan. Cation Percent Percent Percent exchange sand, coarse organic capacity Location, depth, cm silt, clay fragments Texture pH carbon meq/lOOg 24-44 46,39,15 40 gravel 10 cobbles gravelly loam 5.6 0.5 10.1 44-100 54,29,17 40 gravel 5 stones 5 cobbles gravelly fine sandy loam 5.5 0.4 10.1 Hart's Aspen (.\spen B) 0-9 43,43,14 20 gravel gravelly loam .5.7 1.5 11.2 9-27 44,42,14 20 gravel gravelly loam 5.8 1.3 9.5 27-49 43,43,14 .30 gravel gravelly loam .5.9 0.7 7.6 49-75 44,42,14 30 gravel 10 cobbles gravelly loam ,5.6 0.7 7.3 7,5-100 42,4,5,13 10 gravel loam 5.3 0.6 12.8 Big Meadow Spruce-fir (Spruce-fir A) 3-0 litter 0-19 ,39,48,13 ■30 gravel gravelly loam ,5.8 1.7 12.9 19-47 47-72 43,43,14 62,18,20 72-100 70,8,22 Sinks Road Spruce-fir (Spruce-fir B) 5-0 litter 0-11 29,57,14 30 gravel 10 cobbles 20 stones less than 10 gravelly loam coarse sandv clay loam less than 10 coarse sandy clay loam 15 gravel silt loam .5.5 6.5 5.0 1.0 0.4 0.5 1.3 10.4 7.1 6.2 II -,35 ,35-60 60-85 85-l(K) 29,.57.14 40,46,14 43,44,13 .54.,32,14 20 gravel ,30 gravel •5 cobbles ,30 gravel 5 cobbles 40 gravel 5 cobbles gravel I V silt loam gravelly loam gravelly loam gravelly very fine sandv loam ,5.4 5.4 .5.4 0.7 0.6 0.5 0.3 8.3 7.7 6.4 5.1 March 1980 SCHIMPF ET AL.: SpRUCE-FiR SUCCESSION 25 LfivE, D. 1970. Subarctic and subaljMiie: W'Iumc and what? Arc. Alp. Res. 2; 63-73. M.\c..\rthur, R. H., .a..\d E. O. Wilson. 1967. The theo- ry' of island biogeography. Princeton Univ. Press, Princeton, New Jersey. Marr. ]. \\'. 1967. Ecosystems of the east slope of the Front Range in Colorado. Univ. Colorado .Stu., Ser. in Biol. \o. 8, Bonlder, Colorado. Miller, P. C. 1970. .\ge distributions of .spruce and fir in beetle-killed forests on the White River Table 8 continued. Plateau, Colorado, .^iner. .Midi. Natur. 83- 206-212. Noble, D. L. 1979. Roots of lodgepole pine seedlings reach depth of only 3 to 4 inches their first sea- son. U..S. Dept. .\gric. For. Serv. Res. Note RM- .363. Noble, D. L., anp R. R. Alexa.nder. 1977. Environmen- tal factors affecting natural regeneration of En- gelniann spruce in the central Rockv Mountains. For. Sci. 23; 420-429. Percent l)ase saturation K, ppm P, ppm Fe, ppm Roots Charcoal July percent total nitrogen & depth, cm Great group 64 54 51 4.8 3.2 36 33 cvf cf.fm fc ff 0.09 50 0.06 100 81 86 75 80 .56 51 68 61 286 220 179 172 132 142 90 48 31 35.0 29.0 25.6 24.3 5.6 10.7 9.0 5.5 4.2 27 34 34 42 19 54 51 28 mf.nini -f- cf,cm + cf,cm + cf.cm + ff,fm + mvf mf,cm cc mvf mf,cm cc cvf cf.ftn fc ff Crvoboroll 0.17 top 3 0.06 25 0.03 50 0.03 1(K) 1.29 litter Cryochrept 0.19 top 3 0.07 25 0.07 .50 0.03 1(X) Cryochrept 1.25 litter 38 93 8.8 69 mvf mf,cm fc 0.15 top 3 .36 64 2.1 40 cvf cf.fc + 0.08 25 39 71 1.4 34 cf.cm + 0.04 50 47 33 1.1 29 cf.cm 86 .30 1.2 23 cf,cm 0,03 U)() 26 Great Basin Naturalist Vol. 40, No. 1 Noble, D. L., and F. Ronco, Jb. 1978. Seedfall and es- tablishment of Engelmann spruce and subalpine fir in clearcut openings in Colorado. U.S. Dept. Agric. For. Serv. Res. Pap. RM-200. Odum, E. p. 1969. The strategy of ecosystem devel- opment. Science 164: 262-270. OosTiNG, H. J., AND J. F. Reed. 1952. Virgin spruce-fir forests in the Medicine Bow Mountains, Wyom- ing. Ecol. Monogr. 22: 69-91. Paden, J. W., J. R. Sutherland, and T. A. D. Woods. 1978. Caloscypha fulgens (Ascomyce- tidae:Pezizales): The perfect state of the conifer seed pathogen Geniculodendron pijrifomie (Deu- teromycotina: Hyphomycetes). Can. J. Hot. 56: 2.375-2.379. Patten, D. T. 1969. Succession from sagebrush to mixed conifer forest in the northern Rocky Mountains. Amer. Midi. Natur. 82: 229-240. Pearson, G. A. 1931. Forest types in the southwest as determined bv climate and soil. U.S. Dept. Agric. Tech. Bull. 247. Pease, J. L., R. H. Vowles, and L. B. Keith. 1979. In- teraction of snowshoe hares and woody vegeta- tion. J. Wildl. Manage. 43: 4.3-60. Peet, R. K. 1978. Latitudinal variation in southern Rocky Mormtain forests. J. Biogeogr. 5: 275-289. Ronco, F. 1970. Influence of high light intensity on sur- vival of planted Engelmann spruce. For. Sci. 16: .331-339. Sabo, S. R., and R. H. Whittaker. 1979. Bird niches in a subalpine forest: .\n indirect ordination. Proc. Natl. .\cad. Sci. USA 76:1.3.38-1.342. ScHMiD, J. M., AND R. H. Frye. 1977. Spruce beetle in the Rockies. U.S. Dept. .\gric. For. Serv. Gen. Tech. Rep. RM-49. ScHMiD, J. M., AND T. E. Hinds. 1974. Development of spruce-fir stands following spruce beetle out- break. U.S. Dept. Agric. For. Serv. Res. Pap. RM- 131. Smith, K. G., and J. A. MacMahon. Bird communities along a montane sere; Community structure and energetics. Submitted to Auk. Stahelin, R. 1943. Factors influencing the natural re- stocking of high altitude burns by coniferous trees in the central Rocky Mountains. Ecology 24: 19-.30. Stokes, W. L. 1963. Geological map of northwestern Utah. Washington, D.C. Veatch, A. C. 1907. Geography and geology of a por- tion of southwestern Wyoming. U.S. Geol. Survey Prof. Paper 56. Wardle, P. 1968. Engelmann spruce {Picea engehnannii Engel.) at its upper limits on the Front Range, Colorado. Ecology 49: 48.3-495. Waring, R. H., and B. D. Cleary. 1967. Plant moisture stress: Evaluation by pressure bomb. Science 155: 1248-1254. Whipple, S. A., and R. Dix. 1979. Age structure and successional dynamics of a Colorado subalpine forest. Amer. Midi. Natur. 101: 142-158. Wicklow-Howard, M. C, and J. Skujins. 1979. In- fection of Engelmann spruce seeds by Gen- iculodendron pijriforme in western North Ameri- ca. Mycologia (in press). UTAH FLORA: MALVACEAE Stanley L. Welsh' .■\bstract.— This paper is the third in a series dealing with a revision of the flora of Utah. Treated herein are 9 genera and 23 species, including both coniinonly cultivated, escaped, and indigenous representatives. Proposed new taxa include Sphuemkcdnwssiilariifolia (H. and \.) Rydh. var. inoorei Welsh, Sphaeralcea leptophijUa (Gray) Rydb. var. jancac Welsh, and Sphacnilcca f)sorak>idcs Welsh. This third paper in the .series leading to a of specimens examined by me is indicated revision of the flora of Utah deals with the following the discussion of each species. The small but significant and taxonomically diffi- number in parenthesis is the number collect- cult Mallow family. Especially complex are ed by me. members of the genus Sphaeralcea, ably mon- ographed by T. H. Kearney (1935), and re- Malvaceae Juss. viewed for Utah by J. A. M. Jefferies (1972). Mallow Familv As with previous treatments, the work con- siders not only indigenous species and weeds Herbs or, less commonly, shrubs, usually or established escaped species, but those in- pubescent with branched or stellate hairs, an- troduced species which are commonly grown nual, biennial, or perennial, with mucila- as ornamentals or for other uses. Casually ginous juice; leaves alternate, simple, mostly grown species, such as the okra. Hibiscus es- palmately veined, stipulate; flowers perfect culentus L., have been excluded. Althaea, Hi- (or imperfect), regular, solitary or in thyrsoid hiscus, and Malta are included entirely on cymes, or more or less racemose or pan- the basis of cultivated ornamentals and weeds iculate, sometimes with an involucel of sepal- which have become established in the state. like bractlets; sepals 5, more or less per- Malva neglecta is a pest of cultivated areas. sistent; petals 5, separate, adnate to the Iliamna, Malvastrum, Sida, Sidalcea, and staminal sheath; .stamens numerous, united by Spliaeralcea are represented entirely by in- the filaments (monadelphous); ovary superior, digenous species. Abutilon has one species in- 3- to many-loculed; fruit a capsule or a schiz- troduced and the other native. The number ocarp. 1. Involucel lacking 2 — Involucel of 1 or more bractlets, or if lacking (as in some Spliaeralcea specimens), then the flowers orange (grenadine) 3 2(1). Petals white, pink, or lavender; plants of moist sites, usually at middle and higher elevations Sidalcea — Petals yellow or pink to red; plants of cultivated lands or of arid sites, u.sually at lower elevations Abutilon 3(1). Petals orange or rarely purplish pink; indigenous perennial herbs of arid habitats at middle and lower elevations Spliaeralcea — Petals variously colored, but not orange; indigenous or adventive perennial, biennial, or annual plants or various distribution 4 4(3). Flowers ro.se pink or rarely white; plants indigenous, 7-15 dm tall, perennial, of middle and higher elevations Iliamna 'Life Science Museum and Department of Botany and Range Science, Brigham Young University. Provo. Utah 84602. 27 28 Great Basin Naturalist Vol. 40, No. 1 _ Flowers white, pink, rose, yellow, or other hues; plants differing in one or more ways from above '^ 5(4) Flowers mostly 6-10 cm broad, opening flat; plants tall adventive or cultivated biennials Althaea — Flowers less than 6 cm broad or, if broader then the plants shrubby 6 6(5). Style branches 5, elongate; fruit a capsule; plants low annuals or shrubs Hibiscus — Style branches more than 5, short; fruit a schizocarp; plants annual or biennial 7(6). Style branches filiform, with elongate stigmatic lines; plants annual or biennial Malta — Style branches with capitate or truncate stigmas 8 8(7). Petals yellow, or orange to pink or red; plants annual with awned carpels or subshrubs with unawned carpels Abutilon — Petals yellowish white to lavender or whitish; carpels few to many, not awned; plants spreading annuals or herbaceous perennials 9 9(8). Petals yellow white; leaves reniform-orbicular, merely crenate-serrate Sida — Petals lavender or whitish; leaves palmately cleft, with rounded lobes Malvastrum .\butilon Mill. leafy panicles; involucel lacking; calyx 5- Plants herbaceous, annual or perennial, cleft; corolla yellow to orange pink or red; with stellate or simple hairs; leaves alternate, fruit truncate-cylindric or subglobose, the petioled, cordate at base, not or only obscu- carpels smooth sided, dehiscent nearly to the rely lobed; flowers solitary and axillary or in base; ovules 2 or more per carpel. Plants perennial, with slender spreading or trailing branchlets; carpels 5, lacking awn-beaks; plants rare, known only from Washington County A. parvidum Plants annual, with robust erect stems; carpels usually more than 10, each with a long divergent awn; plants uncommon, in agricultural regions A. theophrasti Abutilon parvulum Gray. Perennial, the stems slender and spreading or trailing, the caudex woody, grayish tomentose with min- ute stellate hairs, the branchlets pilose; leaves 0.5-5 cm long, ovate, cordate basally, den- tate and sometimes obscurely 3-lobed; pe- duncles slender, axillary, 1 -flowered, longer than the leaves; calyx lobes ovate-acuminate, reflexed in fruit; petals orange pink to red or sometimes yellowish, 4-6 mm long; carpels 5, somewhat tomentose, to 8 mm long. Known in Utah only from Veyo, Washington County (Meyer 4111), Colorado to California, and south to Texas and Mexico, 1(0). Abutilon theophrasti Medic. Velvet leaf. Annual, the stems robast, erect, velvety and cinereous with short, soft hairs; leaves 3-10 cm long (from sinus to apex) and as broad or broader, orbicular-ovate, cordate at the base, abruptly acuminate at the apex, velvety pub- escent; peduncles shorter than the leaves, one to few flowered; calyx lobes broadly ovate- acuminate; petals yellow, to about 6 mm long; carpels 10 or more, each with a long di- vergent awn. Adventive weedy species of dis- turbed or cultivated areas, occasional in Utah and Washington counties (to be expected elsewhere); widespread in North America; native to Europe; 3(0). Althaea L. Plants herbaceous, biennial, with coarse stellate hairs; leaves alternate, petiolate. March 1980 Welsh: Utah Flora, Malvaceae 29 cordate at the base, lobed; flowers sohtarv or in racemes; invohicel of 6-9 bractlets, con- nate at the base; calyx 5-cleft; corolla of vari- ous colors; fruit flattened wheellike, invested by the calyx, the numerous carpels separating at maturity. Althaea rosea Cav. Hollyhock. Coarse biennials to 20 dm tall or more, the stems erect, stellate-hairy; leaves (3-) 5- to 7-lobed, mostly 3-15 cm long (from sinus to apex) and often much broader; flowers shortly pedicel- late, 6-12 cm wide or more, variously col- ored, often rose to pink or lavender, or some- times white, usuallv with a dark center; calyx lobes triangular, investing the fruit at matu- rity, the involucel calyxlike; carpels numer- ous, stellate along the margins, and reticulate on the sides, 5-7 mm long. Cultivated orna- mental, persisting and escaping, to be ex- pected in all counties in Utah; widespread in North America; introduced from China; 15(0). Hibiscus L. Plants herbaceous or woody, annual or perennial, with stellate or simple hairs; leaves alternate, petiolate, obtuse to truncate or cordate basally, lobed to incised; flowers ax- illary, solitary; involucel of 5-10 distinct bractlets; calyx 5-cleft, more or less accres- cent in fruit; fruit a loculicidal capsule, the carpels 5; seeds several in each locule. 1. Plants annual; calyx strongly veined; petals cream colored, with a purple center H. trionum Plants shrubs; calyx herbaceous, not distinctly veined; petals variously colored, but usually rose pink to lavender H. si/riacus Hibiscus syriacus L. Althaea; Rose-of-Sha- ron. Shrubs, 20-40 dm tall or more, glabrous or softly stellate-hairv; leaves 2.5-8 cm long, 1.5-6 cm wide, triangular-ovate to rhombic, strongly 3-ribbed, commonly 3-lobed; flowers axillary, 4-7.5 cm wide; bractlets usually 5, linear, about as long as the calyx, glabrous to obscurely hairy; corolla variously colored and often double; fruit oblong-ovoid, to 25 mm long. Cultivated ornamental, rarely per- sisting; widely cultivated in North America; introduced from eastern Asia; 3(i). Hibiscus trionum L. Flower-of-an-Hour. Annual, commonly 1.5-5 dm tall, the lower branches often prostrate, coarsely hispid-stel- late to glabrate; leaves 3-lobed or more com- monly 3- to 5-parted, the main lobes cuneate basally, the middle lobe the largest; flowers solitary, axillary, mostly 3-6 cm wide; bract- lets usually 10, linear, often coarsely hispid, much shorter than the fruiting calyx; corolla cream colored to yellowish, with a purple Ltnter, closing in shade. Weedy species of Lultivated land at lower elevations; wide- spread in North America; adventive from :entral Africa; 8(i). Iliamna Greene Plants herbaceous, perennial, sparingly and minutely stellate-hairy; leaves alternate, pe- tiolate, cordate to truncate basally, the mar- gin lobed; flowers in thyrsoid panicles; in- volucel of 3 narrow, persistent bractlets; calyx 5-cleft; fruit a loculicidal capsule, the carpels many; seeds usually 3 in each locule. Wiggins, I. L. 1936. A resurrection and revi- sion of the genus lUamna Greene. Contr. Dudley Herb. 1: 213-229. Iliamna rivularis (Dougl.) Greene. Wild Hollyhock. {Malta rivularis Dougl. ex. Hook.; Sphaeralcea rivularis (Dougl.) Torr. ex. Gray; Phymosia rivularis (Dougl.) Rydb.). Perennial, the stems few to many from a woody caudex, mostly 7-15 dm tall, minutely stellate-puberulent, green; leaves 3- to 7- lobed, cordate to truncate basally, 2.5-15 cm long (from petiole apex to tip), 2-16 cm broad, the lobes triangular, crenate-serrate, finely stellate; pedicels mostly less than 1 cm long; bractlets linear-lanceolate, shorter than the calyx; calyx lobes 3-5 mm long (to 8 mm 30 Great Basin Naturalist Vol. 40, No. 1 long in fruit); petals rose pink (rarely white), 20-37 mm long; carpels 6-10 mm long in fniit, hispid and stellate. Along streams, on foothills, in mountain bnish, ponderosa pine, aspen, and spruce-fir communities, 1440-2900 m elevation, in Daggett, Davis, Duchesne, Iron, Juab, Piute, Salt Lake, San- pete, Sevier, Summit, Tooele, Utah, Wasatch, and Weber counties; Colorado, Idaho, Ne- vada, and Washington; 40(vi). Malva L. Plants herbaceous, annual, biennial or per- ennial, from taproots, the pubescence simple to branched or stellate; leaves alternate, pe- tiolate, usually more or less cordate basally, commonly lobed; flowers in axillary clusters (sometimes solitary) or in subterminal pan- icles; involucel of 3 narrow to broad per- sistent bractlets; calyx 5-cleft; fruit a schiz- ocarp, the carpels mostly 10-15. 1. Petals commonly 1.5-2 cm long; bractlets of involucel ovate to oblong . M. sylvestris Petals asually less than 1 cm long; bractlets of involucel linear to narrowly lanceolate 2 2(1). Stems prostrate spreading from the caudex; leaves obscurely lobed; plant a common weedy species M. neglecta Stems erect; leaves definitely lobed; plant cultivated, rarely escaping M. verticillata Malva neglecta Wallr. Cheeses; Mallow. Annual or biennial, the stems prostrate- spreading, commonly 1-6 dm long, stellate- hairy; leaf blades reniform-orbicular, 0.6-3 cm long (from sinus to apex) or more, and much broader, crenate and not at all to only shallowly 5- to 7-lobed, the petioles to 20 cm long or more; flowers clustered (or solitary) in the axils; bractlets linear; calyx (3) 4-6 mm long at anthesis, the lobes acuminate; petals white to pink or lilac, about twice as long as the sepals; carpels hairy, roimded on the back. Weeds of disturbed sites and cultivated land, in much of Utah (specimens known from Cache, Iron, Kane, Salt Lake, San Juan, Summit, Utah, and Washington counties); widespread in North America; adventive from Eurasia; 22(ii). Note: Two other species, M. parviflora L. and M. rotundifolia L., might be present in Utah. They are similar to M. neglecta but have petals subequal to the sepals. Malva parviflora has glabrous petal claws, whereas in M. rotundifolia the claws are bearded. Malva sylvestris L. High Mallow. Biennial, the stems ascending, mostly 3-10 dm tall, rough hairy to glabrate; leaf blades 3-8 cm long or more and often broader, orbicular to cordate or reniform, crenate and with 5-7 lobes, the petioles to 10 cm long or more; flowers clustered in the leaf axils; bractlets ovate to elliptic; calyx 5-7 mm long at an- thesis, the lobes short and broad; petals 15-20 mm long, rose purple; carpels glabrous or nearly so, sharp edged. Cultivated orna- mental, rarely escaping (Utah Co., Larsen 7152 BRY); widespread in North America; adventive from Europe; 1(0). Malva verticillata L. Curled Mallow. An- nual, the stems erect, mostly to 10 dm tall or more, sparingly stellate-hairy; leaf blades mostly 1.5-7 cm long and as broad or broad- er, orbicular to reniform, undulate-crisped and distinctly 5- to 7-lobed, long-petioled; flowers solitary or clustered, subsessile or some pediceled; bractlets linear to narrowly lanceolate; calyx 3.5-5 mm long, the lobes acuminate; petals white, only somewhat sur- passing the sepals; carpels glabrous, the edges rounded. Cultivated ornamental, rarely es- caping (Washington Co., Galway in 1934 BRY); widely scattered in the United States; adventive from the Old World; 1(0). Our ma- terial belongs to var. crispa L. Malvastrum Gray Plants herbaceous, annual, stellate-hairy; leaves alternate, petiolate, the blades sub- cordate to tmncate basally, palmately lobed; flowers solitary in the axils or in terminal bracted clusters; involucel of usually 3 slen- March 1980 Welsh: Utah Flora, Malvaceae 31 der bractlets; calyx 5-cleft, the lobes long- acuminate; carpels 10-15; fruit a schizocarp. Malvastrum exile Gray. {Malveopsis exile (Gray Kuntze; Eremalche exile (Gray) Greene; Sphaeralcea exile (Gray) Jepson). An- nual, the stems spreading-decumbent to pro- strate, branching from near the base, 0.3-4 dni long, rather sparingly stellate-hairy; leaf blades suborbicular, 0.8-3.2 cm wide, palma- tely 3- to 5-cleft, with rounded or cuspidate teeth; petioles 1-5 cm long; bractlets narrow- ly lanceolate to sublinear; calyx 3-5 mm long; petals whitish to pinkish or lavender, only somewhat surpassing the sepals; carpels transversely wrinkled. Open sites in black- brush and creosote brush communities, 850-1200 m elevation, in Garfield (report probably erroneous) and Washington coun- ties; Arizona and southern California; 6(0). SiDA L. Plants herbaceous, perennial, from spread- ing rhizomes, densely stellate-canescent; leaves alternate, petiolate, crenate-serrate, not or obscurely linear, deciduous bractlets; calyx 5-lobed; carpels 5-10, 1 -seeded; fruit a schizocarp. Sida hederacea (Dougl.) Torr. Alkali-Mal- low. {Malva hederacea Dougl.; A4. californica Presl.; Disella hederacea (Dougl.) Greene). Perennial, the stems from elongate rhizomes, decumbent to prostrate, the surface obscured by overlapping stellate hairs, 1-4 din long; leaf blades reniform to orbicular, often oblique, dentate, obscurely if at all lobed, the petioles 0.3-2.5 (3) cm long; bractlets sub- linear; calyx 5-7 mm long; petals yellowish (fading orange), 10-12 mm long; carpels reti- culate on the sides. Saline meadows and seeps, at lower elevations in Emery, Salt Lake, Tooele, Uintah, and Utah counties (and probably elsewhere); Washington south to California, Texas, and Mexico; 6(i). SiDALCEA Gray Plants herbaceous, perennial, from tap- roots or short rhizomes, usually stellate and somewhat hirsute; leaves alternate, petiolate often dimorphic, the lowermost merely pal- mately lobed, the upper ones commonly cleft and with linear lobes; flowers borne in semi- spicate racemes, of two types, those of plants with perfect flowers the largest; involucel lacking; calyx 5-cleft; carpels 5-10, 1-seeded, tardily separating. Hitchcock, C. L. 1957. A study of the per- ennial species of Sidalcea. Univ. Wash. Publ. Biol. 18: 1-79. Roush, E. M. F. 1931. A monograph of the genus Sidalcea. Ann. Mo. Bot. Card. 18: 117-244. 1. Petals white or merely pinkish-tinged, often drying yellow; anthers bluish pink; plants rhizomatous; stems hirsute below S. Candida Petals pink to lavender; anthers usually yellow to white; plants rhizomatous or not; stems hirsute to glabrous or tomentose below 2 2(1). Plants from rather fleshy taproots; stems commonly hirsute below; calyx hirsute with pustulose hairs (at least in part) S. neomexicana - Plants often rhizomatous; stems stellate to glabrous below; calyx seldom with pustulose hairs S. oregana Sidalcea Candida Gray. Plants from slen- der rhizomes, the stems 4-10 dm tall, glabrous to hirsute with simple hairs below, more or less stellate above; leaf blades 6-20 cm wide, the basal ones shallowly 5- to 7- lobed and coarsely crenate, the upper ones divided into 3-5 entire segments; calyx 7-10 mm long, variously stellate-hairy and glandu- lar puberulent; petals white to pinkish, often drying yellow, 12-20 mm long; carpels about 3 mm long. Stream banks, lake shores, and seeps, 1410-2750 m, in Beaver, Garfield, Grand, Iron, Millard, Piute, Salt Lake, San Juan, Sevier, Summit, Uintah, Utah, and Wasatch counties; Wyoming and Colorado west to Nevada and south to New Mexico. Our materials have been treated as belonging to two more or less and at least partially sym- patric varieties; 25(vi). 32 1. Great Basin Naturalist Vol. 40, No. 1 Calyx rather uniformly hairy from base to apex of the lobes; plants of wide distribution S. Candida var. Candida Calyx more hairy at the base than on the lobes, the lobes often subglabrous; plants mostly from mountainous portions of middle Utah S. Candida var. glahrata Var. Candida. (S. Candida var. tincta Cock- erell). Known from Beaver, Grand, Iron, Salt Lake, San Juan, Summit, and Wasatch coun- ties; Colorado, New Mexico. Var. glahrata C. L. Hitchc. Known from Iron, Millard, Piute, Salt Lake, Sevier, Sum- mit, and Uintah counties; Wyoming, Colo- rado, and Nevada. Sidalcea neomexicana Gray. Plants from enlarged taproots or fascicled roots, the stems 2-9 (10) dm tall, hirsute below (or rarely glabrous) with simple or bifurcate hairs; leaf blades L5-11 cm wide, the basal ones cre- nate to shallowly 5- to 7-lobed, the cauline ones divided usually into 5 laciniate to entire segments; calyx 5-10 mm long, usually with some simple pustulose hairs interspersed with stellate ones; petals rose pink (fading blue- purple), 11-19 mm long; carpels 2-3 mm long. Wet Meadows, stream banks, and seeps, at 1370 to 2150 m in Box Elder, Garfield, Juab, Piute, Salt Lake, Sanpete, Sevier, Sum- mit, Utah, and Wasatch counties; Oregon, Idaho, and Wyoming south to California, Arizona, and Mexico. Hairs of lower stem nearly all simple; calyx coarsely and rather densely hirsute to coarsely hairy, lacking appressed stellate hairs; upper stems usually glabrous S. neomexicana var. neomexicana Hairs of lower stem often forked; calyx often with fine appressed stellate hairs in addition to the coarse ones; upper stems often stellate hairy S. neomexicana var. crenulata Var. crenulata (A. Nels.) C. L. Hitchc. (S. crenulata A. Nels., type from Juab, Utah; S. neomexicana ssp. crenulata (A. Nels.) C. L. Hitchc). Known from Box Elder, Juab, Salt Lake, Sanpete, Sevier, Summit, Utah, and Wasatch counties; Oregon, Idaho, and Ne- vada; lO(ii). Var. neomexicana. Known from Box Elder, Garfield, Piute, San Juan, Sevier, Utah, and Wasatch counties; Wyoming, Colorado, Ari- zona, and Mew Mexico; Mexico; 12(ii). Sidalcea oregana (Nutt.) Gray. {Sida ore- gana Nutt. ex T. & G.; S. nervata A. Nels.). Plants from a taproot, lacking or rarely with rhizomes, the stems 3-11 dm tall or more, glabrous or usually appressed-stellate hairy below, appressed-stellate above; leaf blades 2.5-17 cm wide, the basal ones shallowly 5-to 7-lobed and coarsely crenate, the cauline ones deeply lobed, with 3-7 coarsely toothed to entire lobes; calyx 3.5-9 mm long, various- ly stellate-hairy and sometimes bristly; petals 7-23 mm long, pale pink to rose pink (fading blue purple); carpels 2.5-3 mm long. Mead- ows, stream banks, and open woods, at 1680 to 2750 m in Cache, Juab, Salt Lake, San- pete, Summit, Utah, Wasatch, and Weber counties; Washington and Idaho south to California, Nevada, and Utah. Our materials belong to var. oregana; 32 (ii). Sphaeralcea St. Hil. Plants herbaceous, perennial, from tap- roots or rhizomes, glabrescent to canescent with stellate hairs; leaves alternate, petiolate, sometimes dimorphic, the lowermost merely toothed or palmately lobed (rarely entire), the upper ones cleft to entire; flowers borne in racemose to thyrsoid cymes; involucel of 3 or fewer filiform bractlets; calyx 5-cleft; car- pels 8-20, the seeds 1 or 2 per carpel; fruit a schizocarp, the mature fruit segments divided into a basal indehiscent, reticulate portion and an apical dehiscent portion. March 1980 Welsh: Utah Flora, Malvaceae 33 Jefferies, J. A. M. 1972. A revision of the Kearney, T. H. 1935. North American species genus Sphaeralcea (Malvaceae) for the of Sphaemlcea, Subgenus Eusphaeralcea. state of Utah. UnpubHshed thesis. Brigham Univ. Calif. Fubl. Bot. 19(1): 1-102. Young University. 92 pp. 1. Inflorescence racemose, rarely with more than one flower per node or, if more, as in S. caespitosa, then the plants restricted to Millard County 2 — Inflorescence thyrsoid to thyrsoid-glomerate, with usually more than one flower per node; distribution various 5 2(1). Leaf blades only slightly, if at all, 3- to 5-lobed, the margins irregularly cre- nate-dentate; hairs with rays radiating in more than a single plane; plants sel- dom more than 1.5 dm tall, known only from western Beaver and Millard counties S. caespitosa — Leaf blades distinctly 3- to 5-lobed, -parted, or -divided; hairs of rays radiating in a single plane (except in S. coccinea); plants often 1.5 dm tall or more, of different distribution 3 3(2). Leaves trifoliolate, the leaflets linear to narrowly oblanceolate and entire, or the upper ones simple and entire; plants of southeastern Utah S. leptophylla — Leaves various, but if trifoliolate then the leaflets oblanceolate and entire to toothed, if the uppermost simple then toothed or lobed; distribution various 4 4(3). Lowermost leaves simple and entire or trifoliolate, or some broadlv toothed or lobed; involucel present; rays of hairs radiating in one plane; plants of eastern Wayne County S. psoraloides — Lowermost leaves usually 3- to 5-lobed, the lobes usually toothed or again lobed; involucel present or lacking (caducous); rays of hairs radiating in several planes; plants of broad distribution S. coccinea 5(1). Plants only sparingly pubescent, the herbage bright green 6 — Plants moderately to densely pubescent, the herbage yellowish, whitish, or grayish 8 6(4). Leaves 3- to 5-parted or -divided, the lobes with narrow, regularly pinnatifid margins, the teeth at nearly right angles to the vein; carpels often with transparent lacunae, 4-6 mm high; plants rare, of southern Utah only S. rusbiji — Leaves variously lobed, divided, or parted, the lobes with broader margins ir- regularly toothed or lobed, but not as above; carpels with opaque lacunae, 3-4.5 mm high 7 7(6). Leaves slightly lobed, the margins unevenly toothed or, in .some, deeply parted to divided with the margin coarsely and irregularly lobed, the base subcordate to cuneate; plants of northern Utah S. munroana — Leaves 3- to 5-parted or -divided, the margins regularly cleft, lobed, or toothed, the base subcordate to deeply cordate; plants mostly of southern Utah S. grossulariifolia 8(5). Inflorescence loosely thyrsoid (appearing paniculate), leafy; flowers not numer- ous at each node; peduncles generally elongate; calyx surpassing the fruit; car- pels with reticulae extending onto back of carpel; plants of southwestern Utah . S. ambigua — Inflorescence contracted thyrsoid-glomerate; flowers often numerous at each node, not especially leafy; calyx often shorter than the fruit; carpels with reticulae confined to lateral face of carpel; plants of various distribution 9 34 Great Basin Naturalist Vol. 40, No. 1 9(8). Leaves 3- to 5-cleft, -parted, or -divided; carpels with well-defined reticiilae on less than half of carpel face; plants of all but the northeastern one-fourth of \JlJ^y^ S. grossiilariifolia Leaves shallowly 3- to 5-lobed; carpels with well-defined to nearly obscure re- ticulae on the lower one-third of the carpel; plants mainly of eastern and southern Utah, scattered elsewhere S. parvifolia Sphaeralcea ambigua Gray. Stems arising from a woodv caudex, several to numerous, .3-10 dm tall, whitish to yellowish canescent; leaf blades 1-6 cm long (from sinus to apex), 0.8-5 cm wide, thickish, usually rugose, with veins prominent beneath, ovoid, deltoid, or nearlv orbicular, the base cordate to deeply cordate, obscurely to definitely 3- to 5-lobed, the lobes crenate; inflorescence an open pan- icle, sometimes narrowly thyrsoid; pedicels usually shorter than the calyx; calyx uni- formly pubescent to glabrate, 6-20 mm long at anthesis, the lobes lanceolate to acumi- nate; petals 15-22 mm long, orange to or- ange pink (fading pinkish); carpels 12-16 mm high, the indehiscent portion comprising about one-third of the carpel, prominently reticulate. Creosote bush-blackbrush and mixed warm desert shrub communities, 670-1070 m, in Washington Co.; Nevada, Arizona, and California; and Mexico. Our material belongs to var. ambigua; 10(i). Sphaeralcea caespitosa M. E. Jones. Jones Globemallow. Stems solitary or more com- monly few to several from the summit of a branching woody caudex, 0.2-2.5 dm tall, whitish to grayish canescent; leaf blades 1.2-5.5 cm long, 1.2-6 cm wide, thickish, not nigose, veins apparent but not especially prominent, ovate to deltoid or orbicular, the base tnmcate to obtuse, obscurely if at all lobed, crenate to crenate-dentate; in- florescence thyrsoid, the flowers tightly clus- tered or solitary; pedicels shorter than the ca- lyx; calyx uniformly stellate, the rays of hairs not radiating in a single plane, the lobes lance-acuminate; petals 15-21 mm long, or- ange; carpels 12-14, 4-6 mm high, the in- dehiscent portion forming slightly more than one-third of the carpel, reticulate on the sides. Mixed desert shrub communities (shad- scale, rabbitbmsh, winterfat), mainly on Sevy Dolomite formation, at 1370-1750 m, in Mil- lard and Beaver counties; endemic; 20(iii). Sphaeralcea coccinea (Nutt.) Rydb. Com- mon Globemallow. {Malva coccinea Nutt.; Cristaria coccinea (Nutt.) Pursh; Sida coc- cinea (Nutt.) DC; Malvastrum coccineum (Nutt.) Gray.; Sida dissecta Nutt.; M. c. var. dissectum (Nutt.) Gray; M. dissectum (Nutt.) Cockerell; S. dissecta (Nutt.) Rydb.; S. coc- cinea ssp. dissecta (Nutt.) Kearney; S. coc- cinea var. dissecta (Nutt.) Kearney; M. c. var. elatum Baker; M. elatiim (Baker) A. Nels.; S. elata (Baker) Rydb.; S. c. ssp. elata (Baker) Kearney; S. c. var. elata (Baker) Kearney; M. cockerellii A. Nels.; M. micranthiim W. & S.). Stems solitary or few to many from the apex to a woody caudex, or less commonly from creeping rhizomes, 0.6-4.2 dm tall, white to yellowish canescent; leaf blades 1.1-3.7 cm long, 1.2-5.2 cm wide, usually wider than long, ovate to cordate-ovate in outline, the base often cordate, usually 3-to 5-lobed, with main divisions cleft almost or quite to the base, the lobes usually again toothed or lobed; inflorescence racemose, sometimes paniculate, rarely thyrsoid; pedicels shorter than the calyx; calyx uniformly stellate, the rays or hairs not radiating in a single plane, the lobes lance-acuminate; petals 8-15 mm long, orange; carpels 8-14, 2-3 mm high, the indehiscent part forming two-thirds or more of the carpel, reticulate on the sides and on the back. Blackbrush, shadscale-greasewood, sagebRish, juniper-pinyon, mountain brush, and ponderosa pine communities, 920-2750 m, in all counties (except Morgan and Wasatch?); Saskatchewan and Alberta south to Arizona, New Mexico, and Texas. Our ma- terials have been recognized as belonging to vars. dissecta and elata, but the segregation of these entities appears to have been wholly arbitrary, with intermediates more numerous than the supposed taxa; 152(xviii). Sphaeralcea grossulariifolia (H. & A.) Rydb. Gooseberry-Leaved Globemallow. {Sicki grossulariifolia H. & A.; Malvastrum March 1980 Welsh: Utah Flora, Malvaceae 35 grossulariifoUwn (H. & A.) Gray; S. pedata Torr., in Gray; S. g. ssp. pedata (Torr.) Kearney; S. g. var. pedata (Torr.) Kearney). Stems few to many from a woody caiidex, 1-10 dm tall or more, whitish to yellowish canescent to subglabrous and green; leaf blades 1.3-5 cm long, 1.3-5 cm wide, usually longer than wide, ovate to cordate-ovate in outline, the base cordate to truncate or ob- tuse, usually 3- to 5-lobed, the main division asually cleft or parted to irregularly toothed; inflorescence thyrsoid, with usually more tlian one flower per node; pedicels shorter than to much longer than the calyx; calyx luiiformly stellate, the rays of hairs not radi- ating in a single plane, the lobes ovate to lance- acuminate; petals 8-20 mm long, or- ange or rarely rose pink; carpels 10-14, 2.5-4.5 mm high, the indehiscent portion forming from two-fifths to three-fifths of the carpel, reticulate on the sides. Blackbrush, shadscale, rabbitbrush, sagebrush, juniper- pinyon, and less commonly mountain brush communities, 920-2450 m, in Beaver, Box El- der, Cache, Emery, Garfield, Grand, Iron, Kane, Juab, Millard, Morgan, Piute, Salt Lake, San Juan, Sanpete, Sevier, Tooele, Utah, Wasatch, Washington, and Wayne counties; Washington, Oregon, Nevada, Cali- fornia, and Arizona. Two infraspecific taxa have been segregated, largely on the basis of form of the leaf blades. Intergradation of the phases seems to be complete. Further, S. grossulariifolia appears to form intermediates with S. coccinea, S. parvifolia, and the more northern S. munroana. A phase with green herbage and thin leaves occurs along Glen Canyon. It seems to represent a taxonomic imit worthy of recognition. 1. Herbage bright green; leaves thin-textured; plants of eastern Kane and Garfield, and western San Juan counties S. grossulariifolia var. moorei Herbage gray green to whitish canescent; leaves thick-textured; plants widespread S. grossulariifolia var. grossulariifolia Var. grossulariifolia. This is the common and widely distributed phase of the species in Utah. The report by Kearney (I.e., p. 90) of S. digitata (Greene) Rydb. apparently belongs here; 115(xii). Var. moorei Welsh var. nov. Plantae siniilis var. grossulariifoliae sed differt in folii et caules virides et folii tenues. Holotype: Kane County, Utah, east side of Last Chance Bay, Lake Powell, Entrada Sandstone, S. L. Welsh and N. D. Atwood 11597, 2 May 1972 (BRY). Additional specimens: Kane County, mouth of Escalante River, Lake Powell, S. L. Welsh and G. Moore 11810, 5 June 1972; do. Wil- low Tank, D. A. White 111, 4 May 1962; do, Escalante Canyon, S. L. Welsh and G. Moore 11827, 5 June 1972; do, N. D. Atwood and R. Allen 3211, 24 August 1971; do, Hole-in-the- Rock, B. F. Harrison 12112, 14 May 1953; San Juan Co., 1 mi. E of Hole-in-the-Rock, S. L. Welsh and C. A. Toft 11869, 16 June 1972; do. Three Garden, Lake Powell, ca 1 mi. N of confluence with San Juan Arm, S. L. Welsh 12420, 5 May 1974; do. Comb Wash, S. L. Welsh and N.'D. Atwood 9972, 6 June 1970 (all at BRY). This variety is named to honor Glen Moore, botanist, teacher, collabo- rator, and collector. Sphaeralcea leptophylla (Gray) Rydb, {Malvastrum leptophyllum Gray). Stems few to many from a woody caudes, 2.0-5.5 dm tall, grayish canescent to yellow green throughout; leaf blades 1.0-3.2 cm long, di- gitately 3-lobed, the lobes entire, linear to oblanceolate, 1-4 mm wide, or the upper leaves simple and linear; inflorescence race- mose, elongate, usually with one flower per node; pedicels from much shorter to longer than the calyx; calyx uniformly stellate, the rays of hairs radiating in a single plane, the lobes lance-attenuate; petals 8-12 mm long, orange; carpels 7-9, 3-3.5 mm high, the in- dehiscent portion forming two-thirds-three- fourths of the carpel, coarsely reticulate, ridged, or tuberculate on the back. Black- brush and mixed semidesert shrub commu- nities, 1200-1520 m, in Garfield, Grand, and San Juan counties; New Mexico, Arizona, Texas, and Mexico. Two distinctive phases are recognizable among our materials; they can be distinguished as follows: 36 1. Great Basin Naturalist Vol. 40, No. 1 Plants grayish canescent, the hairs obscuring the surface of stems, leaves, and calvx lobes; leaf lobes narrowly oblong to linear; distribution as for the species S. leptophylla var. leptophylla Plants green to yellow green, the hairs widely separated, not obscuring stems, leaves or calyx lobes; leaf lobes, at least of lowermost leaves, oblanceolate to spatulate- known only from San Juan County, Utah S. leptophyUa var. janeae Var. leptophylla. This is the common form of the species. It is known from Garfield, Grand, and San Juan counties, Utah, and from New Mexico and Arizona; 7(iii). Var. janeae Welsh var. nov. Plantae similis var. leptophylhi sed differt in folii caules et calvces virides et lobos foliorum inferiorum oblanceolatos vel spathulatos. Holotype: San Juan County, Utah, along White Rim road, north of Turks Head, on sandy slopes in blackbnish communitv, Canyonlands Nation- al Park, S. L. Welsh 7064, 17 May 1968 (BRY). This variety is named to honor Jane .\rdis Murray Jefferies, student of Sphae- ralcea in Utah. Sphaeralcea miinroana (Dougl.) Spach in Gray. Munroe Cdobemallow. {Malta miin- roana Dougl. in Lindl.; Nuttallia miinroana (Dougl.) Nutt.; Malvastnirn munrooniim (Dougl.) Gray; S. suhrhomhoidea Rydb.; S. m. ssp. suhrhomhoidea (Rydb.) Kearney; S. m. var. s. (Rydb.) Kearney). Stems several to many from a branching woody caudex, 1.8-7 dm tall or more, yellowish green to some- what grayish canescent, the foliage usually bright green; leaf blades 1-6 cm long, 0.8-6 cm wide, ovate to orbicular or rhombic in outline, the base truncate to obtuse or sub- cuneate, usually 3- to 5-lobed, the sinuses shallow to very deep, the main divisions merely toothed or the lateral ones incised; in- florescence narrowly thyrsoid, usually with more than one flower per node; pedicels usu- ally much shorter than the calyx; calyx uni- formly stellate, the rays of hairs not radiating in a .single plane, the lobes deltoid-ovate to ovate; petals 8-15 mm long, orange; carpels 10-1.3, 2.5-3 mm high, the indehiscent por- tion forming about half the carpel, reticulate on the sides. Mixed desert shrub, or more commonly, in sagebrush and mountain brush conununities, 1370-2450 m, in Box Elder, Cache, Duchesne, Emery, Summit, Tooele, Uintah, Utah, and Wasatch counties; Mon- tana, Idaho, Washington, Wyoming, Nevada, and California. This entity is much like both S. parvifoUa and S. grossulariifolia. The green color of herbage is diagnostic of S. miinroana from both, except for the var. moorei which is not sympatric with S. miinroana; 21(ii). Sphaeralcea parvifoUa A. Nels. Nelson Globemallow. (S. marginata York, ex Rydb.; S. arizonica Heller, ex Rydb.). Stems few to many from a branching woody caudex, 1.5-10 (11) dm tall, grayish canescent, the fo- liage gray green or only somewhat yellow green; leaf blades 1.0-5.5 cm long, 1.2-5.2 cm wide, ovate to orbicular, reniform, or cordate-ovate, the base cordate to truncate or obtuse, usually shallowly 3- to 5-lobed, the si- nuses usually shallow, the lobes crenate-den- tate; inflorescence commonly narrowly thyr- soid, usually with more than one flower per node; pedicels usually shorter than the calyx; calyx uniformly stellate, the rays of hairs not radiating in a single plane, the lobes lance- ovate to deltoid; petals 7-15 mm long, or- ange; carpels 10-12, 3-4 mm high, the in- dehiscent part forming from one-fourth to one-third of the carpel, faintly reticulate on the sides. Blackbrush, salt desert shrub, sage- liRish, pinyon-jiuiiper, and mountain brush communities, at 850 to 2700 m, in Box Elder, Cache and Tooele counties, where probably of recent introduction, and in Duchesne, Emery, Garfield, Grand, Iron, Kane, Piute, San Juan, Sevier, Tooele, Washington, and Wayne counties, where likely indigenous; Nevada, Arizona, New Mexico, and Califor- nia. Sphaeralcea parvifoUa has been com- pared by Kearney (I.e.) with S. amhigua, which it resembles. The relationship of S. parvifoUa in Utah seems to lie with the largely sympatric S. gro.s.sularii folia; 144(xxii). Sphaeralcea psoraloides Welsh sp. nov. Stems few to many from a branching caudex, 1.4-2.4 dm tall or more, sparsely yellowish canescent, the foliage yellow green; leaf blades 1.3-3.5 cm long, 0.4-3.8 cm wide, ob- lanceolate to cuneate-ovate in outline,, cu- March 1980 Welsh: Utah Flora, Malvaceae 37 neate to obtuse or rounded basally, trifoliol- ate or simple to 3-lobed below, deeply 3- to 5-cleft above, the lobes entire to few toothed or lobed, usually more than 5 mm wide; in- florescence racemose, the flowers solitary in tlie upper axils; calvx uniformlv stellate, the rays of hairs radiating in a single plane, the lobes lance-acuminate; petals 10 (8-12) mm long, orange; carpels 10 (fruit unknown). Ephedra-Gmijia commimity on Entrada silts- tone, 1500 m, in Wavne Countv; endemic. Plantae similis S. leptophylla sed differt in foliolos oblanceolata vel laminas super iores confluentes et lobatos; e S. coccinea laminis inferioribus simplicibus vel trifoliolati digi- tatis distinguenda. Caules pauci vel multi e caudicibus raniifi- cantibus 1.4-2.4 dm alti vel plures flavidi-ca- nescentes parce folia et caules luteo-virides; laminae foliorum 1.3-3.5 cm longae 0.4-3.8 . cm latae oblanceolata ad cuneati-ovatas cu- neatae ad obtusas vel rotundatas basaliter tri- foliolatae vel simplicia ad trilobata infra 3-5 fissa profimde supra lobis intergris ad pauci- dentatis vel pauci-lobatis plerumque plus quam 5 mm latis; inflorescentiae racemosae, flores solitari in axilas supras; calyces stellati uniformiter, radius pilos radiantibus in plan- item singularem, lobus calycis lanci-acumi- natis; petala 10 (8-12) mm longa, aurantiaca; carpeli 10 (fructus ignotus). Holotvpe: Wavne County, Utah, Salt Wash, ca 17 mi. due WMW of Hanksville, T27S, R8E, Sec. 24, at 1500 m, on Entrada siltstone, Gmi/ia- Ephedra connnunitv, S. L. Welsh 13348, 1 June 1976 (BRY). Paratvpe: do, S. L. Welsh 13345, 1 June 1976 (BRY). Sphaeralcea riisbyi Gray. Stems few to many from a caudex, or rarely sub- rhizomatous, mostly 2-6.5 (8.5) dm tall, yel- lowish green to somewhat grayish canescent; leaf blades 1.3-3 cm long, 1.2-4 cm wide, ovate to orbicular in outline, the base trun- cate-obtuse to prominently cordate, parted to divided or merely cleft, the lobes again toothed (the teeth spreading at nearly right angles); inflorescence thrysoid to paniculate, with more than one flower per node; pedicels usually shorter (to much longer) than the ca- lyx; bractlets often dark red; calyx imiformly stellate (more densely so than on the her- bage), the rays of hairs not radiating in a single plane, the lobes ovate to lance-ovate; petals 9-18 mm long, orange; carpels 10-12, 4-6 mm high, the indehiscent part forming from one-fourth to two-fifths of the carpel, finely reticulate on the sides. Blackbrush, creosote brush, and mixed warm desert shrub communities, 820-1070 m, in Washington County; Arizona. S. rusbiji forms apparent intermediates with phases of S. grossularii- folia and S. pa wi folia; 4(0). UTAH FLORA: MISCELLANEOUS FAMILIES / Stanley L. Welsh' Abstract.— Considered in this treatment are the families Aquifoliaceae, Canabinaceae, Ericaceae, Krameriaceae, Magnoliaceae, Moraceae, Oleaceae, Pyrolaceae, Resedaceae, Tamaricaceae, and Tiliaceae. These 11 families include 61 cultivated, escaped, and indigenous species. The flora of Utah is both large and diverse. A portion of the diversity is due to the pres- ence of a large number of cultivated species in many plant families. Floras of regions have traditionally avoided inclusion of strictly cul- tivated species. Only those taxa which escape and become acclimated have been treated. Included are the cultivated plants and those species which escape. The present treatment covers all taxa in common cultivation, and es- pecially those which are represented in re- gional herbaria. In Table 1 a list is presented of the families treated herein, the numbers of genera and species, and whether cultivated or indigenous. The list heavily favors the cultivated and/or escaped species, and, because of the status of cultivated species collections, the treatment is likely to be incomplete. It is presented herein for use by students of the flora who want to know the names of culti- vated and of native plant species. Aquifoliaceae Holly Family Evergreen Shrubs or small trees: leaves al- ternate, simple, coriaceous, armed with spiny teeth; stipules minute, caducous; flowers usu- ally imperfect, regular, small and inconspic- uous, solitary or few in axillary cymes; sepals usually 4, more or less connate basally; petals usually 4, distinct or slightly connate basally; stamens or staminodes usually 4 (-9), alter- nate with the petals; pistil 1, the ovary supe- rior, 3- to many-loculed, the carpels as many as the locules; fruit a globose, berrylike drupe with 2-8 bony 1 -seeded divisions. Ilex L. Evergreen; leaves thick and shining; flow- ers small, mostly in few-flowered axillary cymes; staminodia usually present in pistil- late flowers, a rudimentary pistil present in most staminate flowers; fruit usually brightly colored. (Note: Members of this family are known in Utah in cultivation only). Flowers in axillary clusters on branches of the previous year /. oquifolium Flowers in solitary cymes on branches of the current year /. opaca Ilex aquifolium L. English Holly. Tall shnibs to small trees of ornamental plantings, rare in Utah; introduced from the Old World; 1(0). Ilex opaca Art. American Holly. Low to moderate shrubs of ornamental plantings, oc- casional in Utah; introduced from the eastern United States; 1(0). Cannabinaceae Hemp Family Plants herbaceous, with watery juice; leaves alternate or opposite, palmately veined and lobed or divided to essentially compound; stipules persistent; flowers imper- 'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 38 March 1980 Welsh: Utah Flora, Miscellaneous Families 39 feet, the plants dioecious, regular, the stami- nate in open racemes or panicles, the pistil- late in dense clusters; sepals 5, connate in pistillate flowers and enclosing the ovary; stamens 5; pistil 1, 2-carpelled, the styles 2; fruit an achene. Plants strong-smelling, stout, erect herbs; leaves palmately 5- to 9-parted .. Cannabis Plants rough-stemmed clambering vines; leaves coarsely 3- to 7-lobed Humulus CaNxNabis L. Plants dioecious or rarely some mon- oecous; leaves palmately lobed to parted and apparently compound, alternate or the lower opposite; flowers small, inconspicuous, the staminate in leafy panicles in upper axils; sepals 5, oblong; stamens 5; pistillate flowers in small clusters on leafy branches from up- per axils, each flower subtended and enclosed by an acuminate bract, the calyx barelv lobed, surrounding only the base of the ovary; stigmas 2, elongate; fruit a lenticular achene, enclosed within the accrescent bract. Small, E. and A. Cronquist. 1976. A practical and natural taxonomv for Cannabis. Taxon 5: 405-435. Cannabis sativa L. Hemp, Marijuana, Hashish, Pot, Grass. Plants 6-20 dm tall or more, the stems simple or much branched; leaves long petioled, the blades 3- to 7-part- ed, the segments oblanceolate to elliptic, at- tenuate to accuminate apically, sharply ser- rate, mostly 4-12 cm long and 0.4-1.8 cm wide, scabrous and more or less glandular and pubescent; achenes mostly 3.5-4.5 mm long. Cultivated historically in Utah for fiber Table 1. Families, genera, and species treated. No. Genera No. Species Family Cultivated Indigenous .Aquifoliaceae 1 2 Canabinaceae 2 1 1 Ericaceae 6 11 Krameriaceae 1 2 Magnoliaceae 2 4 Moraceae 3 4 Oleaceae 6 14 4 Pyrolaceae 3 7 Resedaceae 1 1 Tamaricaceae 1 3 Tiliaceae 1 7 TOTAL 27 36 25 produced from the stems, the commercial source of hemp; currently sporadic, or else grown illegally for its intoxicant properties. Utah materials are sufficiently rare as to give only hints as to the classification below the species level. It seems likely, however, that, at least historically, two phases have been grown in the state (for a complete review see Small and Cronquist 1. c). Nineteenth-cen- tury plantings for hemp likely belonged to ssp. sativa, demonstrated to have only limited intoxicant ability. At least some of the recent introductions clearly belong to ssp. indica (Lam.) Small & Cronq. (C. indica Lam.), which has demonstrated high intoxicant lev- els. Humulus L. Plants herbaceous, twining, perennial vines; stems scabrous; leaves opposite, broad- ly 3- to 5-lobed; flowers small, inconspicuous, the staminate in axillary panicles; sepals 5, distinct; stamens 5; pistillate flowers in short spikes, in pairs, with each pair subtended by a foliaceous bract; calyx membraneous, un- lobed, closely covering the ovary; stigmas 2, elongate; fruit an achene enclosed by the persistent calyx and accrescent bracts. Humulus americanus Nutt. American Hop. Plants twining, the stems to 20 dm long or longer; leaves ovate to orbicular in out- line, deeply cordate ba.sally, mostly 3-15 cm long and 2.8-16 cm wide, the lobes serrate to doubly so, attenuate to acimiinate apically, rough-hairy above, glandular-dotted beneath; fruiting spikes usually 2-3.5 (4) cm long at maturity. Twining over shrubs and other veg- etation at lower and middle elevations in Duchesne, (iarfield. Grand, Millard, Piute, Salt Lake, Summit, Uintah, Utah, Wasatch, Washington, and Weber counties, and prob- ablv throughout Utah; widespread in North America. The hop of commerce, H. hipulus 40 Great Basin Naturalist Vol. 40, No. 1 L., or European hop, is grown in the United States, where it has escaped and persists. Though not definitely known for Utah, the European hop might occur here. It can be distinguished by its unlobed leaves, or when lobed, the terminal lobe is less than twice longer than broad; 27(v). Ericaceae Heath Family Shrubs or subshnibs; leaves simple, alter- nate, sometimes leathery or persistent; flow- ers perfect, regular, axillary, in terminal clus- ters, or solitary; sepals mostly 4 or 5, distinct or more or less connate; petals mostly 4 or 5, connate or distinct, the corolla rotate to fun- nelform or urn shaped; stamens as many as the corolla lobes and alternate with them or twice as many, the anthers dehiscent by ter- minal pores or by longitudinal slits; pistils 1, the ovary superior or inferior, usually with 4-10 carpels and locules; styles 1, the stigma capitate or lobed; fruit a capsule or a berry. 1. Ovary inferior or apparently so 2 - Ovary superior 3 2(1). Plants prostrate shrublets, rooting along the stems; ovary superior but sur- rounded by the fleshy calyx when ripe and apparently inferior Gaultheria (hiimifusa) - Plants erect or ascending, rooting only at the base; ovary inferior Vciccinium 3(1). Flowers borne in terminal corymbs, white, the segments of the corolla much longer than the short tube; leaves punctate below with yellow-glandular dots ... Ledum - Flowers solitary and axillary or in axillary racemes, rarely terminal, pink to lavender, the segments of the corolla much shorter than the tube; leaves lacking glandular punctae 4 4(3). Corolla broadly saucer shaped or ratate, not constricted at the apex Kahnia Corolla campanulate to urn shaped, often more or less constricted at the throat 5 5(4). Corolla campanulate; anthers lacking appendages; fruit a capsule embedded in a fleshy calyx Gaultheria {humifusa) Corolla urn shaped; anthers 2-awned; fruit a berry Arctostaphylos Arctostaphylos Adams Evergreen prostrate to ascending or erect shnibs, often with purplish to orange brown, smooth bark; leaves alternate, simple, entire, leathery-thickened; flowers in terminal pan- icles or racemes, perfect, regular; sepals usu- ally 5; petals usually 5, united almost to the tips; corolla urn shaped; stamens usually 10, included; anthers opening by falsely terminal pores, each with 2 hornlike appendages; ovary superior, usually 5-loculed; fruit fleshy, berry like, 1- to several-seeded. Adams, J. E. 1940. A systematic study of the genus Arctostaphylos. J. Elisha Mitchell Soc. 56: 1-62. Eastwood, A. 1934. A revision of Arctosta- phylos with keys and descriptions. Leaft. West. Bot. 1: 105-127. Plants with creeping-prostrate stems; leaves obovate-spatulate, commonly less than 1.5 cm long X. uva-ursi Plants with stems ascending to erect; leaves mostlv ovate to lanceolate or elliptic, often more than 2 cm long 2 March 1980 Welsh: Utah Flora, Miscellaneous Families 41 2(1). Calvx and pedicels puherulent with spreading glandular hairs; twigs and leaves puberulent throughout with spreading hairs; plants of Washington County A. pringlei - Calyx glabrous or nearly so; twigs and leaves puberulent or sessile to sparingly stipitate-glandular or almost or quite glabrous; plants of various distribution 3 3(2). Twigs and axis of inflorescence white-puberulent, not glandular; plants of Washington and Kane counties A. piingens Twigs and axis of inflorescence glandular to glandular-puberulent; plants widely distributed A. patulu Arctostaphylos patula Greene. Green-leaf Manzanita. {Uva-iirsi patiila (Greene) Abranis; A. piingens var. phityphylla Gray; A. platyphyUa (Gray) Kuntze; A. obtiisifolia Piper; A. patula var. incarnata Jeps.; A. pi- nctorum Rollins; A. parryana var. pinetorum (Rollins) Weislander & Schreiber). Rounded shrubs with gnarled stems to 15 cm long or more, the bark smooth, cinnamon to reddish brown or purplish in color; branchlets glandular-puberulent and sometimes with long-spreading hairs as well; leaf blades (0.8) 1.8-4.7 cm long, (0.6) 1.5-4 cm wide, ovate to elliptic, lanceolate, or orbicular, obtvise to acute apically, rounded to truncate basally, glabrous or glandular, yellow green; petioles pubescent like the twigs; inflorescence pan- iculate, the axis and bracts glandular-pub- enilent and sometimes with some long hairs; pedicels glabrous; sepals glabrous; corolla pink to white, 5-8 mm long; ovary glabrous; fruit 8-11 mm thick, depressed-globose, glabrous, white to brown, with nutlets sepa- rable or not. Usually associated with ponde- rosa pine at 1520 to 2830 m in Beaver, Duch- esne, Garfield, Iron, Juab, Kane, Millard, San Juan, Sanpete, Sevier, Summit, Tooele, Uin- tah, Utah, Wasatch, and Washington coun- ties; Colorado, Nevada, Oregon, Arizona, and California. Arctostaphylos patula and A. platyphyUa both date as species from the same year, 1891; the question of which has priority is difficult to ascertain; 64(x). Arctostaphylos pringlei Parry. Pink- bracted Manzanita. Rounded, erect shrubs to 20 dm tall or more, the bark smooth, dull red l)rown; branchlets densely glandular-hairy with long-spreading hairs; leaf blades (1.2) 1.8-4.2 cm long, (0.4) 0.8-2 cm wide, elliptic to lance-elliptic or lanceolate, obtuse to acute apically, truncate to rounded or obtuse basally, glandular-pubescent, gray green; pe- tioles pubescent like the twigs; inflorescence paniculate or racemose, the axis and bracts glandular-hairy; corolla pink, 6.5-8.5 mm long; ovary glandular-hairy; ovary glandular- hairy; fruit 6-10 mm thick, ovoid, glandular- hairy, red, with nutlets inseparable. Oak-juni- per community, 1840-2750 m, in Washing- ton County; Arizona, California and Baja California; 4(o). Arctostaphylos pungens H.B.K. Mexican Manzanita. Erect or ascending, rounded shrubs to 20 dm tall or more, the bark smooth, red brown; branchlets canescent with a dense pubescence; leaf blades 1.6-4.7 (6) cm long, 0.5-3.2 cm wide, ovate to ellip- tic or oblong, rounded to acute apically, acute to rounded basally, puberulent on one or both sides, bright green; petioles pub- escent like the twigs; inflorescence pan- iculate, the axis and bracts canescent; pedi- cels glabrous; sepals glabrous; corolla pink to white, 5.5-8.5 mm long; ovary glabrous; fruit 5-8 mm thick, depressed-globose, glabrous, brownish red, with nutlets separable or not. Pinyon, juniper, live oak communities, 920-2750 m, in Washington and Kane (At- wood .3538 BRY) counties; California, Ari- zona, New Mexico, Texas; Mexico; 19(iii). Arctostaphylos uva-ursi (L.) Spreng. Kin- nikinnick, Bearberry, Sandberry. {Arbtitus uva-ursi L.; Uva-ursi procumhens Moench; Mairania uva-ursi (L.) Desv.; U. buxifolia S. F. Gray; A. officinalis Wimm. & Grab.; A. procumhens in Mey. & Elkan; U. uva-ursi (L.) Britt. in Britt. & Br.; A. media Greene; A. uva-ursi var. coactilis Fern. & Macbr.; A. uva-ursi var. adenotricha Fern. & Macbr.). Prostrate shrub with stoloniferous rooting steins, mat-forming, the branches ascending, the intemodes usually apparent, puberulent and sometimes glandular, the bark exfoliating exposing dull brown imder bark; leaf blades 42 Great Basin Naturalist Vol. 40, No. 1 (0.6) 1-2.7 (3) cm long, 0.3-1.2 wide, oblan- ceolate to spatulate, rounded apically, cu- neate to acute basally, glabrous or pub- enilent, especially on the margins, green; inflorescence racemose, the axis and bracts glandular; pedicels glabrous or sparingly puberulent; sepals glabrous; corolla pink to white, 4-5.2 mm long; ovary glabrous; fruit 6-11 mm thick, globose, bright red, with sep- arable nutlets. Ground layer in coniferous forests, at 2140-3350 m, in Daggett, Duch- esne, Garfield, Salt Lake, Sevier, Summit, Uintah, and Wasatch counties; Alaska and Yukon east to the Atlantic and south to Cali- fornia, New Mexico, Illinois, and Georgia; Eurasia; 15(ii). Gaultheria L. Prostrate shrubs, the branches rooting; leaves alternate, thin, serrulate; flowers ax- illary, solitary, perfect, regular; calyx 5- lobed, united, enlarging and becoming fleshy at maturity; corolla campanulate, the lobes shorter than the tube; stamens usually 10, in- cluded, the filaments flattened, tapering to the apex; anthers opening by terminal pores, not awned; ovary superior, usually 5-loculed; fniit a loculicidally dehiscent capsule en- closed by the fleshy expanded calyx. Gaultheria humifusa (Grab.) Rydb. Al- pine Wintergreen. {Vaccinium humifusum Grab.; G. myrsinites Hook.). Prostrate, scarcely woody plants with creeping, rooting stems to 2 dm long, glabrous or puberulent; leaves 0.6-1.5 cm long, 0.4-1.3 cm wide, oval to ovate or elliptic, rounded to obtuse api- cally and basally, serrulate; flowers solitary, axillary; calyx glabrous; corolla 3-4 mm long, campanulate, pink; fruit 5-7 mm thick, sub- globose, red. Ground layer in coniferous for- ests and margins, 2900-3350 m, in Duchesne and Summit counties, and possibly elsewhere; Colorado westward to California and north to Alberta and British Columbia; 5(i). Kalmia L. Low shrubs with puberulent branches; leaves opposite, evergreen, leathery, decur- rent, entire, revolute, glaucous beneath; flow- ers in terminal leafy-bracted corymbs or soli- tary, perfect, regular; calyx 5-lobed, the segments almost distinct; corolla bowl shaped, the lobes shorter than the tube, the tube with 10 pouches in which the anthers are enclosed in bud; stamens usually 10, the filaments flattened, hairy below; anthers opening throughout, unawned; ovary superi- or, 5-loculed; fruit a septicidally dehiscent capsule. Kalmia microphylla (Hook.) Heller. Bog Laurel. {K. glauca var. microphylla Hook.; K. polifolia var. microphylla (Hook.) Rhed.). Erect slender shrubs, 0.7-1.5 dm tall; leaves 0.6-1.8 (3) cm long, 0.2-0.8 (1.2) cm wide, lance-oblong to elliptic, revolute, shining and green above, grayish beneath; corymbs most- ly 2- to 6-flowered, the pedicels 1-3 cm long; sepals glabrous, ciliate; corollas 11-14 mm broad, pink; capsules 4-6 mm broad. Alpine meadows and lake margins, 2900-3800 m, in Daggett (?), Duchesne, Summit, and Uintah counties; Alaska and Yukon south to Califor- nia and Colorado; ll(i). Ledum L. Erect or spreading shrubs with glandular- pubenilent branchlets; leaves alternate, ever- green, leathery, entire, revolute, pale below; flowers in terminal corymbs, perfect, regular; calyx small, the segments almost distinct; co- rolla rotate, the 5 petals distinct or nearly so; stamens usually 5-10, the filaments almost filiform, usually hairy below; anthers opening by terminal pores, unawned; ovary superior, 5-loculed; fruit a septicidally 5-valved cap- sule, opening at the base. Note: At least some species of this genus are poisonous to live- stock. Ledum glandulosum Nutt. Trapper's Tea. Plants mostly 5-15 dm tall, the branchlets puberulent and glandular dotted; leaves 1.1-3.4 (4) cm long, 0.4-1.4 (1.8) cm wide, el- liptic to oblong, rounded to acute apically and basally, green above, pale to grayish be- neath, glandular, the margin more or less revolute; flowers white, the segments to 5 mm long or more; pedicels commonly 1-2.5 cm long, puberulent near the base; capsules 3-6 mm long, puberulent and glandular. Meadows, stream banks, and bogs in open forest, 2600-3050 m, in Duchesne^ Salt Lake, March 1980 Welsh: Utah Flora, Miscellaneous Families 43 Summit, and Uintah counties; British Cohim- bia east to Montana and south to Cahfornia, Nevada, and Wyoming; 9(o). Vaccinium L. Decumbent-ascending to erect shrubs; leaves alternate, deciduous, or more or less evergreen, entire or serrulate, flat, green or pale beneath; flowers solitary, axillary, or in terminal clusters, perfect, regular; calyx 4- to 6-lobed, united at the base; corolla urn shaped or campanulate, the 4-6 lobes shorter than the tube; stamens 8-12, the fllaments usually glabrous; anthers opening by pores at the ends of tubular beaks, usually 2-awned; ovary inferior, usually 4-locular; fruit a sever- al-seeded berry. Camp, W. H. 1942. A survey of the Ameri- can species of Vaccinum, subgenus Eu- vaccinium. Brittonia 4: 205-247. 1. Branches bright green and angled; plants often less than 3 dm tall 2 - Branches neither bright green nor angled, or sometimes irregularly angled when dry; plants often more than 3 dm tall 3 2(1). Fruit red; grooves of branches usually glabrous; leaves often less than 12 mm long V. scoparium - Fniit blue black or black; grooves of branches usually puberulent; leaves often over 12 mm long V. myrtillus 3(1). Flowers in clusters of 2-4, or solitary; leaves entire; calyx deeply lobed, the lobes persistent in fruit V. occidentale Flowers solitary in leaf axils; leaves more or less serrate; calyx shallowly lobed, the lobes deciduous in fruit 4 4(3). Plants mostly 1-3 dm tall; leaves serrate above the middle and uncon- spicuously below the middle, mainly 1-3 (4) cm long, oblanceolate to obovate . V. caespitosiwi Plants mostly 4-7 dm tall or more; leaves serrate to the base or nearly so, commonly 2-6 cm long, elliptic to ovate V. membranaceum Vaccinium caespitosum Miehx. Dwarf Huckleberry. Plants mostly 1-3 dm tall; twigs brownish, somewhat angled, pub- erulent or glabrous; leaves 0.7-4 cm long, 0.3-2 cm wide, oblanceolate to obovate, ob- tuse or less commonly acute to rounded api- cally, usually cuneate basally, serRilate from tip to below the middle; flowers solitary, ax- illary, whitish to pink, the corollas 5-6 mm long, twice as long as thick; calyx obscurely lobed, the lobes deciduous in fruit; berries blue glaucous, subglobose, 5-8 mm broad, edible and good. Streamsides, meadows, and rock outcrops, 2,227-3,416 m elevation, in the Uinta Mountains and Boulder Mountains, in Daggett, Duchesne, Garfield, Summit, and Uintah counties; Alaska and Yukon east to Newfoundland and New Hampshire, and south to California and Colorado. Materials from Utah have previously passed under the names V. membranaceum Dougl. (see below) and V. globulare Rydb. The latter is not known for the state; 10(i). Vaccinium membranaceum Dougl. Moun- tain Huckleberry. Shrubs mostly 3-7 dm tall or more; twigs brownish, glabrous or pub- CRilent; leaves 1.8-7 cm long, 1-3.4 cm wide, elliptic or less commonly ovate or obovate, acute to obtuse apically, acute to rounded basally, serrate almost throughout; flowers solitary, axillary, yellowish pink, the corollas about 6 mm long, about one-third longer than broad; calyx obscurely lobed, the lobes deciduous; berries purple, not glaucous, 7-9 mm broad, edible and good. Slopes in aspen- conifer and spruce-fir woods, 2,500 to 2,775 in, in Cache, Carbon, Duchesne, Salt Lake (?), and Summit counties; British Columbia southward to California, Idaho, and Mon- tana; 6(0). Vaccinium myrtillus L. Dwarf Billberry. (V. oreophilum Rydb., in part, the type from 44 Great Basin Naturalist Vol. 40, No. 1 the Uinta Mountains.) Plants mostly 0.5-3 dm tall; twigs seldom numerous and broomlike, green, sharply angled, puberulent; leaves 1.1-3.9 cm long, 0.6-1.6 cm wide, ovate to lanceolate or elliptic, acute to obtuse api- cally, obtuse to roimded basally, serrulate al- inos't or quite from base to apex; flowers soli- tary, axillary, pink, the corollas 4-5 mm long; calyx shallowly lobed; berry usually bluish, 5-8 mm broad. Ground layer in coniferous forests, 2750-3200 m, in the Uinta, Wasatch, and LaSal moimtains (Daggett, Duchesne, San Juan, Summit, and Uintah counties), where evidently not common; British Colum- bia and .\lberta south to Arizona and New Mexico; Eurasia. Vaccinium myrtillus is a near congener of the very common V. scopa- riwn and can be distinguished by the larger size of its leaves and flowers and by the pub- erulent stems; 6(i). Vaccinium occidentale Gray. Western Huckleberry. Plants mostly 2-6 dm tall, the twigs round, usually glabrous; leaves 0.6-2.1 cm long, 0.4-1.2 cm wide, oblanceolate, rounded to obtuse apically, acute basally, en- tire; flowers 2-4, or less commonly solitary in the axils, pinkish, the corollas 3.5-6 mm long; calvx definitely lobed, the lobes persistent in fniit; berries blue, glaucous, 4-6 mm thick. Meadows, streamsides, and forest margins, 2750-3100 m, in the Uinta Mountains in Daggett (?), Duchesne, Summit, Uintah, and Wasatch counties; British Columbia south to California and Idaho; 12(iv). Vaccinium scoparium Leiburg. Grouse- berry. (V. myrtillus var. microphijUum Hook.; V. microphijllwn (Hook.) Rydb., not Rein.; V. enjthrococcum Rydb.). Plants mostly 1-2.5 dm tall, the twigs numerous, broomlike, sharply angled, usually glabrous; leaves 0.6-1.3 cm long, 0.3-0.7 mm wide, ovate, ob- tuse to acute apically, roiuided to obtuse ba- sally, serrulate throughout; flowers solitary, axillary, pinkish, the corollas 2.5-3.5 mm long; calyx very shallowly lobed; berry bright red, drying red purple, 4-6 mm thick. Com- mon component of ground layer in con- iferous forests and forest margins, 2450-3200 m, in the Uinta Mountains in Daggett, Duch- esne, Summit, Uintah, and Wasatch counties; British Columbia and Alberta south to Cali- fornia and Colorado; 20(i). Krameriaceae Ratany Family Shrubs, with divaricate branches; herbage grayish pubescent; leaves alternate, simple, entire extipulate; flowers perfect, irregular, solitary, axillary; pedicels usually with 2 op- posite foliacious bracts; sepals 4 or 5, un- equal; petals 5, the upper 3 long clawed, dis- tinct or partially connate and often purplish in color, the 2 others broad, thick, sessile, usually greenish and glandlike; stamens 4, free or adnate to claw of upper petal, the an- thers dehiscent by pores; ovary superior, 1- loculed; ovules 2; fruit an indehiscent pod, armed with prickles. A family of the Western Hemisphere of a single genus with about 25 species from South America to southern United States. Krameria L. A single genus with characteristics of the family. Branchlets modified as thickened thorns 0.8-1.2 mm in diameter at base; spines of fruit barbed at apex only K. graiji Branchlets not modified as thorns or if so then less than 0.6 mm in diameter; spines of fruit with barbs scattered or, rarely, barbless K. pawiflora Krameria grayi Rose & Painter. White Ra- tany. Shrubs branched, 2.5-6 dm tall and as wide; leaves 5-7 (25) mm long, 1-3 mm wide, lance-ovate to lanceolate, elliptic or oblong, more or less spinulose-tipped, tomen- tose on both surfaces; pedicels not glandular- pubescent; upper petals 2.5-3.5 mm long, 0.3-0.5 mm wide, yellowish with a purplish tip; sepals 4.5-6.5 mm long, villous-pilose dorsally, pilose to glabrate within, purplish; prickles of the fruit 2-6 mm long at maturity, each with a whorl of barbs at the apex; pods March 1980 Welsh: Utah Flora, Miscellaneous Families 45 subglobose, 6-10 mm in diameter, hirsute over the surface and on bases of prickles. Blackhmsh and creosote bush communities at 670-1170 m in western Washington Co.; Cal- ifornia, Nevada, Arizona, New Mexico, Texas, and Mexico; 2(i). Krameria parvifolia Benth. Range Ratany. (K. glandulosa Rose and Painter; K. parvi- folia var. glandulosa (Rose and Painter) Macbr.; K. iniparata (Britton) Macbr.) Shrubs, intricately branched, 2-6 dm tall and as wide; leaves 3-15 mm long, 0.3-1 mm wide, linear to oblong, callous- to spinulose- tipped, tomentose on both surfaces; pedicels glandular or not; upper petals 2.5-2.8 mm long, 0.7-1.2 mm wide, yellowish; sepals 4-6 mm long, strigulose dorsally, glabrous within, pinkish to purplish; prickles of fruit 2-5 mm long, retrorsely barbed along the rachis; pods subglobose, 5-9 mm in diameter, pilose-hir- sute on the surface. Joshua tree, blackbrush, creosote bush, and bursage communities, 750-1600 m, in Washington Co.; California, Nevada, Arizona, New Mexico, Texas, and Mexico. The materials demonstrate variation in glandular condition of pedicels, sepals, and bracts. The variation seems to be haphazard, with little or no correlation with other fea- tures or with ecology. Hence, included herein as synonyms are those names involved with recognition of glandular and nonglandular phases; 16(i). Magnoliaceae Magnolia Family Deciduous or evergreen trees or shrubs; leaves alternate, simple, entire or lobed, stip- ulate, the stipules enclosing the buds, de- ciduous or caducous, and leaving a circular scar; flowers regular, perfect, solitary, termi- nal and axillary, large and showy, the floral parts spirally arranged; sepals often 3, the petals 6 to many; stamens numerous, sepa- rate, hypogynous, the anthers 2-loculed; pis- tils several to many, each 1-loculed and 1- carpelled; style 1, the stigma 1; fruit a follicle or samara. 1. Leaves lobed, truncate or broadly retuse at the apex; flowers borne after the leaves Liriodendron Leaves entire, acute, or acuminate; flowers borne before or after the leaves Magnolia Liriodendron L. Trees, the leaves large and 4-lobed; flowers large, inconspicuously colored; sepals 3, soon reflexed; petals 6, ascending to erect, forming a tuliplike corolla; anthers extrorse; pistils many, en masse becoming conelike, the indi- vidual samaras eventually deciduous. Liriodendron tulipfera L. Tulip Tree; Yel- low Poplar. Deciduous, cultivated trees to 40 m tall or more, the tmnks to 10 dm in diam- eter or more; leaves long-petioled, the blades 6-15 cm long and almost as wide; flowers solitary, terminal; sepals green; petals 3.7-6 cm long, yellow green, with a basal orange spot within; samaras narrow, 3-4 cm long. Occasional shade tree in more moderate low elevation portions of Utah; introduced from the eastern United States; 8(o). Magnolia L. Trees or shrubs; leaves large, entire; flow- ers large, conspicuous or inconspicuous; se- pals 3, colored like the petals; petals 6-12, erect or spreading; anthers introrse; pistils many, en masse becoming conelike, the indi- vidual follicles finally dehiscent. Plants shrubs or small trees, deciduous; flowers showy, cream to pink or suffused with rose or lavender, borne before the leaves appear M. soulangeana Plants moderate to large trees, deciduous or evergreen; flowers greenish and inconspicuous or, if showy, then white in color and the trees evergreen 2 46 Great Basin Naturalist Vol. 40, No. 1 Plants evergreen, the leaves dark green, leathery; flowers white M. grandiflora Plants deciduous, the leaves not both dark green and leathery; flowers greenish ygllovv ^- acuminata Magnolia acuminata L. Cucumber-tree. Deciduous trees to 30 m tall or more; leaves deciduous, short-petioled, the blades 8-25 (3) cm long and 4-15 cm wide; flowers solitary, terminal; perianth greenish yellow, 5-8 cm long. Cultivated shade tree, uncommon, hardy in the major cities of the state; in- troduced from the eastern United States; 2(o). Magnolia grandiflora L. Bull Bay. Ever- green trees to 30 m tall; leaves evergreen, short-petioled, the blades mostly 8-20 cm long and 3-8 cm wide; flowers solitary, ter- minal; perianth white, mostly 8-12 cm long. Cultivated ornamental, uncommon, not hardv except in favorable sites in moderate to warm portions in Utah; introduced from southeastern United States; l(o). Magnolia soulangeana Soul. Showy Mag- nolia. Shrubs or small trees to about 4 m tall; leaves deciduous, short-petioled, the blades mostly 8-14 cm long and 3.5-10 cm wide; flowers solitary, terminal; perianth cream to pink or suffused with rose or lavender, 6-12 cm long or more. Cultivated ornamental, oc- casional in more moderate climatic areas of Utah; a hybrid of M. denadata Descr. and M. lili flora Descr., both native of China; 3(o). MORACEAE Mulberry Family Deciduous trees or shrubs with milky juice; leaves alternate, simple, pinnately or pal- mately veined, entire, serrate, or lobed, stipu- late, the stipules small and distinct or each pair forming a cap over the bud and leaving a scar around the stem; flowers imperfect, minute, regular, borne in cymes or much modified inflorescences; perianth of usually 4 sepals; staminate flowers with usually 4 (2 in Ficus) stamens, the filaments distinct; pistil- late flowers with or without a 4-lobed per- ianth; pistil 1, the ovary superior to inferior, 1-loculed, the styles and stigmas 2 (1 in Ma- dura)., fruit a multiple (Morus, Madura) or a syconium (Ficus). 1. Fniit a fleshy hollow receptacle with flowers borne inside (syconium); leaves palmately veined and lobed; cultivated plants of Washington County, and of greenhouses elsewhere Ficus - Fruit a multiple (formed of several flowers and a common axis); leaves various . 2 2(1). Leaves crenate-serrate, palmately veined and often palmate lobed as well; flowers, both sterile and fertile, borne in catkinlike spikes; fruit seldom more than 1 cm thick Morus Leaves entire, pinnately veined, not lobed; flowers borne in dissimilar in- florescences, the sterile in racemes, the fertile in globular heads; fruit globular, more than 5 cm thick Madura Ficus L. Trees or large shrubs; leaves alternate, simple, palmately veined and lobed, the stip- ules forming a circular scar around the stem; flowers minute, numerous, borne inside a hol- low receptacle which ripens to form a syco- nium; staminate perianth 2- to 6-parted, with 1 or 2 stamens; pistillate perianth reduced or lacking; receptacles perfect or imperfect; fruits of individual flowers of achenes. Ficus carica L. Common Fig. Deciduous trees to 5 m tall, rarely more, often sprawling in age; leaves prominently veined, thick, to 25 cm long or more and to 20 cm broad, 3- to 5-lobed, the lobes undulate-serrulate; fruits obovoid, mostly 2.4-4.5 cm long and 2-3 cm thick. Cultivated fruit plant in Washington March 1980 Welsh: Utah Flora, Miscellaneous Families 47 (and formerly Garfield, at Hite) County, frost sensitive elsewhere except under glass; in- troduced from the Mediterranean region of the Old World. This is the fig of commerce; 5(iii). Maclura Nutt. Dioecious trees with hard yellow wood; leaves entire, the stipules minute, the scar not encircling the stem; staminate flowers nu- merous in loose, peduncled, axillary heads or umbels, the calyx 4-parted and with 4 sta- mens; pistillate flowers coherent in dense, globose, axillary heads, the calyx 4-lobed, the single filiform style very long; fruit a globose multiple. Maclura pomifera (Raf.) Schneid. Osage Orange. {Toxylon pomiferum Raf.) Trees to 10 m tall, rarely more; stems usually armed with stout thorns 1-2 cm long; leaves petio- late, the blades 5-10 cm long and 1.8-6.5 cm wide, ovate, entire, rounded to obtuse ba- sally, attenuate to acuminate apically; clus- ters of staminate flowers 2.5-3.5 cm across; heads of pistillate flowers 2-2.5 cm across; multiple fruit mostly 8-14 cm thick. Culti- vated ornamental and botanical curiosity of low elevation regions in Utah, long per- sisting; introduced from the eastern states. The wood of this tree is very strong, and has served as a source of bows for American In- dians and others; 5(i). Morus L. Dioecious trees; leaves palmately veined, serrate to dentate, sometimes lobed; stipules lanceolate, the scar not encircling the stem; flowers monoecious or dioecious, those of both sexes borne in stalked, axillary, catkin- like clusters; calyx 4-parted; stamens 4; styles 2, deeply parted; fruit a multiple. Leaves glabrous above and beneath or pubescent beneath only along main veins and/or in vein axils; our common mulberry M. alba Leaves pubescent over much of the lower surface, scabrous above; rarely cultivated M. nisra Morus alba L. White Mulberry. Cultivated ornamental and shade tree to 10 m tall or more; leaves obliquely ovate and crenate-ser- rate or irregularly lobed, mostly 3.5-14 cm long and 2.5-10 cm wide, truncate to sub- cordate basally, acute to acuminate apically, glabrous above and below except along veins and in vein axils; fruit 1-2 cm long and 0.6-1 cm thick, white, pink, red purple, or nearly black. Persisting and occasionally escaping in most of Utah at lower elevations; introduced from China; widespread in North America. This plant was introduced to southern Utah to provide food for silkworms in an attempt to develop a silk industry. The fruit is edible, but is consinned mainly by birds. Reports from Utah of red mulberry, M. rubra L., be- long here. Red mulberry is easily recognized by the densely hairy lower and scabrous up- per leaf surfaces. So-called fruitless phases are known; 24(v). Morus nigra L. Black Mulberry. Small trees to about 10 m; leaves cordate-ovate, crenate-serrate, seldom lobed, 5-20 cm long, 3-15 cm wide, cordate basally, obtuse to acuminate apically, scabrous above and hairy over veins and at least some intervein areas below; fniit 1-2.5 cm long and to 1 cm thick, purple to black. Sparingly cultivated orna- mental, mainly in warm regions of Washing- ton County; widely cultivated in temperate regions of the earth for its fruit; introduced from Asia; 5(o). Oleaceae Olive Family Trees or shrubs; leaves opposite (or rarely alternate), simple or pinnately compound, stipulate; flowers perfect or imperfect, borne in axillary or terminal racemose, paniculate, or thyrsoid inflorescences; calyx commonly 4- 48 Great Basin Naturalist Vol. 40, No. 1 lobed or absent; corolla usually of 4 united or distinct petals, or lacking; stamens 2, distinct; pistil 1, the ovary superior, 2-carpelled and 2-loxuled; style 1, or lacking, the stigmas 1 or 2; fruit a berry (Ligustrinn), drupe (Forestiera), loculicidal capsule {Syringa, For- sythia), circumscissile capsule (Menodora), or samara {Fraxiniis). 1. Leaves pinnately compound; fruit a samara Froxinus Leaves simple, or rarely compound; fruit various 2 2(1). Leaves ovate to orbicular, crenate-serrate; fruit a samara; plants indigenous Fraxiniis Leaves various, but seldom ovate to orbicular and crenate-serrate; fruit a drupe, capsule, or berry; plants cultivated or indigenous 3 3(2). Shrubs with yellow flowers appearing before the leaves; plants cultivated Forsythia Shmbs, subshrubs, or trees with flowers variously colored, but if yellow then not as above, and appearing with or after the leaves (before in Forestiera) 4 4(3). Corolla none or rudimentary, the flowers often unisexual; fruit a drupe; shrubs of stream banks in southeastern Utah Forestiera - Corolla well developed, the flowers perfect; fruit a berry or a capsule 5 5(4). Corolla yellow; fruit a membranous, circumscissile capsule; plants indigenous subshrubs of southern Utah Menodora - Corolla commonly lavender to red, purple, white, or cream; fruit a loculicidal capsule or a berry; plants cultivated shrubs or trees 6 6(5). Flower clusters usually less than 6 cm long; flowers white to cream; fruit a berry Ligustrwn Flower clusters usually 6-30 cm long or more; flowers lavender to red, purple, lilac, white or cream Syringa Forestiera Poir. Sprawling indigenous shrubs; leaves oppo- site, simple, serrate to entire; flowers incon- spicuous, polygamo-dioecious, borne sessile or in cymes, appearing before the leaves; ca- lyx minute, unequally 5- to 6-cleft, or lack- ing; corolla lacking, or rarely with 2 or 3 pet- als; stamens 2 or 4; ovary 2-loculed, with 2 ovules per locule; style slender; stigma 1; fniit a dnipe. Forestiera pubescens Nutt. Desert Olive. (F. neo77iexicana Gray; Adelia neomexicana (Gray) Kuntze; A. parvifolia Gov.). Shrubs to 2 m tall or more; leaves (0.8) L5-5.5 cm long, (0.3) 0.5-2 cm wide, oblanceolate to el- liptic, entire to serrulate; staminate flowers sessile; pistillate flowers pedicellate; drupe 5-7 (8) mm long, ellipsoid, blue black. Sandy terraces along the Colorado and San Juan riv- ers and tributaries, 1280-1750 m in Grand and San Juan counties; California eastward to Oklahoma and Texas, and south to Chihuahua. The fruit is eaten by fox and by coyotes, and tlie purple-stained, stone-laden fecal pellets are to be found far from the riv- ers. Long known as F. neomexicana, our ma- terials form a portion of a complex whose definition includes those portions known as F. pubescens, and that name has priority; 8(iii). Forsythia Vahl Cultivated shrubs; leaves opposite, simple or some compound, entire to serrate; flowers perfect, showy, borne in axillary cluster of 3-5, or solitary, appearing before the leaves; calyx 4-lobed; corolla 4-lobed, campanulate; stamens 2, inserted at corolla base; ovary 2- loculed, with several ovules per locule; fruit a loculicidal capsule, with many winged seeds (ours seldom fruiting). March 1980 Welsh: Utah Flora, Miscellaneous Families 49 Forsythia suspensa (Thunb.) Vahl. C.old- Fraxinus L. en-bell. {St/rin^a suspciisa Thunb.). Shrub to 2 m tall or more; branchlets somewhat 4- Deciduous, cultivated and/or indigenous angled; leaves 6-10 cm long, ovate to lanceo- trees or .shrubs; winter buds often prominent, late, acute apically, cuneaie to rounded ba- g^y to brown or black; leaves opposite, pin- .sally, usually serrate; flowers to 25 mm long, nately compound (simple in F. anomala); golden yellow; fruit lance-ovoid, to 15 mm flowers perfect or uni.sexual, inconspicuous, long, .seldom developing. Cultivated orna- borne in panicles; calyx 4-lobed or lacking; mental, common, persisting but not spread- toroUa lacking or of 2 or more, usually dis- ing at lower elevations throughout Utah; tinct petals; stamens commonly 2; ovary 2-lo- widespread; introduced from China. Numer- <-uled; styles 1; stigmas 1 or 2; fruit a samara, ous horticultural varieties are present; 2(o). 1. Leaves normally simple, sometimes with 1 or 2 leaflets below the terminal one; indigenous shrubs or .small trees of eastern and southern Utah F. anomala — Leaves normally pinnately compound with 5-9 or more leaflets; trees, either indigenous or cultivated 2 2(1). Branchlets, petioles, and axis of panicle commonly spreading hairy, seldom glabrous; leaflets usually 5 or fewer; trees, indigenous in southwestern Utah, cultivated elsewhere F. velutina 2. Branchlets, petioles, and axis of panicle variously hairy or glabrous, but seldom spreading hairy; leaflets usually 7 or more; trees, cultivated and sometimes escaping 3 3(2). Flowers appearing after leaves formed; corolla present F. ornus — Flowers appearing before leaves formed; corolla lacking 4 4(3). Fiiiit with calyx persisting as a campanulate cap; anthers oblong; leaflets usually 5-7 5 — Fruit with calyx early deciduous or lacking (except in F. qiiadrangulata); anthers often cordate; leaflets u.sually 9-11 or more 6 5(4). Petiolules of middle and lower mature leaflets wingless nearly their entire length; winter buds black; leaf scars horseshoe shaped; wing of fruit terminal, not or only slightly decurrent F. americana — Petiolules of middle and lower mature leaflets winged nearly to the base; win- ter buds brown; leaf scars semicircular or shield shaped; wing of fruit decur- rent to below the middle F. pennsylvanica 6(4). Branchlets 4-sided, 4-angled; bark broken into plates; flowers with a minute, deciduous calyx F. qtiadrangulata — Branchlets terete, not or only slightly 4-angled; bark smooth or irregularly roughened; flowers with calyx lacking 7 7(6). Leaflets glabrous or somewhat hairy along veins beneath; commonly cultivated tree F. excelsior — Leaflets definitely pubescent beneath, especially along the veins, the long red- dish hairs extended onto and along the leaf rachis; unconunon to rarely cultivated tree F. nigra Fraxjnus americana L. White Ash. Moder- usually 7 (5-9) 6-15 cm long, petiolulate, ate to large trees; branchlets terete, green to ovate to lanceolate, acuminate apically, cu- brown, glabrous; winter buds black; leaflets neate to rounded ba.sally, entire to serrate, 50 Great Basin Naturalist Vol. 40, No. 1 glaucous beneath and usually glabrous; an- thers oblong, apiculate; calyx persistent; co- rolla lacking; samaras (20) 25-35 (50) mm long, 4-7 mm wide, the wing not decurrent along the terete base. Shade tree of lower elevations in Utah; introduced from eastern North America; 5(o) Fraxinus anomala Torr. ex Wats. Single- leaf Ash. Shrub or small tree, commonly 2.5-4 m tall, usually with many stems; branchlets 4-angled; leaves glabrous, ovate, crenate-serrate to subentire, 1.5-6.5 cm long, 1-6 cm wide, acute to obtuse or subcordate basally, acute to rounded or emarginate api- cally, sometimes 2- or 3-foliolate or transi- tional to simple; flowers usually perfect; an- thers oblong: calyx campanulate, persistent; petals lacking; samaras winged almos^ to the base, 12-27 mm long, 5-11 mm wide. Mixed desert shrub, mainly on rimrock or along drainages, and in pinyon-juniper woodland, 900-2150 m, in Emery, Garfield, Grand, Iron. Kane, San Juan, Uintah, Washington, and Wayne counties; Colorado, New Mexico, Arizona, and California; 80(xv). Fraxinus excelsior L. European Ash. Mod- erate to large trees; branchlets terete, glabrous; winter buds black; leaflets 7-11, 5-12 cm long, .sessile, ovate to oblong or lan- ceolate, acuminate apically, cuneate basally, serrate, green Ijeneath, glabrous except along midrib, the hairs sometimes extending to the rachis; flowers polygamous; anthers ovoid; calyx lacking; corolla lacking; samaras 25-35 (40) mm long, 5-11 mm wide, the blade de- current almost or quite to the base of the flattened body. Shade tree of habitations and streets at lower elevations throughout Utah; introduced from Europe; 11(0). Fraxinus nigra Marsh. Black Ash. Moder- ate trees; branchlets terete, glabrous; winter buds black; leaflets 7-11, mostly 6-12 cm long, sessile, lanceolate to oblong, obtuse to rounded basally, long-acuminate apicallv, serrate, green and glabrous except reddish- hairy along veins, the pubescence extending along the leaf rachis; flowers dioecious; an- thers oblong; calyx lacking; corolla lacking: siunaras mostly 25-35 nun long and 6-10 mm broad, the blade decurrent to the ba.se of the flattened body. Sparingly cultivated shade tree at lower elevations in at least the major population centers; introduced from eastern North America; 4 (o). Fraxinus ornus L. Flowering Ash. Small to moderate trees; branchlets terete; winter buds gray to brownish; leaflets usually 7 (7-11), mostly 2.5-7 cm long, petiolulate, lance-ovate to obovate (terminal one), rounded to obtuse basally, acuminate api- cally, crenate-serrate, glabrous except along midrib; flowers perfect; calyx present, per- sistent, with 4 triangular-acuminate, spread- ing lobes; petals present, linear; samaras 20-25 mm long, 3-6 mm wide, the blade ter- minal on the terete base. Rarely cultivated shade and ornamental tree of lower eleva- tions in Utah; introduced from Europe; 2(o). Fraxinus pennsylvanica Marsh. Red Ash. Moderate trees; branchlets terete, pubescent to glabrous, sometimes glandular; winter buds olive to brown; leaflets usually 7 (5-9), 6-15 cm long, petiolulate, lanceolate to lance-oblong, acuminate apically, acute to obtuse or rounded basally, serrate to entire, green and glabrous or hairy (especially along the veins) beneath; anthers oblong, apiculate; calyx campamulate, persistent; corolla lack- ing; samaras 27-40 (50) mm long, the blade decurrent to the middle of the terete body or below. Common shade tree of lower eleva- tions throughout Utah, persisting and escap- ing, in Box Elder, Cache, Davis, Iron, Juab, Millard, Salt Lake, Utah, and Washington counties; introduced from eastern North America. The escaped plants have become established along streams and on lake mar- gins at lower elevations. Much of our mate- rial has glabrous branchlets and petioles, and has been designated as F. pennsylvanica var. lanceolata (Borkh.) Sarg. (F. lanceolata Brokh.). This phase is known as green ash; 28 (o). Fraxinus quadrangulata Michx. Blue Ash. Small to moderate trees; branchlets sharply 4-angled, glabrous; winter buds black; leaflets 7-11, mostly 5-12 cm long, petiolulate, lan- ceolate to ovate-lanceolate, acute to rounded basally, acute to acuminate apicallv, serrate, glabrous except along the midrib or rarely hairy over the lower surface; flowers perfect; calyx minute, caducous; corolla lacking; an- thers cordate-oblong, blunt; samaras 20-40 (50) nmi long, the blade decurrent to the base of the flattened body. Sparingly cultivated March 1980 Welsh: Utah Flora, Miscellaneous Families 51 shade tree, at lower elevations in Utah; in- troduced from eastern North America; l(o). Fraxinus vehitina Torr. Velvet Ash, Ari- zona Ash. [F. pcnnsylvcmica Marsh, ssp. veln- tina (Torr.) G. N. Miller] Moderate trees; hranchlets terete, denselv spreading hairv to merelv sparingly so, or glabrous; winter buds brown; leaflets 3-5 (or leaves simple), lan- ceolate to ovate, elliptic, or orbicular, pe- tiolulate, cuneate to acute basally, acuminate to rounded apicallv, serrate, glabrous or hairv over the lower surface; flowers imperfect; ca- lyx campanulate, persistent; corolla lacking; anthers oblong, apiculate; samaras 16-34 mm long, 4-6 mm wide, the blade decurrent about half way along the terete body. In- digenous tree of stream courses and flood plains in Washington and Iron counties, and cultivated there and elsewhere in Utah; Ari- zona and New Mexico. The phase with co- riaceous leaflets has been treated as var. co- riacea (Wats.) Rehd. (F. coriacea Wats.), but seems not to be worthy of taxonomic recogni- tion, at least in Utah; 25(ii). Note: The shrubby Fraxinus dipetala Hook. & Am. is reported for Utah in Kearney & Peebles, 1961. Flora of Arizona, Supplement p. 1063. The related F. cuspi- data Torr. is known from adjacent Mohave and Coconino counties, Arizona, and might occur in Utah. Both species have corollas present; the former has two petals and the latter has four. LiGUSTRUM L. Shrubs; leaves opposite, simple, entire; flowers perfect, white, showy through small, borne in terminal penicles, appearing after the leaves; calyx 4-toothed; corolla 4-lobed, funnelform; stamens 2, inserted on the co- rolla tube; ovary 2-loculed, 1- or 2-seeded; fruit a berry. Ligustrum vulgare L. Common Privet. De- ciduous or semievergreen shrub to 3 m tall or more, with puberulent to glabrate branchlets; leaves 2-6 cm long, 0.8-2 cm wide, oblong to elliptic or ovate-lanceolate, glabrous; panicle dense, 3-6 cm long; corolla tube shorter than the lobes, white; anthers exserted; fruit 6-8 mm long, black, ovoid to subglobose. Culti- vated hedge plant throughout Utah at lower elevations, persisting and escaping; in- troduced from Europe; 3(o). Menodora Humb. & Bonpl. Subshrubs; leaves alternate or the lower- most opposite, simple, sessile or nearly so; flowers perfect, arranged in cymes; calyx 5-to 15-lobed; corolla yellow, subrotate, 5- to 6- lobed; stamens 2, inserted on the corolla tube; ovary 2-loculed, with 2-4 ovules per locule; style slender, the stigma capitate; fiiiit a circumscissile capsule. Steyermark, J. A. 1932. Revision of the genus Menodora. Ann. Missouri Bot. Card. 19: 87-176. Menodora scabra Gray. Plants erect or as- cending, commonly 2-3.5 dm tall, woody at the base only; leaves 0.5-2.9 cm long, 0.2-0.5 cm wide, narrowly elliptic to oblong or lan- ceolate, glabrous or scaberulous; calyx mi- nutelv puberulent, the lobes linear; corolla bright yellow, subrotate, the lobes 5-9 mm long; capsule 8-12 mm thick, membranous; seeds 4-5 mm long. Pinyon-juniper commu- nity, known in Utah only from Garfield, San Juan, and Washington counties; California, Arizona, New Mexico, Texas, and Mexico; 3(1). Syringa L. Shrubs or small trees; leaves opposite, simple, petiolate; flowers perfect, in terminal or lateral panicles; calyx campanulate, 4- toothed to nearly truncate, persistent; corolla tubular, the limb 4-lobed and rotate or nearly so; stamens 2, inserted on the corolla tube; ovary 2-loculed, each locule with usually 2 ovules; style with a 2-lobed stigma; fruit a loculicidal capsule. Flowers cream to whitish, borne in large panicles; corolla tube 1-2.2 mm long, only half as long as the calyx; fragrance musky, not that of lilac; plants flowering in summer, often treelike 2 Flowers lilac, violet, purpli.sh, or white; corolla tube mostly 6-12 mm long or more, several times longer than the calyx; fragrance usually of lilac; plants commonly shrubs, flowering in spring or .summer 3 52 Great Basin Natuhalist Vol. 40, No. 1 2(1). Leaves ovate, rounded or subcordate basally, the veins prominent on the lower surface S. amiirensis — Leaves lanceolate to elliptic or ovate-lanceolate, obtuse to cuneate basally, the veins not prominent S. pekinensis 3(1). Panicles from terminal buds; leaves of current season borne on branch with panicle; plants flowering in summer S. villosa — Panicles from lateral (or terminal) buds, the terminal buds often lacking; leaves of current season not borne on the branch with panicle; plants flowering in springtime 4 4(3). Leaves ovate to cordate, the base subcordate to obtuse; our most common species S. vulgaris — Leaves lanceolate to elliptic or ovate, obtuse to cuneate basally; common to uncommon 5 5(4). Leaves mostly less than 4 cm long, some often irregularly lobed; individual panicles short, mostly 7 cm long or less S. persica — Leaves often over 4 cm long, entire; individual panicles usually 8-12 mm long S. X chinensis Sijringa amiirensis (Rupr.) Rupr. Amur Li- lac. Shrubs or small trees to 5 nun tall or more; leaf blades 3.5-13 cm long, 1.3-8 cm wide, ovate, rounded to obtuse of short acuminate basally, acuminate apically, the lower surface hairy to glabrous, the veins prominent; petioles mostly 1-2 cm long; pan- icles 10-15 cm long, the clusters of panicles usually much longer; flowers cream to white; stamens exserted. Sparingly cultivated orna- mental of lower elevations in Utah; in- troduced from Japan; flowering in summer; 4(0). Sijringa \ chinensis Willd. Chinese Lilac. Shrub to 4 m tall or more, with spreading and often arching branches; leaves 2.5-8 cm long, 1.5-4 (5) cm wide, ovate-lanceolate, ob- tu.se to cuneate basally, acuminate apically, glabrous, the veins not prominent; petioles 0.5-1.5 cm long; panicles mostly 8-12 cm long, the clusters of panicles much longer; flowers purple lilac, or otherwise; stamens in- cluded. Commonly cultivated ornamental al- most throughout Utah; introduced from the Old World. This plant is evidently of hybrid origin, having resulted from a cross between S. persica and S. vulgaris, (j.v.; flowering in springtime; 3(o). Syringa pekinensis Rupr. Peking Lilac. Shrub or small tree to 5 m tall or more, with spreading branches; leaves 5-12 cm long, 2-4 (6) cm wide, lanceolate to ovate, cuneate ba- sally, acuminate apically, glabrous, the veins not prominent; petioles 1.5-3 cm long; pan- icles mostly 8-15 cm long, the clusters of panicles to 30 cm long or more; flowers cream to yellow white; stamens exserted. Un- common, cultivated ornamental in northern Utah, but to be expected elsewhere; in- troduced from China; flowering in early sum- mer; l(o). Syringa persica L. Persian Lilac. Shrub to 2 m tall, with upright to arching branches; leaves 1.5-6 cm long, 0.6-3 cm wide, lan- ceolate to elliptic, sometimes lobed, cuneate to obtuse basally, acute to acuminate api- cally, glabrous, the veins not prominent; pet- ioles 0.5-1 cm long; panicles mostly 3-7 cm long; flowers usually lilac but purple phases are known; stamens included. Uncommonly cultivated ornamental, especially in northern Utah; introduced from Asia Minor; flowering in springtime; 5(o). Syringa villosa Vahl. Shrub to 3 m tall, rarely more, with erect branches; leaves 4-15 cm long (or more), 2.5-9 cm wide, ovate to elliptic, acute basally, abruptly acuminate apically, spreading hairy below, especially along the prominent veins; petioles 0.8-2 cm long; panicles mostly 10-18 cm long; flowers March 1980 Welsh: Utah Flora, Miscellaneous Families 53 pink lilac to white; stamens included. Spar- ingly but widely planted ornamental, mainly in northern Utah; introduced from China; tlowering in summer; 4(o). Syringa vulgaris L. Common Lilac. Shrubs to 4 m tall or more, the branches usually erect; leaves 3-12 cm long. 1.5-8 cm wide, ovate to cordate, cordate to rounded, trun- cate or obtuse basally, acute to acuminate apically, glabrous; petioles 0.8-3 cm long; panicles mostly 10-20 cm long; flowers lilac or white, seldom purple; stamens included. Abundantly cultivated ornamental, long per- sisting, in most of Utah; introduced from Eu- rope; flowering in springtime. Many horticul- tiual forms are known; 7(o). Pyrolaceae VVintergreen Family Suffrutescent or herbaceous perennials; leaves simple, alternate, opposite, or appear- ing whorled, evergreen or much reduced and lacking chlorophyll; flowers usually perfect, regular, or irregular; calyx with 4 or 5 more or less distinct sepals; corolla with 4 or 5 more or less distinct petals (united in Ptew- spora): stamen twice as many as the petals, the anthers pendulous, opening by appa- rently terminal pores or by slits, or the an- thers erect, awnless or 2-awned; pistil 1; ovary superior, 4- or 5-loculed; style 1; fruit a capsule. 1. Plants lacking chlorophyll; leaves reduced and scalelike, reddish, brownish, purple, or yellowish when fresh, often drying dark Ptewspom Plants with chlorophyll (rarely without); leaves not reduced to scales, except rarely, commonly evergreen 2 2(1). Flowers solitary, the petals rotate or nearly so Moneses Flowers few to several, the petals concave 3 3(2). Stems leafy, though short, the leaves apparently whorled; flowers corymbose; staminal filaments dilated near the base; styles very short or lacking ChimaphUa Stems leafy at base only; flowers in elongate racemes; filaments not especially dilated at the base; styles in most species over 2 mm long Pyrola Chimaphila Pursh Low shrubs from creeping rhizomes, the stems erect or ascending; leaves evergreen, leathery, apparently whorled or some alter- nate; flowers (1) 2-several, borne in peduncu- late, umbellate corymbs; sepals usually 5, dis- tinct nearly to the base, persistent; petals usually 5, distinct, rotate-cam panulate; sta- mens usually 10, the filaments dilated and cil- iate near the base; anthers awnless, opening by falsely terminal pores on short tubes; ovary superior, 5-lobed and 5-loculed; fruit a loculicidallv dehiscent capsule. Cimaphila umbellata (L.) Bart. Pipsis- sewa. Prince's Pine. {Pyrola innbellata L.; C. occklentalis Rydb.; C. umbellata ssp. occiden- talis (Rydb.) Hulten). Plants (1) 1.5-2.5 (3) dm tall, the stems glabrous, only somewhat woody; leaves 1.5-4.5 (6) cm long, 0.5-1.5 (2) cm wide, elliptic to oblanceolate, cuneate ba- sally, sharply serrate, shining above, pale be- neath, glabrous; peduncles 4-7 (10) cm long, glabrous or minutely glandular-puberulent, often suffused with red purple; pedicels glandular-puberulent or merely puberulent; flowers 1-6 or more, umbellate-corymbose; sepals erose-ciliate; petals 5-7 mm long, pink; stamens with expanded bases ciliate; capsules 5-7 mm broad. Coniferous forests, 2300-2750 m, in Duchesne, Summit, Uintah, and Washington counties; .\laska, southward to California and Mexico, east to New Mexico and Colorado, and in the eastern United States; Eurasia. Our materials are referable to var. occidentalis (Rvdb.) Blake; 4(0). 54 Great Basin Naturalist Vol. 40, No. 1 MoNESES Salisb. Rhizomatous herbs; leaves with chloro- phyll, leathery, persistent, mainly basal, but sometimes opposite or in whorls; flowers soli- tary, nodding, borne on a long peduncle; se- pals usually 5, persistent; petals usually 5, dis- tinct, spreading; stamens usually 10, the filaments tapering to the apex, the anthers awnless, nodding, opening by means of ap- parently terminal pores; ovary superior, 5- loculed, the stigma borne on an elongate, glabrous style; fruit a loculicidal capsule. Moneses uniflora L. Single Delight, Wax- flower. (M. reticulata Nutt.; M. uniflora var. reticulata (Nutt.) Blake). Plants 0.4-1.7 dm tall; leaves (including petioles) 0.8-4 cm long, 0.6-2 cm broad, serrate to crenate-serrate; peduncles 3-15 cm long, usually with 1 or 2 bracts along its length; flowers 1.3-2.5 cm broad, white to cream; sepals 1.5-2.5 m long, ciliate; petals 7-11 mm long, spreading; style 2-4 mm long; capsule 5-8 mm broad. Moist sites in coniferous forest, 2450-3050 m, in Beaver, Carbon, Duchesne, Emery, Juab, Salt Lake, and Utah counties; widely distributed in North America; Eurasia; 9(ii). Pterospora Nutt. Plants herbaceous saprophytes, devoid of chlorophyll, tall, reddish or purplish brown, the stems arising from a bulbous cluster of coralloid roots; leaves alternate, simple, scalelike, colored like the stems; flowers nu- merous, borne in an elongate raceme, nod- ding; calyx 5-lobed; corolla urn-shaped, the tube much longer than the lobes; stamens 10, the filaments flattened, tapering to the apex, glabrous, the anthers with 2 recurved awns, dehiscent almost throughout; ovary superior, 5-loculed, the stigma borne on a short thick style; fruit a loculicidal capsule. Pterospora andromeda Nutt. Pinedrops. Plants erect, the stems simple, 2-8.5 (10) dm tall, reddish brown, succulent, arising from a cluster of roots to 5 cm in diameter, glandu- lar-hairy, leafy only near the base; racemes 3-35 cm long or more; flowers 5-8 mm long, nodding, axillary; pedicels 5-15 mm long, re- curved; sepals oblong, glandular; corolla pale yellow, depressed urn-shaped; capsule 8-12 (14) mm broad, 5-lobed, depressed globose. Coniferous forest, 2300-2900 m, in Daggett, Duchesne, Garfield, Grand, San Juan, Sum- mit, Uintah, and Washington counties, and to be expected at higher elevations elsewhere; widely distributed in North America; 15(iii). Pyrola L. Rhizomatous herbs; leaves with chloro- phyll, leathery, persistent, all basal or appar- ently so, or rarely lacking and the plants then partially or completely saprophytic; flowers regular to irregular, borne in terminal ra- cemes; sepals 5, united at the base; petals 5, distinct, usually concave, deciduous; stamens 10, the filaments tapering to the apex, the an- tliers unawned, pendulous, opening by means of apparently terminal pores; ovary superior, 4-loculed, the stigma borne on a straight or curved style; fruit a loculicidal capsule. Copeland, H. F. 1947. Observations on struc- ture and classification of the Pyroleae. Madrono 9: 65-102. 1. Styles straight or nearly so; pores of anthers sessile; stigma usuallv much broader than the style 2 Styles bent or curved; pores of anthers usually borne on short tubes; stigmas only slightly broader than the styles 3 2(1). Styles 2 mm long or less, not (or seldom) exserted from the flower; flowers not secund; petals pinkish to cream P. minor Styles over 2 mm long, exserted from the flower; flowers secund; petals greenish white P. secunda 3(1). Flowers pink to purplish; sepals longer than broad P. asarifolia Flowers pale, greenish yellow; sepals broader than long P. virens March 1980 Welsh: Utah Flora, Miscellaneous Families 55 Pyrola asarifoUa Michx. Liver-leaf Win- tergreen. {P. rotimdifolia var. bracteata (Hook.) Gray; P. asarifoUa var. hractcata (Hook.) Jeps.; P. rotimdifolia var. purpurea Bunge; P. asarifoUa var. purpurea (Bunge) Fern.; P. iiiearnata Fisch. in DC; P. asari- foUa var. inearnata (Fisch.) Fern.; P. asari- foUa var. ovata Farw.; P. uliginosa T. & G. ex Torr.; P. rotimdifolia var. uliginosa (T. & G.) Gray; P. asarifoUa var. uUginosa (T. & G.) Farw.; P. ekita Nutt.; P. bracteata var. hiUi J. K. Henry). Plants 1.3-4 dm tall; leaves basal or essentially so, the blades 1.3-7.5 cm long, 1.1-7.3 cm wide, oval, rotund, elliptic, or obovate, subcordate to rounded, obtuse, or acute basally, rounded to obtuse or emargi- nate apically, entire to serrulate; petioles 1-9 cm long; racemes mostly 2- to 12-flowered; pedicels 3-8 mm long; sepals longer than broad, 1.5-4 mm long; petals pink to pur- plish, 5-7 mm long; anthers pink, the pores on short tubes; style curved, with a flaring collar below the stigma. Coniferous and de- ciduous woods, often along streams, or less commonly in meadows, 1750-2750 m, in Daggett, Duchesne, Emery, Garfield, Grand, Iron, Juab, Piute, Rich, Salt Lake, Summit, Uintah and Washington counties (and likely elsewhere); Alaska east to Newfoundland and south to California, New Mexico, South Da- kota, and New England; Asia. Varietal status of Utah materials is not clear; .30(iv). Pyrola minor L. Lesser Wintergreen. {Amelia tninor (L.) Alef.; Erxlebenia minor (L.) Rydb.; P. minor var. conferta C. & S.; P. conferta (C. & S.) Fisch. ex Ledeb.). Plants 0.8-2.4 dm tall; leaves basal, the blades (0.4) 1.1-3.3 cm long, (0.6) 0.9-2.5 cm broad, oval, elliptic, or ovate, obtuse to rounded or sub- cordate basally, obtuse to rounded apically, crenate to subentire; petioles 0.2-3 cm long; racemes mostly 5- to 13-flowered; pedicels 2-3 mm long; sepals 1-1.5 mm long, erose to subentire; petals pale pink to cream, 3.5-4.5 mm long; anthers with pores sessile; style straight, very short, not exserted from the co- rolla, with a more or less distinctive collar below the stigma. Wet stream sides and other moist sites, usually in coniferous forests, 2150-2750 m, in Beaver, Daggett, Duchesne, Garfield, Juab, Salt Lake, Sevier, Siunmit, Uintah, and Washington counties; Alaska and Yukon east to Greenland and south to Cali- fornia and Colorado; circumboreal; ll(i). Pyrola secimdti L. One-sided Wintergreen. {Ramisehia secimda (L.) Garke; Actinocijclus secundus (L.) Klotzsch; P. secimda var. obtii- sata Turcz.; Orthilia secimda var. obtusata (Turcz.) House; P. secunda var. pumila Paine; P. secimda f. eiicycla Fern.). Plants 0.6-1.8 (2.1) dm tall; leaves basal or rarely some cauline, or sometimes with a naked stem below the leaves, the blades 1.3-4 (5) cm long, 1-3 cm wide, ovate, oval, elliptic, or orbicular, obtuse ro rounded basally, acute to obtuse or rounded apically, crenate-ser- rate; petioles 0.6-2 cm long; racemes mostly 4- to 15-flowered, the flowers secund; pedi- cels 2-5 mm long; sepals 0.5-1.5 mm long; petals greenish white, 4-6 mm long; anthers with pores sessile; style straight, exserted from the corolla, lacking a collar. Ground layer in usually coniferous forests, 2000-3350 m, in Box Elder, Carbon, Daggett, Duchesne, Garfield, Juab, Kane, Piute, Salt Lake, San Juan, Sanpete, Summit, Uintah, Utah, and Washington counties, broadly distributed in North America; Eurasia. Segregation of our materials into the various proposed in- fraspecific categories seems unwarranted; 33(vi). Pyrola virens Schweigg. in Schweigg. & Koerte. Greenish Wintergreen. {P. chlorantha Sw.; P. chlorantha var. saximontana Fern.; P. virens var. saximontana (Fern.) Fern.; P. chlo- rantha var. paucifolia Fern.; P. virens f. paucifolia (Fern.) Fern.; P. chlorantha f. paucifolia (Fern.) Camp). Plants 0.9-2.5 dm tall; leaves basal, the blades 0.6-3.5 cm long, 0.5-3 cm broad, elliptic, oval, or obovate, ob- tuse to rounded basally, rounded to obtuse apically, crenate-serrate to subentire; petioles 0.8-6 cm long; racemes mostly 2- to 9-flow- ered; pedicels 3-8 mm long; sepals 0.5-1.5 mm long; petals greenish yellow, 5-7 mm long; anthers yellowish, the pores on elongate tubes; style curved, with a flaring collar be- low the stigma. Coniferous or deciduous woods, often in moist sites, 2150-2750 m, in Daggett, Duchesne, Piute, Salt Lake, Sum- mit, and Uintah counties; widely distributed in North America; Eurasia; 8(i). 56 Great Basin Naturalist Vol. 40, No. 1 Resedaceae Mignonette Family Annual or perennial herbs with watery sap; leaves alternate, simple, or pinnately to siibpalmately divided; flowers perfect, ir- regular, borne in terminal racemes; sepals (4) 5-6 (8), distinct; petals (4) 5-6 (8), unequal in size, the upper one the largest, appendaged; stamens 8 or more, borne on the upper side of a rounded disk, the anthers 2-loculed; pis- til 1, the ovary superior, 1-loculed, with usu- ally 3 (2-6) carpels; style lacking; fruit a cap- sule, usually open at the tip before maturity. Reseda L. Erect or ascending annual or perennial herbs from a taproot; leaves alternate; flow- ers greenish yellow; sepals subequal; petals imequal; pistils 1, the carpels usually 3, open toward the apex. Reseda lutea L. Yellow Mignonette. Plants simple or much branched, glabrous; leaves pinnatifid or subpalmately divided; flowers greenish yellow, numerous, borne in elongate racemes; petals usually 6, each commonly with 3 connate or distinct appendages; ovary and capsule usually with 3 apical lobes. Cul- tivated ornamental; rarely escaping in Utah; 1(0). Tamaricaceae Tamarisk Family Shrubs or small to moderate trees; leaves alternate, scalelike, exstipulate, entire; flow- ers mostly perfect, regular, borne in spikelike racemes arranged in panicles; sepals 4 or 5, overlapping; petals 4 or 5, separate, more or less overlapping, arising from the base of a nectiferous disk; stamens usually as many or twice as many as the petals, the anthers 2-lo- culed; pistil 1, the ovary superior, unilocular, usually 3 or 5 carpelled, the placentation bas- al; stigmas 2-5, separate; ovules 2 per pla- centa; fruit a capsule, the seeds comose. Baum, B. R. 1967. Introduced and natural- ized tamarisks in the United States and Canada (Tamaricaceae). Baileya 15: 19-25. Tamarix L. Deciduous or evergreen shrubs or trees, the branchlets deciduous; leaves clasping or sheathing; flowers small, shortly pedicelled; petals white to pink or lavender, inserted be- low the disk; capsules dehiscent by 3-5 valves. 1. Leaves sheathing; evergreen trees of moderate size, restricted to Washington County T. ophylla Leaves not .sheathing, at most merely clasping; deciduous trees of small size or merely shrubs of broad distribution 2 2(1). Flowers 4-merous, or the stamens sometimes more than 4; stamens emerging gradually from the di.sk-lobes; plants uncommon both in cultivation and as e.scapes T. pairiflora Flowers 5-merous, or the stamens sometimes more than 5; stamens inserted un- der disk near the margin between the emarginate lobes; plants abundant, cultivated and otherwise T. ramosissima Tamarix aphylla (L.) Karst. Athel Tamar- isk. {Thuja aphylla L.) Trees to 10 m tall and 6 dm in diameter or more, the bark reddish brown to gray; branchlets jointed; leaves sheathing, minute, evergreen; bracts longer than the pedicels; flowers 5-merous; sepals entire, the inner ones slightly larger; petals elliptic-oblong to ovate, 2-2.2 mm long, early deciduous or with 1 or 2 persisting; sta- minal filaments inserted between the disk lobes. Cultivated sparingly in Washington County, where it seldom flowers; native to •Africa and the Middle East; introduced in California, Nevada, Arizona, and Texas; 2(i). Tamarix parviflora DC. Small-flowered Tamarisk. Shrubs or small trees to 5 m tall; bark brown to deep purple; branchlets not jointed; leaves merely sessile, not .sheathing. March 1980 Welsh: Utah Flora, Miscellaneous Families 57 deciduous with the branchlets; bracts longer than the pedicels, more or less translucent; flowers 4-merous; sepals erose-denticulate, the outer two keeled and acute, the inner flat or slightly keeled and obtuse; petals oblong to ovate, 1.9-2.3 mm long, persistent; stainin- al filaments arising gradually from disk-lobes. Cultivated and naturalized along streams and seeps, in Emery, Kane, Utah, and Washing- ton counties, and to be expected elsewhere; introduced from southern Europe and now widespread in Canada and the United States; 7(i). Tamarix ramosissima Ledeb. Branched Tamarisk; Salt Cedar. (T. gaUica authors, not L.; T. pentandra authors, not Pall.). Shrubs or small trees to 6 m tall, or rarely more; bark reddish brown; branchlets not jointed; leaves merely sessile, not sheathing, deciduous with the branchlets; bracts longer than the pedi- cels, scarious but scarcely translucent; flow- ers 5-merous; sepals erose-denticulate, the outer 2 narrower than the inner, all more or less acute; petals obovate, 1-1.8 mm long, persistent; filaments inserted under the disk near the margin between the emarginate lobes. Cultivated and naturalized along seeps, streams, and reservoirs, almost throughout Utah (Carbon, Davis, Duchesne, Emery, Gar- field, Grand, Juab, Kane, Millard, San Juan, Sevier, Tooele, Uintah, Utah, Wasatch, Washington, Wayne, and Weber counties); introduced from Eurasia, now widespread in the southern United States; 99(xix). Tiliaceae Linden or Bas.swood Family Trees; leaves alternate, simple, serrate to obscurely lobed, usually oblique, stipulate; flowers regular, perfect, borne in cymes; sep- als 5, distinct or more or less connate; petals 5, alternate with the sepals; stamens numer- ous, the filaments free or connate in bundles of 5-10; ovary superior, 5-loculed; fruit dru- paceous. TiLIA L. Cultivated trees; leaves long-petioled, the blades obliquely cordate, serrate or doubly so, sometimes obscurely lobed; flowers in long-peduncled cymes, the peduncle adnate at its base to a ligulate bract; sepals 5; petals 5; stamens numerous, distinct or in 5 clusters, sometimes bearing petaloid staminodia oppo- site the petals; ovary 5-loculed, the stigma 5- lobed; fruit subglobose, 1- to 3-seeded. 1. Branchlets and petioles densely white-hairy; leaf blades white stellate hairy beneath T. tomentosa Branchlets and petioles glabrous or nearly so; leaf blades variously pubescent or glabrous 2 2(1). Leaf blades hairy (sometimes thinly so) over the lower surface and usually along the veins beneath '^ Leaf blades glabrous beneath, except in vein axils 5 3(2). Leaf blades densely white or brown stellate hairy beneath T. heterophylla Leaf blades variously hairy but the surface not obscured by hairs 4 4(3). Hairs of lower leaf surface stellate, at least some; flowers with staminodes T. neglecta Hairs of lower leaf surface all simple; flowers without staminodes T. platijpliyllos 5(2). Leaf blades deflnitely glaucous beneath, usually less than 8 cm long; flowers lacking staminodes T. cordata - Leaf blades green or merely pale green beneath, the largest usually more than 8 cm long; flowers with or without staminodes 6 58 Great Basin Naturalist Vol. 40, No. 1 6(5). Flowers with staminodes; leaves serrate to doubly serrate with long-acuminate teeth, the largest blades on flowering stems to 10 cm long or more T. atnericana Flowers without staminodes; leaves serrate with short acute teeth, the largest blades on flowering stems usually less than 10 cm long T. europaea Tilia atnericana L. American Linden. Moderate to large trees of streets and other ornamental plantings, common in Salt Lake, Utah, and Weber counties, and probably grown elsewhere; indigenous to the eastern states and Canada; 9(o). Tilia cordata L, Small-leaved European Linden. Small to large trees of ornamental plantings; common in Box Elder, Cache, Juab, Salt Lake, Utah, and Weber counties; widely cultivated in North America; in- troduced from Europe; 12(o). Tilia X europaea L. Common or European Linden. Moderate to large trees of ornamen- tal plantings, uncommon in Utah; indigenous to Europe. This tree is reputed to be a hybrid derivative of T. cordata x T. platijphyllos; 2(0). Tilia heterophylla Vent. White Basswood. Large ornamental trees, uncommon in Utah; indigenous to the eastern United States; 2(o). Tilia neglecta Spach. Moderate to large or- namental trees, uncommon in Utah; in- digenous to the eastern United States and Canada. This taxon resembles, and apparent- ly intergrades with, T. atnericana, with which it is very closely allied; 2(o). Tilia platyphyllos Scop. Large-leaved Linden. Moderate to large ornamental trees, common in Salt Lake, Utah, and Weber counties, and probably elsewhere; indigenous to Europe; 8(o). Tilia tomentosa Moench. Silver Linden. Moderate to large ornamental trees, moder- ately common in Cache, Juab, Salt Lake, Utah, and Weber counties; indigenous to eastern Europe and Asia Minor; 5(o). THE TAXONOMIC STATUS OF THE ROSY BOA LICHANURA ROSEOFUSCA (SERPENTES: BOIDAE) John R. Ottley', Robert W. Murphy-, and Geoffrey V. Smith' Abstract. — Evidence is presented indicating that lAchanura roseofusca and Lkhanum triiirgata are conspecific. Data include the report of an intermediate specimen from El Arco, Baja California Norte, a site midway between the previously known peninsular ranges of the two species; captive hybridization provides additional support for the conclusion. The close relationship of the boas Lich- anurci triiirgata Cope and Lichanura roseo- fusca Cope has long been recognized, due principally to the overlap of most scale char- acters and because the desert boa L. roseo- fusca gracia Klauber appears to be an inter- mediate between the two species (Klauber 1931). The problem in establishing their rela- tionship stems from the rather broad gaps be- tween their known ranges in central Baja California and southwestern Arizona. Subsequent to the description of L. r. gracia, Klauber (1933) reported a single spec- imen from Guaymas, Sonora. This specimen agrees exactly with L. trivirgata in coloration but has scale counts resembling those of L.r. gracia. He stated that the specimen might be considered an intergrade of L. trivirgata and L.r. gracia. This is somewhat surprising since he restricted L. trivirgata to the cape region of Baja California, thus necessitating a trans- gulfian dispersal of trivirgata to facilitate hybridization. Gorman (1965) reemphasized the wide variation in meristic characters within the genus, as first demonstrated by Stejneger (1891), and referred to Klauber s (1933) scale counts and color descriptions as evidence indicating that the populations from southern Arizona, Sonora, and southern Baja California are all one form, L. trivirgata (all have three primary stripes of chocolate brown on a light drab background). The variation seen in the genus led Klau- ber (1931, 1933) to speculate that we might be dealing with a single, polytypic species, L. trivirgata. He suggested, however, that before such a designation be considered we should await the collection of more material from regions of potential hybridization. Gorman (1965) and Bostic (1971) com- mented on new material from the range gaps and stated that the basis was yet lacking for uniting the two species because of the great uniformity of L. trivirgata throughout its range and the absence of obvious intergrades. The range gaps were shown to be separations of approximately 160 km (100 miles) in both central Baja California and southwestern Ari- zona. In spite of these appraisals, several au- thors (Miller and Stebbins 1964, Lowe 1964, Soule and Sloan 1966) have proposed, in ad- vance of adequate evidence, to imite the two species. The needed evidence is reported in this paper. During the summer of 1979, an unusual specimen of L. trivirgata (Fig. 1) was collect- ed at the town known as El Arco, Baja Cali- fornia Norte (28°02'N, 113°27'W). The spec- imen, taken as it was crossing the road in front of the military base on 17 July at 2225 hours by Kenneth A. Stockton, is unique for two reasons. First, its coloration" and scale counts are intermediate between the two spe- cies. Second, the geographic location of El Arco is midway between the previously re- ported limits for the two species (Bostic 'Life Science Museum, Brighani Young University, Provo, Utah 84602. ^Department of Biology. UCLA, Los Angeles, California 90024. 'Alia Mira .\nimal Clinic, VLsta, California 9208.3. Color characters with numbers refer to the color-name charts by Kelly (1958). 59 60 Great Basin Naturalist Vol. 40, No. 1 1971). Scale counts are as follows: 224 ven- trals, 48 subcaudals, 41 dorsal scale rows, 15-14 supralabials, 15-16 infralabials, and 10-11 oculars. The specimen is an adult male measuring 577 mm total including the 85 mm tail. The coloration and color pattern consists of three primary stripes of deep brown (No. 56) on a ground of light gray olive (No. 109). When one considers all these characters, the El .\rco specimen appears to be the obvious intergrade spoken of by Gorman (1965). Al- though this report essentially closes the range gap on the Baja California peninsula, a gap vet remains between the Kofa Mountains and Organ Pipe Cactus National Park in south- western Arizona. Fowlie (1965) has indicated in a range map that trivirgata and grcicia overlap in the region of the Growler Moun- tains southwest of Ajo. If two subspecies are in fact found together in the area, we would expect to see the effects of intergradation. No such evidence has ever been reported or are we aware of any specimens that substantiate such a claim. We must therefore (juestion the validity of Fowlie's range for gracia in the Ajo region. Notes on Captive Breeding Recent captive breeding experiments have produced enlightening results. In April 1975 a male L.t. roseofusca from San Diego, Cali- fornia, was bred to a female L.t. trivirgata from Cabo San Lucas, Baja California Sur. On 7 August 1975 three young were born, two of which died within a few days; how- ever, the third specimen, a male, is alive at the time of this writing and in our possession (Fig. 2). Coloration and color pattern consist of three primary stripes of medium brown (No. 56) on a light olive gray (No. 112) back- ground. The stripes are moderately serrated, yet fairly uniform. Scale counts are as fol- lows: ventrals 2.31, subcaudals 47, dorsal scale rows 41, supralabials 14-14, infralabials 17-15, and oculars 10-10. Another cross, in- volving a male L.t. trivirgata from near San Bartolo, Baja California Sur, and a female of the same subspecies from the vicinity of Esta- cion Ortiz, Sonora, occurred in March 1976. Four young were born on 29 July 1976. A fe- male from that litter (Fig. 3) yet remains in our possession. Coloration and pattern are of Fig. 1. Dorsal view of a Lkhunum triim^ala x wscofusai iiitcnnfcliato tioin El Arco. Baja California \ortt March 1980 Ottley et al.: Rosy Boa 61 Fig. 2. Dorsal view of a Lichaniini tritirgata x roscufiisca hybrid; male parent is a L. t. wscofttsca from San Diego, California, and the female parent is a L. t. triiirgata from Cabo San Lucas, Baja California Sur. Fig. .3. Dorsal view of a Lichanura t. triiin^dta transgnlfian cross; male parent is a L. t. trivirgata from near San Bartolo, Baja California Sur, and the female parent is a L. t. trivirgata from Estacion Ortiz, Sonora. 62 Great Basin Naturalist Vol. 40, No. 1 three primary stripes of chocolate brown on a cream ground. The stripes are uniform with shghtly serrated edges. Scale counts are as follows: ventrals 220, subcaudals 44, dorsal scale rows 38, supralabials 13-13, infralabials 15-14, and oculars 10-11. The female L.t. tri- virgata transgulfian cross and a male desig- nated as L.t. gracia from near Punta Prieta, Baja California Norte, were observed cop- ulating on 16 May 1979. On 24 October 1979 three young were bom, all males, each bear- ing well-delineated medium brown stripes and a ground of color intermediate between the parents. Acknowledgments We thank Wilmer W. Tanner and Kent M. Van De Graaff for their constructive criti- cisms and comments in reviewing this paper, Vickie R. Ottley for typing the manuscript, and Lawrence E. Hunt, Kenneth A. Stockton, and Dale M. Stockton for their help and companionship in the field. Scientific collec- ting permit 30/832/79 was issued by Ignacio Ibarrola Bejar, director general of the De- partamento de la Conservacion de la Fauna Silvestre. Summary In coloration the El Arco specimen and the captive bred trivergata x roseofusca hybrid are very similar, differing only in the latter having moderately serrated stripes. These data support our consideration of the El Arco specimen as an intermediate. Since El Arco is situated in a geographical region midway be- tween "pure" trivergata and roseofusca, in- dicating a continuous range, we find no alter- native to considering the two taxa as being conspecific. The binomial Lichanura trivir- gata Cope has priority over L. roseofusca Cope by publication date. Accordingly, we recognize L. trivirgata as a single, polytypic species with four subspecies (L. t. trivirgata Cope, L. t. roseofusca Cope, L. t. gracia Klauber, and L. t. bostici Ottley). Literature Cited BosTic, D. L. 1971. Herpetofauna of the Pacific Coast of north central Baja CaUfomia, Mexico, with a de- scription of a new subspecies of PhyUodactylus xanti. Trans. San Diego Soc. Nat. Hist. 16(10):237-263. FowLiE, J. A. 1965. The snakes of Arizona. Azul Quinta Press, Fallbrook, California, 164 pp. Gorman, G. C. 1965. The distribution of Lichanura tri- virgata and the status of the species. Herpetolo- gica 21(4):2a3-287. Kelly, K. L. 1958. ISCC-NBS color-name charts illus- trated with centroid colors. National Bureau of Standards, NBS Circular 533. Klauber, L. M. 1931. A new subspecies of the Califor- nia Boa, with notes on the genus Lichanura. Trans. San Diego Soc. Nat. Hist. 6(20):305-318. 19.33. Notes on Lichanura. Copeia (4):214-215. Lowe, C. H. 1964. The vertebrates of Arizona: Anno- tated check lists. Tucson, University of Arizona Press. Miller, A. H., and R. C. Stebbins. 1964. The lives of desert animals in Joshua Tree National Mon- ument. Berkeley and Los Angeles, University of California Press. Ottley, J. R. 1978. A new subspecies of the snake Lich- anura trivirgata from Cedros Island, Mexico. Great Basin Nat. .38:411-416. SouLE, M., AND A. J. Sloan. 1966. Biogeography and dis- tribution of the reptiles and amphibians on is- lands in the Gulf of California, Mexico. Trans. San Diego Soc. Nat. History 14(11):137-156. Stejneger, L. 1891. On the snakes of the California genus Lichanura. Proc. U.S. Nat. Mus. 14:511-515. HESPEROPERLA HOGUEI, A NEW SPECIES OF STONEFLY FROM CALIFORNIA (PLECOPTERA: PERLIDAE) Richard \V. Baiiinaiin' and Bill P. Stark- .\bstr.\ct.— a new species of Hcspcropcila is named from northern California. The adult male and female, nymph, and egg are described and figured. Hespcroperla lioouci adds a second species to this previously monotypic genus. The genus Hcsperoperla Banks (1938) was not accepted by other workers until the re- cent world catalog (lilies 1966). Studies by Needham and Claassen (1922), Claassen (1940), and Frison (1942) placed six species in synonymy luider Acroneiiria pacifica Banks, including Hesperoperia obscura (Banks), the designated type species. Stark and Gaufin (1976), in their revision of the Perlidae, confirmed that Hesperoperia was indeed a separate genus with one valid species, Hesperoperia pacifica (Banks). In the fall of 1976, Charles L. Hogue of the Natural History Museum, Los Angeles County, sent a distinctive female perlid to the senior author for identification. When it proved to belong to Hesperoperia, the help of numerous colleagues, including Bill P. Stark, was enlisted. The types of several species in the A. pa- cifica synonymy were examined, including: A. pacifica Banks, A. nigrita Banks, A. pu- mila Banks, and A. obscura Banks. The spe- cific name nigrita suggested that its type might be dark and distinctive, but this was not the case. The type locality of A. pumila. Three Rivers, California, which is near Vis- alia, is a short distance south of the known range of this new species, but the specimen was positively H. pacifica. Several additional specimens were sub- sequently found that confirmed that this was an undescribed species of Hesperoperia with a known range that included most of north- em California. Both species of Hesperoperia occur in the same streams with no intergradation; thus they are sympatric species. Hesperoperia hoguei, n. sp. Figs. 1-9 Male.— Macropterous. Length of fore- wings (18-20 mm; length of body 16-18 mm. Dorsum of head mostly yellow, with brown U-shaped pattern connecting ocelli, posterior lateral margins brown behind compound eyes, sometimes with faint brown area near frontal margin. Pronotum with broad yellow median stripe, lateral margins dark brown, rugosities distinctive and slightly lighter than lateral margins (Fig. 4). Legs brown. Wings light brown, veins brown. Abdomen yellow with narrow brown lateral stripes. Tergum 10 with median tergite, segment covered with medium-length hairs. Sternum 9 with large quadrangular hammer. Paraprocts sclerotized, broad basally, apex narrow and pointed (Fig. 2). Aedeagus with two bands of large spinules, apical band broad, consisting of 12 or more closely set spinule rows, basal band narrow, interrupted on both dorsomesal and ventromesal surfaces; small spinules pres- ent in patches near base and at dorsal apex (Figs. 5-6). Female.— Macropterous. Length of fore- wings 26-28 mm; length of body 24-26 mm. Color pattern similar to male. Subgenital plate strongly produced, with more darkly sclerotized area along mesoposterior margin 'Monte L. Bean Life Science Museum and Department of Zoology, Brigham Young University, Provo, Utah 84602. 'Department of Biological Sciences, Mississippi College, Clinton, Mississippi 39056. 63 64 Great Basin Naturalist Vol. 40, No. 1 Fig. 1. Hesperoperht /logi/ei, mature nymph, habitus. March 1980 Baumann, Stark: California Stonefly 65 2 3 11 5 Figs. 2-6. Hesperoperla hogtici: (2) male terminalia, dorsal; (3) female terminalia, ventral; (4) adult, head and pro- notum; (5) aedeagiis, dorsal (.50X, inset lOOX); (6) aedeagiis, lateral (SOX, inset lOOX). 66 Great Basin Naturalist Vol. 40, No. 1 (Fig. 3). Vagina, spermathecum, and acces- sory glands membranous. Egg.- Outline oval; cross-section circular. Collar stalked, margin flanged and irregu- larly incised; collar end of egg with regular indentations which end at terminal margin. Chorion smooth. Micropyles arranged cir- cumlinearly in apical end opposite of collar (Figs. 7-9). Nymph.— General color dark brown, pat- terned with yellow markings. Occiput with an irregularly spaced row of spinules. Post- ocular fringe present. Head mostly dark, with distinctive inverted yellow W-shaped pattern located anterior to compound eyes, frontal margin entirely yellow. Pronotum with later- al setae sparse or absent. Abdominal terga with numerous small intercalary spinules. Cerci with fringe of spines at segmental joints, never longer than segments; few tiny intersegmental spinules present. Proventricu- lus with teeth in 12 longitudinal bands; acces- sory bands and structures absent. Thoracic and anal gills present (Fig. 1). Diagnosis.— Hespemperla fwguei can be easily separated from H. pacifica in the nym- phal stage by the difference in head pattern and the abdominal spinulation. Hesperoperla pacifica exhibits a large, inverted, mushroom- shaped pattern anterior to the compound eyes that terminates in an enlarged base on the mesoanterior margin, and H. hoguei bears an inverted W-shaped pattern and a separate broad yellow band running the full width of the anterior margin. Intercalary spinules are entirely absent from the abdominal terga of mature H. pacifica nymphs but are numerous in H. hoguei specimens. Claassenia sabulosa nymphs also have an inverted W-shaped pat- tern on the head, but they possess a complete occipital ridge. Adults of H. pacifica have a plain yellow brown pattern on their head and pronotum. Those of H. hoguei have a broad yellow me- dian area set off by distinctive dark lateral margins. The external genitalia are quite sim- ilar, but the basal spinule band on the ae- deagus of the males is different. In H. pacifi- ca the band is only broken ventrally, but in H. hoguei the band is broken both dorsally and ventrally. Types.- Holotype: $ , Gibson Creek, 800 ft, 1 mile west of Ukiah, Mendocino Co., Cal- ifornia, 6-IX-1976, C. L. Hogue. Allotype: Toadtown, 3000 ft, 4 miles SW Stirling City, Butte Co., California, 9-IX-1976, C. L. Hogue. Paratypes: Butte Co.: Paradise, 25-V-1966, Lowe, 2 females (CSUC); Tehama Co.: Big Chico Creek, Hwy. 32, 14-IX-1979, G. L. Boles, 13 males, 7 females (GLB) (BYU). Ad- ditional specimens: Butte Co.: Big Chico Creek, 580 ft, V4 mile below Salt Springs, Bidwell Park, P-VIII-1972, M. W. Kainu, nymph (UCD); 2 miles SW Stirling City, 20- VI- 1979, J. A. Stanger, nymphs (BYU). Plumas Co.: Sulphur Creek, Hwy. 89, 5-VII- 1979, B. P. Stark and K. W. Stewart, nymphs Figs. 7-9. Hesperoperla hoguei: (7) egg, lateral (200X); (8) egg, collar end {400X); (9) egg, micropyles (700X). March 1980 Baumann, Stark: California Stonefly 67 (BPS). SJiasta Co.: unnamed spring tributary to Lake Britton, 20-IX-1978, G. L. Boles, nvniphs (GLB); South Fork Bear Creek, 12- v'lI-1979, G. L. Boles, nymphs (GLB). Te- hama Co.: Big Chico Creek, Hwv. 32, 14- V- 1978, G. L. Boles, nymph (BYU),' same data, 2-IX-1978, nymphs (2 females emerged 28- IX- 1978) (GLB) (BYU). Holotype and allotype deposited at the Natural History Museum, Los Angeles Coun- ty- Etymology.— This species is named in honor of Dr. Charles L. Hogue, Senior Cura- tor of Entomology, Natural History Museum, Los Angeles County. He has collected many interesting stoneflies during his studies on the torrenticolus insects of the New World. Acknowledgments We thank the following individuals for the help that we received during this study: Ger- ald L. Boles, California Department of Water Resources, Red Bluff, California (GLB); Dr. Charles L. Hogue, Natural History Museum, Los Angeles County, California (LACM); Dr. David H. Kistner, California State University, Chico, California (CSUC); Dr. Robert O. Schuster, University of California, Davis, California (UCD); Jean A. Stanger, Brigham Young University, Provo, Utah (BYU); Dr. Kenneth W. Stewart, North Texas State Uni- versity, Denton, Texas. The drawings were made by Connie A. Be- van Bhagat. Literature Cited Banks, N. 1938. A new genus of Perlidae. Psyche 45:136-1.37. Claassen, p. W. 1940. A catalogue of the Plecoptera of the world. Mem. Cornell Agr. Exp. Station 232:1-235. Prison, T. H. 1942. Descriptions, records and systematic notes concerning western North .American stone- flies (Plecoptera). Pan-Pac. Entomoi. 18:61-73. Illies, J. 1966. Kataiog der rezenten Plecoptera. Das Tierreich, Berlin, 82, 632 pp. Needham, J. G., AND P. W. Claasse.n. 1922. The North American species of the genus Acronetiriti (Order Plecoptera). Canad. Entomoi. 54:249-255. Stark, B. P., and A. R. Gaufin. 1976. The nearctic gen- era of Perlidae (Plecoptera). Misc. Pubi. Entomoi. Soc. Amer. 10:1-77. REPRODUCTION IN THREE SYMPATRIC LIZARD SPECIES FROM WEST-CENTRAL UTAH John B. Andre' and James A. MacMahon' .\bstr.-vct.- Data on reproduction by the lizards Uta stanshiiriana. Ciotaphytus colkiris, and Cnciindophorus tigris are presented from a communitv where they are sympatric in west-central Utah. Data are compared to a previous studv of these species at the same site and to data from other sites in the United States. Lizard reproductive data from geographi- callv separated populations are important to ecologists attempting to explain a highly var- iable species characteristic. Reproduction by Uta stansburiana has been well documented in the literature (Fautin 1946, Medica and Turner 1976, Nussbaum and Diller 1977, Tanner 1965, Tinkle 1961, 1967, Turner et al. 1970, 1973, 1976, Worthington and Ar- vizo 1973, Parker 1974, Parker and Pianka 1975, Tinkle and Hadley 1975, and Goldberg 1977). Reproductive studies of Ciotaphytus coUaris and Cnetnidophorotis tigris are scarce (Fautin 1946, Turner et al. 1969, Pianka 1970, Burkholder and Walker 1973, Parker 1973, and Vitt and Ohmart 1977). This paper presents data on the reproduction by three lizard species {Uta stansburiana stansbu- riana, Crotaphytus collaris bicinctoris, and Cnemidophorus tigris tigris), coexisting in west-central Utah. We are cognizant of the limitations im- posed by the short period covered by our col- lections. Despite this, there is clearly a need for carefully collected data on reproduction by lizards, or other taxa, so that one might gather such data into a body of information used to address general evolutionary theorv (Tinkle 1969a, b. Tinkle et al. 1970). Methods Lizards were collected in Tule Valley, Mil- lard County, Utah (lat. 39°13'N,' long. 113°27'W). Tule Valley, bordered on the east and west by mountain ranges, is typical of the basin and range topography of the Great Basin Desert (MacMahon 1979). Bajadas (coa- lesced alluvial fans) slope from the bases of both mountain ranges to the playa that cov- ers most of the valley floor. The study site was located in the Tetra- dy?nia glabrata and Atriplex confertifolia communities described by Fautin (1946). The common plant species were T. glabrata, A. confertifolia, Chrysothamnus viscidiflorus, Artemisia spinescens. Ephedra nevadensis, Ceratoides (Eurotia) lanata, and Hilaria jamesii. The substrate was mostly small rocks embedded in packed soil, with localized areas containing large boulders, which were used as basking /perching sites by C. coUaris. Lizards were collected (shot) throughout the day, at three-week intervals between 1 April and 29 August 1976. Specimens were preserved in 10 percent formalin within two hours of collection. Analysis of reproductive state (for females) and measurement of snout- vent length (SVL) were made in the labora- tory. The sex of each specimen was determined by dissection. The reproductive tracts of the females were removed and the number of corpora lutea, yolked follicles and/ or oviduc- al eggs were recorded. Estimates of clutch size were based on the number of yolked fol- licles > 2.5 mm diameter and/ or oviducal eggs and corpora lutea for U. stansburiana 'This paper is a contribution from the Department of Biology and the Ecology Center, Utah State University, Logan, Utah 84322. Reprint requests sh be sent to James A. MacMahon. Present address for John B. Andre is Cape Romain National Wildlife Refuge, R.R. 1, Box 191, Awendaw, South Car, should rolina 68 March 1980 Andre, MacMahon: Utah Lizards 69 and volked follicles > 5.0 mm diameter and/or oviducal eggs and corpora liitea for C. coUam and C. tigris. Results Mean clutch size and mean SVL of sex- ually mature females of each species are list- ed in Table 1. The relationship between clutch size and SVL is illustrated for U. stanshuriona in Figure I. Analyses of similar data for C. coUaris and C. tigris showed no significant correlation (F-tests). The line in this figure was determined by linear regres- sion, the correlation coefficient is given for the data set. Uta stanshuriana females reach sexual maturity in their second growing season (10 mo. old) at about 40 mm SVL (the smallest female having yolked follicles was 37.0 mm SVL, see Table 2). Most U. stansburiana emerged from hibernation bv the first week of April and bred shortly after this time. Yolked follicles and oviducal eggs were pres- ent from 1 April to 16 May; only oviducal eggs were found from 6 Jiuie to 29 June. From 17 July through 29 August no yolked follicles or oviducal eggs were found in the females collected. Though yolked follicles and oviducal eggs were present in the fe- males from April to the end of June, females contained the most oviducal eggs between 24 April and 15 May. While we believe that fe- males laid one or two clutches of eggs in 1976, our data are not exten.sive enough on this point. Turner et al. (1970) have warned of the problem of determining clutch fre- quency with too few observations. Crotaplujtus coUaris females are sexually mature at about 85 mm SVL. Yolked follicles were present in the single specimen collected 15 May. All females collected in the first week of June contained oviducal eggs. At the end of June no females contained yolked fol- licles or oviducal eggs. Cnemidophorus tigris females attain sexual maturity abovit 73 mm SVL. Specimens col- lected during the first and last weeks of June CLUTCH SIZE 5 .Ufa • 4 , t • • • 4 4 3 • 10 9 5 f% \ • I V • 2 , t-rr ^4 4 V #^#^ *• V • • • 1 y = -6.448+0.216 X r = 0.605 36 40 42 44 46 48 50 SNOUT-VENT LENGTH (mm) Fit;. 1. Relationship hctufi-ii tlutcli si/c iiiuiiil)t'n and S\'I, mini) tor I'. sldiisbitrUm Table 1. Clutch size and SVL of adult female Vta stansburiana, Crotaphytus collaris and Cnemidophorus tigris. N SVL (mm) Clutch size Species X Range X Range Uta stanshuriana Crotaphytus coUaris Cnemidopliorus tigris 96 13 15 44.61 90.69 a3.13 .39.5-49.0 85.5-99.0 73.0-96.0 2.99 2-5 5.38 3-7 3.07 2-5 70 Great Basin Naturalist Vol. 40, No. 1 contained yolkfed follicles and/or oviducal eggs. After mid-July, no females contained yolked follicles or oviducal eggs. The first hatchlings observed were: U. stansburiana, 17 July; C. collaris, 9 August; C. Tigris, 7 August. Discussion Mean clutch size for U. stansburiana was 2.99 (range 2-5). Fautin (1946) reported a mean clutch size of 4.1 (range 3-5) from the vicinity of our study site. For northern popu- lations (Tooele Coimty, Utah) Parker and Pianka (1975) found a mean clutch size for Vta of 4.6; Nussbaum and Diller (1977) in Oregon found 3.3. Both Parker and Pianka (1975) and Nussbaum and Diller (1977) re- ported that Vta produces one or two clutches per season. At our site, one, and for some fe- males perhaps two, clutches of eggs were laid by Uta in 1976. Parker and Pianka (1975) also reported that oviducal eggs were present during a three- to four-month period; our data agree. The relationship between SVL and clutch size indicates that larger females produce larger clutches. An F test shows a significant relationship between SVL and clutch size (0.05 level). Other workers report the same relationship between SVL and clutch size for Uta stansburiana from other parts of its range (Tinkle 1961, Turner et al. 1973, Park- er and Pianka 1975, and Goldberg 1977). By mid-July Uta is in postreproductive condition: the reproductive tracts of both males and females have decreased in size. This size decrease is accompanied by an in- crease in the size of the fat bodies. Fat bodies continue to enlarge as the growing season progresses. Little information exists on the reproduc- tion of C collaris. In southern New Mexico, Parker (1973) reported a mean clutch size of 5.3 (range 3-7). Robison and Tanner (1962) reported a mean clutch size of 6.7, but the lizard was collected from many different parts of its range. Our data show a mean clutch size of 5.38 (range 3-7). The relationship between SVL and clutch size is illustrated in Figure 2. A loose correla- CLUTCH ^ SIZE 5 88 90 92 94 96 SNOUT-VENT LENGTH (mm) Fig. 2. Relationship between clutch size (number) and SVL (mm) for C. collaris. 98 lOO Table 2. Monthly clutch size and SVL of adult female Uta stansburiana. Month April May June July August SVL (mm) Clutch size N X Range X Range 21 43.17 37.0-47.5 2.39 2-4 13 44.38 41.5-48.0 3.46 2-5 20 42.73 38.0-47.5 2.45 2-4 20 42.80 38.0-49.0 2.47 2-4 22 43.50 40.0-48.0 2.76 2-4 March 1980 Andre, MacMahon: Utah Lizards 71 tion exists, with larger females producing larger clutches; an F test (0.05 level) is not significant. Yolked follicles and/ or oviducal eggs were present from mid-May to mid-June in C. col- laris females. By the end of June this species is postreproductive; the reproductive tracts are decreasing in size and the fat bodies are enlarging. Cneinidophoms tigris becomes active near the end of April. Mating occurs shortly after. Yolked follicles and/or oviducal eggs were present from June to the first week of July. Bv mid-July this species is postreproductive; they exhibit small reproductive tracts and en- larging fat bodies. Mean clutch size was 3.07 (range 2-5), whereas Fautin (1946) reported a mean clutch size of 6.7 (range 5-9). The relation- ship between SVL and clutch size is similar to that of C. coJIaris, with larger females ten- ding to produce more eggs (Fig. 3), although an F test (0.05 level) is not significant. Pianka (1970) and Vitt and Ohmart (1977) report a relationship between SVL and clutch size that is "loosely correlated" for C. tigris. Acknowledgments This work was made possible by the US /IBP Desert Biome fimded by the Nation- al Science Foundation (Grant GB32139). Linda Finchum typed the manuscript. Robert Bayn executed the figures. Literature Cited BuRKHOLDER, G. L., AND J. M. Walker. 1973. Habitat and reproduction of the desert whiptail lizard, Cncmidophonis tigris Baird and Girard in south- western Idaho at the northern part of its range. Herpetolot^ica 29:76-8.3. Fautin. R. W. 1946. Biotic communities of the northern desert shrub biome in western Utah. Ecol. Mon- ogr. 16;251-31(). Goldberg, S. R. 1977. Reproduction in a mountain pop- ulation of the side-blotched lizard, Uta stansbttr- iana (Reptilia, Lacertilia, Igiianidae). J. Herpetol. 11:31-35. MacMahon, J. A. 1979. North American deserts: Tlieir floral and faiuial components. Pages 21-82 in R. Perry and D. Goodall, eds. Arid land ecosystems: Their structure, fvmctioning and management. Vol. 1. Cambridge University Press, Cambridge. Medica, p. a., and F. B. Turner. 1976. Reproduction by Uta stansburiana (Reptilia, Lacertilia, Igua- nidae) in southern Nevada. J. Herpetol. 10:123-128. Nussbaum, R. a., and L. V. Diller. 1977. The life his- tory of the side-blotched lizard, Uta stansburiana Baird and Girard, in north-central Oregon. Northwest Sci. 50:24.3-260. Parker, W. S. 1973. Notes on the reproduction of some lizards from Arizona, New Mexico, Texas and Utah. Herpetologica 29:258-264. 1974. Home range, growth and population den- sity of Uta stansburiana in Arizona. J. Herpetol. 8:135-139. P.ARKER, W. S., AND E. R. PiANKA. 1975. Comparative ecology of populations of the lizard Uta stansbur- iana. Copeia 1975:61.5-6.32. PiA.NiCA, E. R. 1970. Comparative autecology of the liz- ard Cneinidophorus tigris in different parts of its geographic range. Ecology 51:703-719. RoBisoN, W. G., Jr., a.nd W. VV. Tanner. 1962. A com- parative study of the species of the genus Crota- phi/tus Holbrook (Iguanidae). Brigham Young Univ. Sci. Bull., Biol. Ser. 2(1):1-21. CLUTCH SIZE Cnemidophorus 72 76 80 84 88 y = -1.423 +0.051 X r= 0.333 92 96 SNOUT- VENT LENGTH (mm) 100 Fig. 3. Relationship between clutch size (number) and SVL (mm) for C. tigris. 72 Great Basin Naturalist Vol. 40, No. 1 Tanner, W. W. 1965. A comparative population study of small vertebrates in the uranium areas of the Upper Colorado River Basin of Utah. Brigham Young Univ. Sci. Bull., Biol. Ser. 7(1): 1-31. Tinkle, D. W. 1961. Population structure and reproduc- tion in the lizard Uta stansburiana stejnegeri. Amer. Midi. Natur. 66:206-234. 1967. The life and demography of the side- blotched lizard Vta stansburiana. Misc. Publ. Mus. Zool. Univ. Michigan 132:1-182. 1969a. The concept of reproductive effort and its relation to the evolution of life histories of liz- ards. Amer. Natur. 103:501-516. 1969b. Evolutionary implications of comparative population studies in the lizard Uta stansburiana. Pages 133-154 in Systematic biology, Proc. of In- ternational Conf., National Acad. Sci. Publ. No. 1692, Washington, D.C. TiNKJLE, D. W., .\ND N. F. H.\uLEY. 1975. Lizard repro- duction effort: caloric estimates and comments on its evolution. Ecology 56:427-4.34. Tinkle, D. W., H. M. Wilbur, and S. G. Tilley. 1970. Evolutionary strategies in lizard reproduction. Evolution 24:5.5-74. Turner, F. B., G. A. Hoddenbach, P. A. Medica, and J. R. La.n.nom. 1970. The demography of the lizard, Uta stansburiana Baird and Girard, in southern Nevada. J. Anim. Ecol. .39:50,5-519. Turner, F. B., P. A. Medica, and B. W. Kowalewsky. 1976. Energy utilization by a desert lizard (Ufa stansburiana). US/IBP Desert Biome Monogr. No. 1. Utah State Univ. Press, Logan. Turner, F. B., P. A. Medica, J. R. Lannom, and G. .\. Hoddenbach. 1969. A demographic analysis of fenced populations of the whiptail lizard, Cne- midophorus tigris, in southern Nevada. South- west. Nat. 14:189-202. Turner, F. B., P. A. Medica, and D. D. Smith. 1973. Reproduction and survivorshp of the lizard, Uta stan.iburiana, and the effects of winter rainfall, density and predation on these processes. US/IBP Desert Biome Res. Memo. RM 74-26, Utah State Univ., Logan. ViTT, L. J., and R. D. Ohmart. 1977. Ecology and repro- duction of lower Colorado River lizards: II. Cne- midophorus tigris (Teiidae), with comparisons. Herpetologica 33:223-234. WoRTHiNGTON, R. D., AND E. R. Arvizo. 1973. Density, growth and home range of the lizard Uta stansburiana stejnegeri in southern Dona Ana County, New Mexico. Great Basin Nat. .33:124-128. HAPLOPAPPUS ALPINUS (ASTERACEAE): A NEW SPECIES FROM NEVADA Loran C'. Anderson' Abstr.\ct.— The new species, llaplopapptis alpinus of section Toncstus, is forniallv described and illustrated. It is endemic to the high mountains of central Nevada. Vegetative and floral morphology of related species is detailed. The new species appears to he most closely related to H. exirnius but also demonstrates close affinity to H. aberrans (all three are diploids with n = 9). In North America, Haplopappus (Aste- raceae) contain.s about 95 .species represented in 17 sections. Additional species— including the type species, H. ghttinosus—Sire in South .\merica. Chronio.somally, two major group- ings can be identified in the genus (Anderson at al. 1974). One group is generally her- baceous and chromosomallv based on x = 4, 5, or 6. The other, more woody group, is based on x = 9. These groupings are poorly distinguished by growth form. Woodiness in the "herbaceous group" is seen in sections Isoconia and Hazardia (correlated with their xeromorphy?), and reduced woodiness is seen in the "shrubby group" in sections Tonestus and Stcnotus where their growth forms are apparently related to their montane or alpine habitats. The generic integrity of this assemblage has been challenged. Many feel that Hap- lopappus is unnatural and should be broken up (Shinners 1950, Anderson 1966, Turner and Sanderson 1971, Clark 1977, Urbatsch 1978); some would raise each section to ge- neric standing. Others feel many of the sec- tions are interrelated; i.e., the South Ameri- can taxa and Hazardia (Grau 1976). Jackson (1966) has demonstrated genetic relationship among many of the sections of the "her- baceous group" through intersectional hybr- idizations; he has recently (pers. comm.) hybridized South American taxa with those of .section Hazardia. Until more is known about the biology of Haplopappiis (especially the South American taxa), Hall's con.servative generic treatment (1928) seems better at the moment than the alternative of elevating each section to ge- neric status. Possibly a half dozen genera are represented in the North American material, but I can not envision a precise treatment now. Description of a new species in section To- nestus presents a problem. That section is in the x = 9 group, whereas the type species, H. ghitinosus, is n = 5 (personal count in 1971 from Kew Garden material; Grau 1976). So, in retaining the wider generic inter- pretation at this time, a species of Hap- lopappus will be named that most likely will be transferred to another genus when the complex is better known. Deferring descrip- tion of this species until a comprehensive ge- neric revision is available might possibly withhold additional data that would be sup- portive of the ultimate revision. Methods and Materials Fresh and dried materials were processed as in Anderson, 1964. Five heads were mea- .sured for involucral and floral data. Cytolo- gical methods are those of Anderson, 1966. Plant materials were collected personallv in the field or .supplied by Sherel Goodrich. Vouchers for various measurements and chromosome counts are at FSU. Taxonomy Haplopapfnis alpinus L. C. Anderson & S. Goodrich, sp. nov. 'Department of Biological Science, Florida State University, Tallahassee, Florida .33206. 73 74 Great Basin Naturalist Vol. 40, No. 1 Herba perennis et lignosa, 0.5-2.0 dm alta; stirpes glandulosae; folia in basi obovata vel oblanceolata, serrata vel dentata, 3-7 cm longa, 10-36 mm lata, folia caulina aliquan- temis angustiora et serrata, 3-5.5 cm longa, 8-18 mm lata; inflorescentia vel mon- ocephala vel cyma paucis cum capitibus; in- volucra 10-12 mm longa, circa 7 mm lata, phyllariis 21-28, exterioribus ovatis et folio similibus et glandulosis, interioribus bracteis angustioribus; disci florum 29-55, flavi, co- rollis 5.8-7.1 mm longis, lobis circa 1.3 mm longis, lineis stigmaticis saepissime longitu- dine paribus styli appendicibus; achaenia 4-5 mm longa et pubescentia. Type: Nevada, Nye Co., granitic rocks at 10,600 ft on 11,077-ft peak on Toiyabe Crest between Washington Creek and Aiken Creek, 24 air mi SSW of Austin, 21 Jun 1979, L. C. Anderson 4885 (BRY-holotype!, FSU!, m^!, UC!). Perennial herb, woody only at base, short rhizomatous, (0.5)0.7-1.0(2.0) dm tall; stems branched only in the inflorescence or mon- ocephalous, densely glandular-pubescent; fo- liage dark green, glandular-pubescent, basal leaves obovate to oblanceolate, petiolate, ser- rate to deeply toothed above the middle, 3-7 cm long, 10-36 mm wide, moderately vis- cous, cauline leaves oblanceolate to spatu- late, cuneate or clasping the stem, saliently dentate, 3-5.5 cm long, 8-18 mm wide; in- florescence usually monocephalous (open sites) or with up to 5 heads in an elongate or flat-topped cyme (deep crevices or protected sites); heads campanulate to hemispheric, 10-12 mm long, 7-10 mm wide (pressed), phyllaries 21-28, outer ones nearly as long as involucre, leaflike, broadly ovate, 3-nerved, glandular, slightly spreading, obtuse with small mucro, inner bracts narrower, lanceol- ate-spatulate, with finely ciliate margins, acuminate-cuspidate; ray flowers absent; disk flowers (29)35-50(55), golden-yellow, corollas (5.8)6.4-7.1(7.6) mm long, lobes (1.0)1.3(1.6) mm long, lanceolate, slightly spreading to re- curved; anthers about 2.6 mm long, appen- dages 0.6 mm long, style branches slender, stigmatic lines nearly as long as style appen- dages; achenes cylindric to hisiform, 4-5 mm long, pubescent, pappus dull white, 6-7 mm long; n = 9 (Fig. 1). Infrequent on boulders, talus, or rocky summits near treeline (primar- ily on light-colored granites but occasionally on basalt, andesite, metamorphics, or lime- stone), 9,000-11,000 ft, Toiyabe and To- quima mountains of southern Lander and Nye counties, Nevada. Mid-July-September. Additional specimens examined: Nevada, Lander Co., peak between Aiken and Carsely Creek, S. Goodrich 12137 (FSU, UTC); Nye Co., type locality, S. Goodrich 12126 (FSU, NY, UTC), head of left fork San Juan Creek, S. Goodrich 11997 (BRY, UTC), S. Goodrich 12006 (FSU, UTC), McLeod Creek, S. Good- rich 13437 (BRY, FSU), crest between Tim- Fig. 1. Representative, but somewhat stout, specimen of//, alpinus; drawn largely from Goodrich 12233 (FSU). March 1980 Anderson: A Nevada Haplopappus 75 blin Creek and Marysville Canyon, S. Good- rich 12226 (BRY), toiyabe crest at French VABM, S. Goodrich 12233 (FSU), right fork Stewart Creek, S. Goodrich 13502 (BRY), top of Shoshone Mtn., Toquima Range, S. Good- rich 6 F. Smith 13267 (FSU). All collections but the last came from the Toiyabe Range. Goodrich (pers. comm.) also reports seeing a population on Mt. Jefferson at 11,000 ft at the head of left fork of Barker (Shipley) Creek in the Toquima Range. The species is found for about 23 miles along the crest of the Toiyabe Range from the Lander-Nye County line south to the head of Stewart Creek and is reported for two sites in the To- quimas. At the type locality, H. alpinus occurs on windswept slopes above the treeline with H. macronema, ChrysotJiamnus viscidiflorus, Erigeron compositus, and Eriogonian iim- bellatU7n. Some sites are at or just below treeline, where the species occurs on rocks in scattered Pinus flexilis or Cercocarpus ledi- foUiis. Other alpine endemics from central Nevada that have been found in the vicinity of H. alpinus are Draba arida, Eriogonum ovalifolium var. caelestinum, Geranium toquimense, Hackelia sp. nov., Senecio sp. nov., and Smelowskia holmgrenii. Relationships and Phytogeography This species belongs to section Tonestus and is related to H. aberrons and H. eximius (Figs. 2-4). Comparative floral features for the three species plus the more distantly re- lated H. peirsonii are given in Table 1; all are distinctive. The Nevadan endemic is like H. abcrrans in its eradiate heads of similar size; however, the latter has differently shaped cauline leaves that overtop the racemosely disposed, turbinate to narrowly campanulate heads. Also, the bracts are more numerous, narrower, and somewhat squarrose in H. abcrrans compared to H. alpinus. The new species is more similar to H. eximius in leaf size and shape, and, although the phyllaries are fairly similar, the campanulate heads differ with those of H. eximius, being radiate with shorter involucres. Haploppapus alpinus also differs from H. eximius in the following minor floral features: H. alpinus has stouter corolla tubes (as in Chrijsothamnus spathu- latus versus C. viscidiflorus; illustrated in An- derson 1964); its pappus is shorter than the corolla length, whereas pappus equals corolla length in H. eximius; and it has longer corolla lobes. Original meiotic chromosome counts for the taxa include: H. alpinus, n = 9 {Ander- son 4885, the type collection); H. abcrrans, n = 9 {Anderson 3660 from Blaine Co., Idaho); and H. peirsonii, n = 45 (Anderson 4326 from Inyo Co., California). The count for H. peirsonii agrees with the earlier count by Stebbins, who also reported H. eximius as a diploid (Howell 1950). The other counts rep- resent first reports for those species. Meiosis appeared normal with pairing as bivalents in all instances. Pollen stainability for Anderson '■JU Figs. 2-4. Flowering heads and individual outer bracts; heads scaled to 5 mm bracket, individual bracts slightly enlarged. Fig. 2. H. aherrans (Anderson 3660, FSU). Fig. 3. //. alpinus (C^oodricli 12137, FSU). Fig. 4. //. eximius [Anderson 4320, FSU). 76 Great Basin Naturalist Vol. 40, No. 1 4885 was 97.7 percent, and there were no in- dications of apomixis in H. alpinus. Haplopappus aberrans was originally de- scribed as an aberrant member of section Macronema and was later tentatively as- signed to Tonestus (Hall 1928), but until now it has been considered poorly placed in To- nestus (Cronquist 1955). With the addition of H. alpinus, the section houses H. aberrans more comfortably. Haplopappus alpinus is .somewhat inter- mediate between H. aberrans and //. eximius morphologically and geographically. Harper et al. (1978) consider the Great Basin moun- tain ranges flori.stically as islands in the sur- rounding desert possibly populated by migra- tions from the "mainland" mountain systems of the Rockies or Sierras. The introduction of H. alpinus from eximiuslike precursors from the Sierra Madre seems very plausible. The possible origin of H. aberrans in the Saw- tooth Mountains of Idaho from alpinuslike stock poses an interesting situation. Hap- lopappus aberrans occurs in the Rocky Mountain "mainland" system (in which H. hjallii and H. pijgmaeus of Section Tonestus occur). Still, its affinities lie with H. alpinus and H. eximius. The species could represent the culmination of a migration from the western Sierra mainland across the Great Ba- sin deserts to the Sawtooths and the eastern mainland. These mountain groups were less isolated in the relatively recent past (Harper et al. 1978), and such a migration is plausible. Billings (1978) suggests the alpine flora in the Great Basin may have resulted from "up- ward evolution" of preadapted desert species of lower elevations. This doesn't .seem to ap- ply to H. alpinus or other Tonestus taxa. Bil- lings further suggests that due to reduced habitat diversity in Great Basin mountains there is a trend toward edaphic endemism. Again, H. alpinus does not follow the trend; it has been collected on granite, basalt, meta- morphics, andesite, and limestone in the geo- logically diverse Toiyabe Mountains. Clearly, these alpine areas of central Nevada do need more vegetational work, as Billings (1978) observed. Our ideas of that region may be greatly changed with further study; endem- ism apparently is not as low as Harper et al. (1978) record. Goodrich, for example, has found several imdescribed endemics in his current survey of the region. T.\BLE 1.— .\veraged floral data (and ranges entered parenthetically) for .selected taxa of Haplopappus, section To- nestus. Taxon anc i collection H. aberrans Anderson 3660 H. alpinus H. eximius H. peirsonii Feature Anderson 4885 Goodrich 12137 Anderson 4320 Anderson 4899 Anderson 4326 Involucre Bract number Length, mm 42.0 (37-49) 11.0(10-12) 24.2 (21-25) 10.6(10-11) 23.8 (21-28) 11.0(10-12) 27.2 (25-30) 8.5 (8-9.5) 29.0 (24-35) 9.0 (8.2-10) 27.0 (23-29) 15.2 (13-16) Rav flowers Flower number Flower length, mm Flower width, mm - - - 13.6(11-19) 10.5 2.2 15.0 (12-20) 10.2 2.1 21.8(21-23) 10.4 4.4 Disk flowers Flower number Flower length, mm Lobe length, mm Stigmatic area— total style branch, % 38.0 (29-48) 7.6 (7.0-8.2) 1.0(0.9-1.1) 51.7 (50-54) 41.6 (.35-49) 6.3(5.8-7.1) 1.3(1.0-1.6) 50.0 (43-55) 44.4 (29-55) 7.1 (6.4-7.6) 1.3(1.2-1.4) 48.1 (45-50) 59.2 (54-69) 7.1 (6.6-7.4) 1.1(1.0-1.2) 56.3 (44-62) 51.6(44-60) 6.4 (6.0-7.0) 1.0(0.7-1.2) 53.7 (50-56) 55.8 (45-75) 8.4 (7.0-9.5) 1.1(1.1-1.2) .38.1 (33-43) March 1980 Anderson: A Nevada Haplopappus 77 Acknowledgments Appreciation is expressed to Sherel Good- rich for guiding me to the alpine collection sites and for supplying habitat data and sev- eral collections. Melanie Darst prepared the line drawings; Dr. Walter Forehand is thanked for assistance with the Latin diag- nosis. This study was supported by National Science Foundation grant DEB 76-10768. Literature Cited Anderson, L. C. 1964. Taxonomic notes on the Chrijso- thamnus viscidiflorus complex (Astereae. Com- positae). Mandrono 17:222-227. 1966. Cytotaxonomic studies in Chitj.sothdmiuis (Astereae, Conipositae). .\nier. ]. Bot. 53:204-211. .\nderson, L. C, D. W. Kyhos, T. Mosquin, A. M. Powell, .\nd P. H. Raven. 1974. Chromosome numbers in Conipositae. IX. Haplopappus and other .\stereae. .\mer. J. Bot. 61:66.5-671. BiLLLNGS, D. W. 1978. .\lpine phytogeography across the Great Basin. Great Basin Nat. Mem. 2:105-117. Clark, W. D. 1977. Chemosystematics of the genus HaMrdia (Conipositae). J. Ariz. .\cad. Sci. 12:16. Cro.nquist, .v. 1955. Conipositae. V. In: C. L. Hitch- cock, A. Cronquist, M. Ownbey, J. W. Thompson (eds.). Vascular plants of the Pacific Northwe.st, Univ. of Washington Press, Seattle. Grau, J. 1976. Chromosomenzahlen von Siidanierikanis- chen Haplopappus ,\rten. Mitt. Bot. .Miinchen 12:40.3-410. IL\LL, H. M. 1928. The genus Haplopappus, a phyloge- netic studv in the Conipositae. Carnegie Inst. Publ. .389:i-.391. Harper, K. T., D. C. Freeman, W. K. Ostler, and L. G. Klikoff. 1978. The flora of Great Basin moun- tain ranges: diversity, sources, and dispersal ecol- ogy. Great Basin Nat. Mem. 2:81-103. Howell, J. T. 1950. Studies in California Aplopappus. Leafl. West. Bot. 4:84-88. Jackson, R. C. 1966. Some intersectional hybrids and relationships in Haplopappus. Univ. Kans. Sci. Bull. 46:47.5-488. Shlnners, L. H. 1950. Notes on Texas Conipositae, IV, V. Field & Lab. 18:25-42. Turner, B. L., and S. Sanderso.n. 1971. Natural hybridi- zation between the Composite "genera" Machae- ranthera and Haplopappus (sec. Blepharodon). Amer. J. Bot. .58:467. Urbatsch, L. E. 1978. The Chihuahuan Desert species of Ericaincria (Compositae: .\stereae). Sida 7:298-303. MISCELLANEOUS PLANT NOVELTIES FROM ALASKA, NEVADA, AND UTAH Stanley L. Welsh' and Sherel Goodrich' \BSTRACT - Described as new to science are Ahwnia ar^ilhsa Welsh & Goodrich, from Utah and Colorado; An- drosace alaskana Gov. & Standi, var recdae Welsh & Goodrich, from Kohlsaat Peak, Alaska; Lepidium osten Welsh & Goodrich, from Beaver Gounty, Utah; Lygode.mia entrada Welsh & Goodrich, from Grand Connty, Utah; Pedto- cactm des-painii Welsh and Goodrich, from Emery County, Utah; and Senecio toiyabensis Welsh and Goodrich, from the Toiyabe Range in Nye Coimty, Nevada. Several undescribed and unnamed taxa have accumulated at the herbarium of Brig- ham Young University in recent years. Mainly they represent materials which have been treated within other taxa, or they are oddities of exceedingly limited areal extent. Some have been known for long periods of time. Others are only recently discovered. All are unique in one or more ways and of suffi- ciently limited apparent distribution as to be candidates for inclusion on lists of sensitive species. Abronia argillosa Welsh & Goodrich, sp. nov. Plantis similis Abronia fragranti Nutt. et Abronia elliptica in habitas sed differt in an- thocarpus alarum nuUis rostro nuUo gracilior- ibus, in receptaculo conico et fructibus in- structis superioris foliis floribus et anthocarpis glabris, et floribus paucioribus. Plantae perennes e caudicibus ramifican- tibus gracilibus (6) 15-30 cm altae; caules glabri ad basim nibellos saepe frondosi om- nino; folia (5) 15-35 mm longa, 3-35 mm lata elliptica cvato obovata vel suborbiculares glabra; pedunculi 1-8 cm longi glabri vel puberulentes raro; bractae 7-15 mm longae, 6-15 mm latae ovales vel orbiculares sca- riosae glabrae vel ciliolatae; flores 15-22 in quoque inflorescentiam; perianthi tubus 10-15 mm longus viridis glaber vel raro pub- enilentus limbus ca. 6 mm latus albidus; re- ceptaculiun breve conicum ferens fnictus in dimidio superiore anthocarpus sine alis sine rostris plicatus leviter vel nullus 7-9 mm longus 3-4 mm latus scariosus glabri rugosus alborostratus; semina 2.5-3 mm longa 1 mm lata. Holotype: Utah, Grand Co., T22S, R24E, Sec 18, ca. 6 miles due south of Cisco at ca. 4300 feet elev., on Mancos Shale Formation, in an Atriplex community, S., E., and M. Welsh 16689, 30 May 1979 (BRY, 8 isotypes to be distributed). Paratypes: Utah, Grand Co., Fifteen miles east of Thompson, B. F. Harrison et al 10403, 16 June 1941 (BRY); first escarpment north of Thompson, west of Sego Canyon, S. L. Welsh 6943, 1 May 1968 (BRY); T22S, R24E, Sec 7, ca. 4 mi. south of Jet. 50-6 and U-128, S. L. Welsh and K. Taylor 14637, 28 April 1977 (BRY); T18S, R25E, Sec 27, ca. 20 mi. NE of Cisco, S. L. Welsh 14916, 8 June 1977 (BRY). Uintah Co., Ca. 2 mi. S of Dragon, S. L. Welsh 5379, 13 May 1966 (BRY). Colorado, Mesa Co., ca. 5 mi. W of Mack along US Hwy 6-50, L. C. Higgins and S. L. Welsh 1034, 14 June 1967 (BRY); ca 13 Km due NW of Mack, T9S, R104W ca. Sec 11, A. Cronquist 11427, 25 May 1976 (BRY;NY). The clay verbena, Abronia argillacea Welsh and Goodrich, is restricted to the Grand River Valley, and less commonly in the drainage of the White River in east-cen- tral Utah and west-central Colorado, where it occurs on heavy soils derived from Mancos Shale and Green River formations. It seems probable that the taxon was taken much pri- or to the specimens cited above, but was 'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 78 March 1980 Welsh, Goodrich: Plant Novelties 79 KHT Fig. L Abronia argillosa: A, habit of plant. 80 Great Basin Naturalist Vol. 40, No. 1 overlooked due to the superficial resem- blance to phases of A. elliptica A. Nels. and to A. fragrans Nutt. ex Hook. The plants are glabrous or essentially so and possess orbicular leaves which are glaucous. The genus has been revised by Gal- loway (1975), who distinguishes A. elliptica (common and widespread in Utah) from A. fragrans (cited from San Juan County only) by technical characters of the anthocarps. The following key will distinguish A. argil- losa from those taxa. 1. Anthocarps with 2 wings, these folded together to form a deep groove; plants of western Colorado and westward A. elliptica A. Nels. — Anthocarps wingless, or, if winged, then the wings not folded together; plants of eastern Utah and eastward 2 2(1). Anthocarps beaked, deeply grooved or narrowly winged; leaf blades variable in shape, glandular-puberulent to glandular-pubescent; plants usually of sandy soils in San Juan Co., Utah, and eastward in Colorado A. fragrans Nutt. ex Hook — Anthocarps beakless, slightly or not at all folded; leaf blades mainly orbicular, glabrous; plants usually of clay soils in Uintah and Grand counties, Utah and Mesa Co., Colorado A. argillosa Welsh & Goodrich Androsace alaskana Gov. & Standi, ex Hul- ten var. reedae Welsh & Goodrich, var. nov. Similis Androsace alaskana Gov. & Standi, in scapis numerosis floribus sessilibus solitariis vel binatum, sed differt scapis numerosioribus brevioribus gracilioribus pubescentioribus, foliis integris parvioribus dense villosis, et flo- ribus parvioribus. Herbae annuae vel biennis; scapi 25-40 in quoque rosellam 10-45 mm longi filiformes 0.3-0.5 mm in diametrum pubentes parce vel dense pilis furcatis apprime infra flores juxta; folia 5-10 mm longa 1-3 mm lata linearia vel spathulata integra vel denticulata cum 1-2 dentibus glabra infra vel pilis ad costam juxta apicem supra dense villosa pilis multi- cellulosis furcatis vel simplicibus apprime ul- tra medium; flores solitari vel imparibus ali- quando bractaea subtenda tubus calycis subaequali; tubus calycis 2-3 mm longus, dentibus circa 1.5 mm longis; corolla alba tubo calyce subaequali lobus circa 2 mm longis; capsulae maturae ignotae. Holotype: Alaska, Lat. 62°12'N., Long. 152°47'W, ca. 2 mi. SSW of Kohlsaat Peak, near VABM 5048, at 4900 feet (1495 m) elev., on rocky ridge top, K. Reed 5857, 29 June 1977 (BRY, isotype at Leningrad). Par.\type: Yukon Territory, Canada, Mount St. Elias Quad.: Outpost Mt. at south end of Kluane Lake, 60°56'N, 138°22'W, at ca. 2140 m., D. F. Murray 3014, 22 Julv 1969 (BRY; ALA). The materials herein segregated as var. reedae represent the slender peduncled, sub- entire- to entire-leaved, smaller, flowered high elevation phase of A. alaskana in interi- or Alaska and southwestern Yukon. Tliat the morphological differences noted in the pro- tologue might be the result of ecological re- sponse has been considered. The tendency to entire leaves and less pronounced ciliate mar- gins, along with slender peduncles and flow- ers that seem to average smaller, indicate a syndrome of characteristics which should re- ceive at least some taxonomic recognition. The plant is named to honor the collector of the holotype, Katherine Reed of Ancho- rage, Alaska. Lepidium ostleri Welsh & Goodrich, sp. nov. Habitu Lepidio nana S. Wats sed differt in inflorescentia longiore floribus numer- osioribus foliis longioribusque pubentioribus et sepalis pubentioribus. Herbae perennes pulvinatae caespitosae caudice ramoso folium basim marcescentibus; caules fnictifri 10-35 mm alti hirsuti; folia 4-15 mm longa hirsuta linearia et integra vel basalia 3-5 lobatis vulgo caulina nulla vel pauca; racemi circa 1 cm longi in florem et March 1980 Welsh, Goodrich: Plant Novelties 81 mm Fig. 2. Androsace alaskana: A, adaxial side of leaf; B, abaxial side of leaf; C, elose-iip of stem and inflorescence; D, habit of plant. 82 Great Basin Naturalist Vol. 40, No. 1 :pi.^ I mm mm '•'iiLv-M /•; Fig. 3. Lepidiiim ostleri: A, habit of plant; B. close-up of leaves showing variation; C, close-up of flower; D, close- up of fniit; E, close-up of inflorescence. March 1980 Welsh, Goodrich: Plant Novelties 83 1-2 cm longi in fructem 5-25 floribus; pedi- celli in fructem 2-3 mm longi; sepala 1.3-2 mm longa obtusa hyalina aspre pilosa saepe purpurascentia; petala 2-3 mm longa alba purpiirascentia; fnictus 2.5-3 mm longa, 2.3-2.5 mm lata ovata late usque ad 1 mm crassa sinus 0.2 mm profundus; stylus 0.4-0.7 mm longus. Holotype: Utah, Beaver Co., T27S, R13W, Sec 23 (SW'/4), San Francisco Moun- tains, at Frisco, on rocky ridge, in pinyon- jimiper community, K. Ostler and D. Ander- son 1258, 6 June 1978 (BRY). Paratypes: Utah, Beaver Co., T27S, R13W, Sec 23 (NW1/4), near Frisco, 6900 feet, rocky slopes, K. Ostler and D. Anderson 1210, 1 June 1978 (BRY); do, T27S, R13W, Sec 16 (SE^/i), San Francisco Mountains, near Utah Hwy 21, at 5800 ft, on west slope, dry limestone, Atriplex community, K. Ostler 1415, 19 June 1978 (BRY); do, west slope of San Francisco Mountains, bristlecone pine, ponderosa pine, Douglas fir, white fir com- numity, K. Ostler 1588, 5 July 1978 (BRY). The obvious relationship of this perennial dwarf species lies with the Lepidium nantim, an endemic of Nevada. The longer racemes, greater flower number, and dense pub- escence constitute the most important diag- nostic features. The petals of Lepidium ostleri are white, while those of L. nanum are yel- low to cream yellow. While probably of less importance than other features, the color of the petals becomes important when taken with the differences in raceme, flower num- ber, and pubescence features. The species is named in honor of its collec- tor. Dr. Kent Ostler, an enthusiastic collector and botanist. Lygodesmia entrada Welsh & Goodrich, sp. nov. Ab Lygodesmia grandiflora Nutt. in caul- ibus rigidioribus ramosissimis foliis brevior- ibus et paucioribus et radiis albis differt. Herbae perennes caudice subterraneo ramosissimo omnino usque ad 45 cm altae; folia integra linearia vel acicularia 5-30 mm longa; pedunculi potius numerosi bracteati elongati 12-20 cm longi in capitulum termi- nans; bracteae involucrorum hyalinae exte- riores 5-10 mm longae fimbrillatae interiores circa sex 16-18 mm longae apex puberulus; radii albi circa 3 cm longi; pappus barbel- latus sordidus setae 10-15 mm longae; ach- enia costata glabra. Holotype: Utah, Grand Co., T24S, R19E, Sec 25, Tusher Canyon, ca. 15 mi. due WNW of Moab, 4800 feet elev., Entrada Sandstone Formation, juniper community, S. L. and S. L. Welsh 16725, 3 June 1978 (BRY, four iso- types to be distributed). This white-flowered material has been identified by A. S. Tomb (pers. comm.) as a probable triploid assignable to Lygodesmia arizonica Tomb. The triploid hypothesis can- not herein be questioned because of lack of knowledge concerning the cytological nature of the plants in question. However, despite the ultimate disposition of these plants fol- lowing future determinations of chromosome numbers, the strikingly different morpholo- gical features dictate taxonomic recognition, if for no reason other than the fact that the plants are so different from other plants of Lygodesmia in Utah. A second collection at BRY, here assigned to L. entrada, is J. S. Allen 132, from north of Courthouse Wash Ridge in Arches National Park. Dried flower remnants appear to be pink, but the tall nidularius habit and defi- nitely ligneous stems and branches are appar- ent. Lygodesmia arizonica Tomb is a low her- baceous plant usually of more southern distri- bution in Utah. Even in late anthesis the stems are herbaceous and lack the character- istic bird's nest appearance of L. entrada. Lygodesmia entrada differs from L. grandi- flora in ways similar to those discussed for L. arizonica. Pediocactus despainii Welsh & Goodrich, sp. nov. Ab Pediocacto hradyi L. Benson differt in spinis paucioribus brevioribus gracilioribus et floribus colorum. Plantae carnosae hemisphericae depressae 3-6 cm in diametrum 4-8 cm longae; tuber- culi ovata numerosa ordinata in serialia cir- cularia vel spiralia; areolae spinis 8-14; spin- ae 2-5 mm longae serialia stellatim; pilis coactis instructis interdum; flores 2.5-4 cm diametrum fragrantes ad apicem gerenti; se- pala numerosa; petala numerosa albida suf- fusa rosea et flava; stamina numerosa lutea 84 Great Basin Naturalist Vol. 40, No. 1 Fig. 4. Lygodesmia entrada: A, habit of plant; B, close-up of achene. March 1980 Welsh, Goodrich: Plant Novelties 85 cm B /JHiw,y Schedl), Ozopanon Hagedorn {Dnjocoetiops Schedl), Scolijtogenes Eichhoff ( = Cnjpliahmiorplius Schauhiss). Stepluinopodius Schedl {^Crijphdloniinictes Browne), and Xijlechintis Chapuis (^Sijuaina.'iinulus Nimberg). Genera new to science and their tvpe-species include: Anaxylchont.'i {Tomicits tnin- ccittis Erichson), Apoxi/lchorus (Xt/lcborus mancus Blandford), Crifphulogenes {Cri/pluilo^enes euphorbiae Wood), Er- nochidius {Cn/phahis corpulcntu.s Sampson), lladrodcniius (Xijlebonis globus Blandford), Leptoxt/lebonis (Phloeo- trootis sordicauda Motschulsky), Micropenis (Xijlebonis theae Eggers), Taphrodasus [Xijlebonis penorthi/liis Schedl), and Taiirodemiis (Xijlebonis sharpi Blandford). The new name Hijliirdn'ctoniis corticiiiiis is presented to replace H. araiicariae (Schedl 1972). Dnjocoetes coffeae Eggers is transferred to Eiilepiops. The following genera are treated in a revised context: Cn/togeniiis. Dnjocoetes, Eiilepiops, Enioporiciis, Enwponis, Xijlebonis, and Xylechinus. Cnjphalo- genes euphorbiae and C. exiguus (Sri Lanka) are named as new to science. In a review of the genera of Scolytidae in the world fauna, several problems that relate to synonymy were encountered. The new svnonvmv listed in the above abstract is re- ported here in order that names might be used in their new context before the generic revision is completed. In addition, several genera are treated in a sense somewhat dif- ferent from the traditional. The basis for these departures is established. The genera are treated alphabetically for convenience of reference. They include representatives from the subfamily Hylesininae, tribe Tomicini {Acmntiis, Hyhirdrectonus, Xylechinus) and from the subfamily Scolytinae, the tribes Ipini {Acanthotornicus), Dryocoetini {Cijrto- genitis, Eulepiops, Ozopemon), Xyleborini (Xyleborus), and Cryphalini {Cosmoderes, Er- noporictis, Ernoporus, Scolytogenes, Steph- anopodius). Nine new genera represent the tribes Xyleborini {Anoxyleborus, Apoxyle- borus, Hadrodcmius, Leptoxyleborus, Micro- perns, TapJirodasus, Taurodemus) and Cryphalini (Cryphalogenes, Ernocladius). Xylechinosomus Schedl is removed from syn- onymy with Pteleobius. The new name Hy- hirdrectonus corticinus is presented to re- place the junior homonym H. araucariae (Schedl 1972). The species Cryphalogenes eu- phorbiae and C. exiguus (Sri Lanka) are named as new to science. Acanthotornicus Blandford Acanthotornicus Blandford, 1894, Trans. Ent. Soc. Lon- don 1894:89 (Type-species: Acanthotornicus spin- osus Blandford, monobasic) Isophthorus Schedl, 1938, Archiv Naturgesch. 7(2): 173 (Type-species: Isophthorus quadrituberculatus Schedl, present designation). .Vpir synonymy In the original description of Isophthorus Schedl, two .species were definitely included and a third species was doubtful, but a type- species was never designated. Since then, Schedl has transferred all three species else- where. To anchor the generic name, Isoph- thorus quadritidierculatus Schedl is here des- ignated as the type-species of Isophthorus. Because this species and Myeloborus bico- nicus Schedl have been transferred to Acan- thotornicus and the unrelated, doubtfiil spe- cies, Pityophthorus heteae Hagedorn, has been transferred to Cryptocarenus, the fix- ation of a type-species requires that Isoph- fliorus be placed in .synonymy under Acan- thotornicus. 'Life Science Museum and Department of Zoology. Brigham Young University. Provo. Utah 84602. Scolytidae contribution number 69. 89 90 Great Basin Naturalist Vol. 40, No. 1 Acrantus Broun Hfliminis Broun, 1881, Manual of New Zealand Coleop- tera 2:720 (Tvpe-species: Homanis inunduliis Broim, monobasic). Preoccupied Acrantus Broun, 1882, Ann. Mag. Nat. Hist. (5)9:409. Replacement name Chaetophorus Fuchs, 1912, Morphologische studien iiber Borkenkafer, II. die europiiischen Hyiesinen, p. 46 (Tvpe-species: Hylesiniis vestitus Mulsant 6c Re\', monobasic). Preoccupied Clwetoptelius Fuchs, 191.3, in Reitter, Wiener Eut. Zeit. .32(Beiheft):4.3. {Replacement name). Xeic synon- !/'"!/ The names Acrantus Broun and Chaetop- telitis Fuchs have been treated as synonyms of Pteleobius Bedel (Schedl 1963:262) and Pseudohijlesinus Swaine (Schedl 1966:75), re- spectively. However, in a review of the char- acters of the type-species of these genera, it was demonstrated (Wood 1978) that Pte- leobius must be placed in the tribe Hylesinini and that Chaetopteliiis and Pseudohijlesinus belong in the tribe Tomicini. For that study, Schedl's (1963:262) placement of Acrantus was not challenged. In a subsequent review of the genera of Tomicini, di.ssection demonstrated that Ho- marus mundulus Broun, type-species of Acr- antus, clearly belongs to the Tomicini and is quite unrelated to Pteleobius. Furthermore, Pseudohijlesinus totally lacks pronotal aspe- rities, it has three distinct sutures on the an- tennal club, and the male frons is not noticea- bly impres.sed. Acrantus, Chaetopteliiis, and Xijlechinosomus all have numerous pronotal asperities, two or four poorly marked sutures on the antennal club, and the male frons strongly impressed and, thus, form a group quite distinct from Pseudohijlesinus. Xijle- chinosomus, which Schedl (1966:75) also placed in synonymy with Pteleobius, has the antennal club less elongate, less strongly compressed, and (apparently) with four ob- •scure sutures and the frontal rectangle much more elongate. Acrantus and Chaetopteliiis have the antennal club more elongate, strongly flattened, and marked by two su- tures and the frontal rectangle comparatively broad. Biological differences also support the continued recognition of Xijlechino.wmus. However, I can find no characters that sepa- rate Acrantus and Chaetoptelius. For this rea- son, Chaetoptelius is placed in synonvmv un- der Acrantus, as indicated above. Acrantus includes mundulus and vestitus, cited above, and most if not all of the species from New Zealand, Australia, New Guinea, and neighboring areas placed by Schedl in Leperisinus and Xijlechinus. Anaxyleborus, n. gen. This genus is distinguished from Eii- wallacea Hopkins and allied genera by the truncate, concave elytral declivity which has a complete, .sharply elevated, circum- declivital costa from base to apex. The discal interstrial punctures are uniseriate; in the su- perficially similar Apoxijleborus they are con- fused. Description.— Antennal club with one su- ture visible on posterior face, anterior face with segment 1 corneous, 2 conspicuous, sometimes rather large. Procoxae contiguous. Protibia armed by more than 11 socketed teeth. Declivity and discal punctures on in- terstriae as described in above diagnosis. Type-species: Tomicus truncatus Erichson. Species assigned previously to the Xijle- borus truncatus group belong here. Apoxyleborus, n. gen. This genus is distinguished from Tauro- demus by the presence of only four to seven socketed teeth on the protibia, by the obliquely truncate elytral declivity, with an abrupt (not acute) circumdeclivital costa, and the face flat to weakly concave. It is distin- gui-shed from the superficially similar Ana- xyleborus by the rather widely separated pro- coxae, by the strongly confused interstrial punctures on the disc, and by the less dis- tinctly concave elytral declivity. Description.— Body stouter than 1.9 times as long as wide. Antennal club with .segment 1 corneous, with no sutures evident on poste- rior face, apical margin of segment 1 on ante- rior face acutely elevated into a continuous co-sta forming a complete circle. Procoxae moderately separated. Protibia armed by four to seven socketed teeth. Elytral disc with in- terstrial punctures strongly confused, declivi- ty as described in above diagnosis. Tvpe-species: Xijleborus mancus Bland- ford. March 1980 Wood: American Bark Beetles 91 Species assigned previously to the Xyle- bonis mancus group belong here. Cosmoderes Eichhoff Cosmoderes Eichhoff, 1878, preprint of Mem. Soc. Roy. Sci. Liege (2)8:49.5 (Type-species: Cosmoderes moiiilUcollis Eichhoff, monobasic) Erioschidias Schedl, 1938, Trans. Roy. Soc. S. Austraha 62:42 (Type-species: Crijphalus sctistriattis L^a, subsequent designation by Wood, 1960, Insects of Micronesia 18(1):21). New sijnoiuit)nj The Beeson Collection at the Forest Re- search Institute, Dehra Dim, India, contains series of three species that were placed by Beeson in Cosmoderes. One, from Samsingh, Kalimpong, Bengal, is labeled monillicoUis Eichhoff; the other two bear manuscript names not yet validated. Beeson 's private notes, of which two volumes treating Scoly- tidae are in my possession, contain no in- dication under this name that he saw the type of moniUicoUis. However, elsewhere in his notes there are several indications that he saw the Eichhoff Collection at Hamburg be- fore it was destroyed during World War II. Blandford also saw the Eichhoff Collection, but there is some doubt (Blandford 1894:86) that he actually examined the type of mon- illicoUis. Both the Beeson and Blandford specimens are congeneric with Erioschidias Schedl. Bee- son's .specimens of monillicoUis match the dis- tinctive characters of Eichhoff's description in every detail. It is, therefore, propo.sed that Erioschidias be placed in synonymy under Eichhoff's name, as indicated above. Cryplialogenes, n. gen. This genus is distinguished from the closely allied Scolytogenes Eichhoff by the 3-seg- mented antennal funicle, by the antennal club with sutures 1 and 2 weakly procurved, marked by .setae, and 1 grooved and partly .septate, and by tlie horizontal venter of the abdomen. Description.— Frons convex, not sexually dimorphic. Eye elongate-oval, entire. Anten- nal scape elongate, simple; fimicle 3-seg- mented; club oval, a slight constriction and groove at suture 1, sutures 1 and 2 moder- ately procurved, 1 partly septate at lea.st on lateral half. Pronotum with basal and lateral margins marked by a fine, raised line; ante- rior slope asperate, anterior margin armed by low, poorly formed serrations. Elytral punc- tures largely replaced by rows of rounded strial and interstrial granules; vestiture of rows of -Strial hair and interstrial scales. Pro- tibia armed by four socketed teeth. Venter of abdomen horizontal. Sexes subequal in size. Type-species: Cryphalogenes euphorbiae Wood. Cryphalogenes euphorbiae, n. sp. This species is distinguished from exiguus Wood by the larger size, by the absence of reticulation of the pronotum (except in ex- treme lateral areas of some specimens), and by the comparatively .smaller pronotal and elytral granules. Male.— Length 1.2-1.4 mm, 2..3 times as long as wide; color dark brown. Frons broadly convex, very feebly so on longitudinal axis; surface largely reticulate, minute punctures moderately, uniformly abundant, most of them feebly granulate. An- tennal club slightly longer than scape. Pronotum 1.0 times as long as wide; sides almost straight and parallel on basal third, rather broadly rounded in front; anterior margin armed by about four to six irregular, poorly formed serrations; summit near middle; anterior .slope rather coarsely aspe- rate, punctured between asperities; posterior areas smooth, shining (except some reti- culation present in extreme lateral areas of some specimens), with close, moderately large, rounded granules, anterior slope of each granule bearing a puncture (punctures usually visible only when light source ceph- alad). Vestiture of fine, .short, semirecumbent hair. Elytra 1.3 times as long as wide, 1.4 times as long as pronotum; .sides almost straight and parallel on basal two-thirds, rather broadly rounded behind; striae not impressed, each puncture largely replaced by a large roimded granule as wide as striae, puncture confined to posterior slope of each granule; interstriae as wide as striae, .smooth, shining, punctures largely replaced by rounded granules of same size and shape as those of striae. Declivity steep, convex; sculpture as on disc. Vestiture of rows of fine, short, strial hair and rows of 92 Great Basin Naturalist Vol. 40, No. 1 erect interstrial scales, each scale slightly shorter than distance between rows, spaced within a row by length of scale, each four to six times as long as wide on disc, two to four times on declivity. Female — Similar to male in all respects. Type locality- Thirty km southeast of Puttalam, Sri Lanka (Ceylon). Type material— The male holotype, fe- male allotype, and 34 paratypes were taken at the type locality on 18-VI-1975, No. 214, from Euphorbia antiquonim, by me; 28 para- types bear the same data except they were taken 24 km SE Puttalam. Additional para- tvpes were taken in Sri Lanka as follows: 21 at 5 km SE Naula, 14-VM975; 14 at 48 km N Naula, 14-VI-1975; 2 at 32 km N Haba- rana, 12-VI-1975; 1 at 8 km SW Kurunegala, 13-VI-1975; and 1 at 11 km W Kikirawa, 19- VL1975; all from the same host and collec- tor. The holotype, allotype, and half the para- types are in the U.S. National Museum. The remaining paratypes are in my collection. Cryphalogenes exiguus, n. sp. This species is distinguished from eu- phorhiae Wood by the smaller size, by the strongly reticulate pronotum, and by the comparatively larger pronotal and elytral granules. Male.— Length 0.8-1.0 mm, 2.2 times as long as wide; color dark brown. Frons as in euphorbiae except more strong- ly convex, granules smaller, less conspicuous. Antennal club with septum in suture 1 less apparent. Pronotum as in euphorbiae except reti- culate, shining, granules in posterior areas proportionately slightly larger. Elytra as in euphorbiae except interstrial scales averaging more slender, those on de- clivity not less than four times as long as wide. Female.— Similar to male in all respects. Type locality.— Thirty km southeast of Puttalam, Sri Lanka (Ceylon). Type material- The male holotype, fe- male allotype, and 43 paratypes were taken at the type locality on 18-VI-1975, No. 214, from Euphorbia antiquonim, by me. Addi- tional paratypes were taken in Sri Lanka dur- ing 1975 from the same host, by me, as fol- lows: 13 at 24 km SE Puttalam, 17-VI; 5 at 5 km SE Naula, 14-VI; 4 at 48 km N Naula, 14- VI. The specimens were taken in indepen- dent galleries in the same stems with eu- pliorbiae. The holotype, allotype, and half the para- types are in the U.S. National Museum. The remaining paratypes are in my collection. Cyrtogenius Strohmeyer Ki/rtogcniits Strohmeyer, 1910, Ent. Blatt. 6:127 (Type- species: Kijrtogeniiis bicolor Strohmeyer, mon- obasic) Ciirtooenius Strohmeyer, 1911, Ent. Bliitt. 7:116. Valid emendation Carposinus Hopkins, 1915, U.S. Dept. Agric. Rept. 99:9, 47 (Tvpe-species: Carposiniis pini Hopkins = Lepicenis nitidus Hagedorn, original designa- tion) Orosiotes Niisima, 1917, Coll. Essays Y. Nawa, p. 1 (Type-species: Orosiotes kumatoensis Niisima, monobasic) Metahylastes Eggers, 1922, Ent. Bliitt. 18:165 (Type-spe- cies: Metahijlastes africanus Eggers, monobasic) Pelicenis Eggers, 1923, Zool. Meded. Roy. Mus. Nat. Hist. Leyden 7:216 (Type-species: Lepicerus ni- tidus Hagedorn, original designation) Taphrohorus Nunberg, 1961, Ann. Mag. Nat. Hist. (13)3:617 (Type-species: Taphrohorus vaticae Nunberg, original designation) Much confusion exists in the literature rel- ative to the identity of this tropical genus. It is characterized by five socketed teeth on the lateral margin of the protibia, by the posteri- or face of the antennal club with only one su- ture, sutures on anterior face procurved, with the pubescence extending to the base, by the narrowly separated procoxae, and by the slightly elevated or armed posterolateral margin of the elytral declivity. Dryocoetes differs from it by the recurved suture 1 on the antennal club, the pubescence never ex- tending to the base, by the contiguous pro- coxae, and by the rounded, unarmed, post- erolateral margins of the elytral declivity. Both genera are phloeophagous and have heterosanguineous polygenous breeding habits in which the male is subequal in size to the female and assists in the formation of new parental galleries. Both genera have been confused with Eulepiops (see below). March 1980 Wood: American Bark Beetles 93 Ernocladiiis, n. gen. This genus is distinguished from the closely allied Ernoporus Thomson by the 3-seg- mented antennal funicle, by the uniseriate in- terstrial setae (interstrial ground vestiture al- ways absent on disc, a few setae sometimes present on declivity), and by the weakly pro- curved (often obscure) sutures of the antennal club. Description.— Frons dimorphic, moder- atelv impressed in male, convex in female. Eye elongate-oval, entire. x\ntennal scape elongate; funicle 3-segmented; club rather large, sutures weakly to moderately pro- curved, aseptate, marked by rows of setae, grooves present or not. Pronotum with basal margin marked by a fine, raised line, lateral margin rounded, without a raised line; aspe- rities in concentric rows, their bases often contiguous or even reduced to a continuous costa. Elytra with basal margins rounded, strial punctures in rows, sculpture conserva- tive; vestiture of rows of minute strial hair and rows of erect interstrial scales, interstrial ground vestiture absent on disc, a few short setae in ground cover sometimes present on declivity. Type-species: Cn/pJialus corptilentus Sampson. Several additional species will be trans- ferred to this genus as soon as their types can be examined. Schedl (1940:590) assigned Cry- phuhis corpulentus to Margadillius, apparent- ly without appreciating the significance of tlie emarginate eye or the fine, raised line on the lateral margin of the pronotum of Marga- dillius species. Ernoporicus Berger Ernoporicus Berger, 1917, Rev. Russc d'Ent. 16:242 (Tvpe-species: Ernoporicus spessivtzevi Berger, monobasic) Eocnjphalm Kurentzov, 1941, Acad. Sci. USSR, Koma- rov Sta. Sci., Orient, p. 230 (Type-species: Eocnj- pluilus seinenovi Kurentzov, monobasic) Ernopocerus Balachowsky, 1949, Fauna de France 5():211 (Type-species: Ernoporus caitcasicus Lind- emann, subsequent designation by Wood, 19.54, Univ. Kansas Sci. Bull. 36:986). Xinc sijnonyiny The complex of genera allied to Ernoporus Thom.son have been poorly known and erro- neously classified, largely due to the paucity of material for study. Following an exam- ination of the type-species of Ernoporicus, Eocryphalus, and Ernopocerus, it was con- cluded that these three congeneric species have the ba.sal and lateral margins of the pro- nottun rounded (without a fine, raised line), the procoxae narrowly separated, the eye short and entire, the antennal funicle 4-seg- mented, and the antennal club with the su- tures procurved and marked only by setae or obsolete (never septate). Ernoporus kanawhae Hopkins of North American and E. fagi (Fab- ricius) and a few species from A.sia also be- long here. The genus Ernoporus is quite dif- ferent, as indicated below. Ernoporus Thomson Ernoporus Thomson, 1859, Skandinaviens Coleoptera Svnoptiskt Bearbitade, p. 147 (Type-species: Bos- trichus tilkic Panzer, original designation) Cryphalops Reitter, 1889, Wiener Ent. Zeit. 8:94 (Type- species; Cn/phalus lederi Reitter =Bostrichus tiliae Panzer, monobasic) Sfcphanorhopalus Hopkins, 1915, U.S. Dept. .\gric. Rept. 99:35 (Type-species: Stephanorhopalus nulodori Hopkins, amended to melodori by Schedl, 1966, Ent. Abh. Mus. Dresden .35:19, original designation) Euptilius Schedl, 1940, Mitt. .Miincher Ent. Ges. .30:.590 (Type-species: Ernoporus concentralis Eggers, original designation). New synonymy Ernoporus Thomson has the basal and lat- eral margins of the pronotum marked by a fine, rai.sed line, the procoxae contiguous, most pronotal asperities arranged in con- centric rows, the antennal funicle 4-seg- mented, the antennal club .sutures strongly procurved to obsolete, and the elytral vesti- ture abundant and conhused. Most of the .spe- cies occur in tropical .\sia except for tiliae, the type-species. In a review of the genera belonging to this complex, it was found that Ernoporus concentralis Eggers falls well with- in the range of variability for Ernoporus. Be- cause concentralis is the type-species of Eu- ptUius Schedl, it is, therefore, necessary to place Schedl's genus in synonymy as in- dicated above. The structure of the pronotum indicates that this genus is quite di.stinct from Ernoporicus, as noted above. 94 Great Basin Naturalist Vol. 40, No. 1 Eulepiops Schedl Eulepiops Schedl, 19.39, J. Fed. Malay St. Mus. 18{,3):344 (Tvpe-species: EiiUpiops glaber Schedl, mon- oba.sic) This genus has been confused with Cyrto- genius Strohnieyer and Dnjocoetes Eichhoff. It differs by the protibia bearing only three socketed teeth on the lateral margin, by the posterior face of the antennal club with two sutures indicated, the anterior face with su- ture 1 straight to recurved and always on the basal fourth. The male is either unknown or dwarfed, deformed, flightless, and does not participate in the formation of new parental galleries. Reproduction is either by con- sanguineous polygyny or possibly by some form of parthenogenesis. The habit is myelo- phagy for the only species observed. Dnjo- coetes coffeae Eggers and its allies belong to this genus. Hadrodemius, n. gen. This genus is distinguished from Eccoptop- tems Eichhoff by the tibiae being of normal size and all bearing socketed teeth, by the normal metatarsi (not compressed), by the declivity being restricted to the posterior half of the elytra, and by the convex to moder- ately impressed, unarmed elytral declivity. Description.— Body very stout, less than 1.8 times as long as wide, usually black. An- tennal club with posterior face unmarked by sutures, on anterior face costa marking apical margin of corneous area usually forming a complete ring. Scutellum visible only on an- terior declivous slope of elytral margins. De- clivity and tibiae as described in above diag- nosis. Type-species: Xylebarus globus Blandford. Members of the Xyleborus globus species group should be referred here. Hylurdrectonus Schedl Hylurdrectomis Schedl, 1938, Trans. Roy. Soc. S. .'Vustra- lia 62;4() (Type-species: Hylurdrectonus piniarius Schedl, monobasic) Xylogopinus Schedl, 1972, Papua New Guinea .\5;ric. J. 23:64 (Type-species: Xylogopinus araucariae Schedl = Hylurdrectonus corticinus Wood, mon- obasic). New synonymy A review of long series of Hylurdrectonus piniarius Schedl, H. araucariae Schedl (1964a:213), and Xylogopinus araucariae Schedl indicates the absence of characters that will separate these two genera. Con- sequently, it is necessary to place Xylogo- pinus in synonymy under the older name as indicated above. This act creates homonymy as indicated below. Hylurdrectonus corticinus, new name Xillogopinus araucariae Schedl, 1972, Papua New Guin- ea Agric. J. 23:64 (Bulolo, Morobe Distr., New Guinea) A long series of this species was collected near Bulolo and compared to the holotype and paratypes in the Forest Research Labora- tory collection at Bulolo. As indicated above, this species must be transferred to Hylurdrec- tonus. The transfer makes this species a jun- ior homonym of H. araucariae Schedl, 1964. Tlie new name Hylurdrectonus corticinus is proposed to replace H. araucariae (Schedl 1972). Leptoxyleborus, n. gen. This genus is distinguished from the allied Theobortis Hopkins and Coptoborus Hopkins by the declivity commencing anterior to the middle of the elytra, its lower half broadly impressed and either flat or shallowly con- cave. If the discal interstrial punctures are uniseriate, then the declivital surface is den- sely covered by small, confused scales; if the declivital setae are hairlike, then the discal interstrial punctures are confused. Description.— Antennal club with two su- tures indicated on posterior face, anterior face with segment 2 comparatively large, sclerotized, convex, apical portion beyond segment 2 flat to concave. Protibiae and metatibiae each armed by six or seven sock- eted teeth. Anterior coxae contiguous. Scutel- lum visible. Declivity as described in above diagnosis. Type-species: Phloeotrogus sordicauda Motschulsky. Other species placed previously in the Xyleborus sordicauda group also belong here. Micropenis, n. gen. This genus is distinguished from Taphro- dastis Wood by the convex elvtral declivity tliat lacks a circumdeclivital costa, by the ab- March 1980 Wood: American Bark Beetles 95 sence of declivital scales, and by the strial pimctures that are arranged in definite rows. Description.— Body slender, at least two times as long as wide, color yellowish or red- dish brown. Posterior face of antennal club with at least one suture visible, apical margin of corneous area never costate. Scutellum not visible. Strial punctures usually in rows. De- clivity convex, variously sculptured, without a costa. Type-species: Xyleborus theae Eggers. Members of the Xijleborus theae species group should be referred here. The name Mi- cropenis was originally coined by F. G. Browne for this group for use in an unpub- lished manuscript a decade ago. Ozopeinon Hagedorn Ozopcmon Hagedorn, 1908, Deutsche Ent. Zeitschr. 1908:382 (Type-species: Ozopemon regius Hage- dorn, monobasic) Dryocoetiops Schedl, 1957, Ann. Mus. Roy. Congo Beige, Tervairen, Ser. 8, Sci. Zool. .56:13 (Type-species; Ozopemon laevis Strohmeyer, monobasic). New sijnonymy A series of Ozopemon laevis Strohmeyer was compared to Eggers's series of this spe- cies and to representatives of eight species of Ozopemon. Although the sculpturing of the pronotum is somewhat unique for the genus, this species appears to fall well within the limits of variability for Ozopemon. For this reason, Dryocoetiops is placed in synonymy as indicated above. Scolytogenes Eichhoff Scolytogenes Eichhoff. 1878, preprint of Mem Soc. Roy. Sci. Liege (2)8:475, 497 (Type-species: Scolyto- genes danvinii Eichhoff. monobasic) Cryphcilomorphus Schaufnss, 1890 (1891), Tijdschr. Ent. 34:12 (preprint 1890 by Martinus N'ijhoff, Hagg) (Type-species: Cryphahnurpbiis conimiinif: Schau- fnss, monobasic). \eiv synonymy Eggers (1929:53) examined the type-speci- mens of the type-species of Scolytogenes and Lepicerus and compared them to the type- specimens of Negritus major Eggers and A'. minor Eggers. He concluded that N. major and N. minor were congeneric with Scoly- todes danvinii Eichhoff. The holotype of S. darwinii apparently was lost when the Stettin Museum was damaged during World War II. In the absence of that type, direct com- parisons are not now possible; however, if it is assumed that Eggers was correct in his ob- servations, then N. major and S. danvinii are congeneric. My examination of the lectotype of A', major and .syntypes of N. ater (type-spe- cies of Negritus) demonstrates that these spe- cies are congeneric; consequently, Negritus must be a jimior synonym of Scolytogenes. Because N. major and A\ ater are also consid- ered congeneric with Cryplmlomorphus com- munis Schaufuss (type-species of Cryphalo- morplius) (Schedl 1957:152), it must also be concluded that Cryphalomorphus is a junior synonym of Scolytogenes. (Note added in press: The list of types in the Schedl Collection at the Vienna Museum, just received, includes the type of S. darwinii. It will be examined as soon as arrangements can be completed.) Stephanopodius Schedl Stephanopodius Schedl, 1941, Rev. Zool. Bot. Afr. .34:.396 (Type-species: Stephanoderes dispar Eggers, sub- sequent designation by Schedl, 1961, Rev. Ent. Mozambique 4:6.33) Cryplwlomimus Browne, 1962, West .\frican Timber Bo- rer Research Unit Rept. 5:75 (Type-species: Hy- pocryphahts ghanaensis Schedl. original designa- tion) Cryphcdmomimetes Browne, 1963, Ann. Mag. Nat. Hist. (1.3)6:242 (Replacement name). Xcii synonymy When Schedl named Htjpocryphalus gha- naensis and then later (Schedl 1964b:305) transferred this species from Cryphalomi- metes back to Hypocnjphalus, he overlooked some very important characters. In this spe- cies and in Stephanopodius, the basal margin of the pronotum bears a fine, raised line, but the lateral margin is rounded and lacks the fine, raised line of Hypocryphalus. In addi- tion, the antennal club is quite different from Hypocryphalus. The species ghanaensis is congeneric with Stephanopodius dispar (Egg- ers) and, as indicated above, should be trans- ferred to that genus. Cryphalomimetes is, therefore, a .synonym of Stephanopodius Schedl and not of Hypocryphalus Hopkins. Taphrodasus, n. gen. This genus is distinguished from Micro- penis Wood by the confused interstrial pimc- tures, by the presence of scales on the elytral 96 Great Basin Naturalist Vol. 40, No. 1 declivity, and by the strongly concave decliv- ity that commences on the basal half of the elytra and is marked on its lateral margins in such a way as to form a blunt, elongate, cir- cumdeclivital costa. Description.— Body slender, at least 2.0 times as long as wide, color reddish brown. Posterior face of antennal club with one su- ture visible, apical margin of corneous area never costate. Scutellum not visible. Strial punctures on disc confused. Declivity as de- scribed in above diagnosis. Type-species: Xylehorus percorthylus Schedl. Taurodemus, n. gen. This genus is distinguished from Xijleborus Eichhoff by the moderately to rather widely separated procoxae, by the rather stout body, by the presence of 9 to 12 socketed protibial teeth, and by the distinctive sculpture of the sulcate elytral declivity. Description.— Body stout, less than 1.9 times as long as wide. Antennal club with segment 1 corneous, without any sutures evi- dent on posterior face, apical margin of seg- ment 1 on anterior face acutely elevated into a continuous costa forming a complete circle. Procoxae moderately to rather widely sepa- rated. Protibia armed by 9 to 12 socketed teeth. Elytral declivity moderately to very strongly sulcate on at least basal half, lateral margins armed by at least one major spine and several smaller tubercles. Type-species: Xijleborus sharpi Blandford. The following species are transferred from Xylehorus to Taurodemus: bicornutus Wood, ebenus Wood, (Bostrichus) flavipes Fabricius, godmani Blandford, pandulus Wood, {Am- phic(iranus) perebeae Ferrari, salvini Bland- ford, sanguinicollis Blandford, sharpi Bland- ford, splendidus Schaufuss, (Bostrichus) varians Fabricius, and varus Wood. Xyleborus Eichhoff Xijleborus Eichhoff, 1864, Berliner Ent. Zeitschr. 8:37 (Type-species: Bostrichus monographus Fabricius, subsequent designation bv Hopkins. 1914, Proc. U.S. Nat. Mus. 48:131) The genus Xyleborus Eichhoff, as inter- preted in recent years by Schedl, contains more than 1400 nominate species, that is, vir- tually all the species in the tribe Xyleborini. However, the diversity of characters and habits within this group suggests the exist- ence of several distinct clusters of species and species groups that could and should be char- acterized as genera. The difficulty in frag- menting the group piecemeal, as has been at- tempted by some workers, is that when one group is removed and elevated to generic rank, the remainder becomes unclassifiable on a logical, phylogenetic basis. In order to remedy this situation, a classification is being composed, based on such constant features as the location of mycetangia, structure of the antennal club, form and armature of the tibiae, characters of the scutellum, and many other features. A deliberate effort is being made to avoid use of adaptive characters such as the surface sculpturing of the pro- notum and elytra. Tentatively, 27 groups are being given ge- neric status within the Xyleborini. Those de- scribed previously include: Ambrosiodmus { = Browneia, Phloeotrogus), Arixyleborus {=Xyleboricus), Cnestus { = Tosaxyleborus), Coptoborus { = Streptocranus), Cryptoxyle- borus, Dryocoetoides, Eccoptoptcrus { = Eu- rydactylus, Platydactylus), Euwallacea, Ka- lantanius, Mesoscolytus, Notoxyleborus, Premnobius { = Prernnophilus), Pseudoxyle- borus, Sampsonius, Schedlia, Theoborus, Webbia { = Prowebbia, Pseudowebbia, Xely- borus), Xyleborinus, Xyleborus {=Anaeritus, Anisandrus, Boroxylon, Bufonus, Coptodryas, Cyclorhipidion, Heteroborips, Phloeotrogus, Progenius, Terminalinus, Xyleborips), and Xylosandrus. The above is mentioned to es- tablish a context into which the seven genera in this tribe, described in this article, can fit. The seven include: Anaxyleborus, Apoxylc- borus, Hadrodemius, Leptoxyleborus, Micro- penis, Taphrodasus, and Taurodemus. Xylechinus Chapuis Xylechinus Chapuis, 1869, Synopsis des Scolvtides, p. 36 (Type-species: Dcndroctontis pilosiis Knoch) Squamasinuhis Nunberg, 1964, Ann. Hist. -Nat. Mus. Nat. Hungarici, Pars Zool. 56:431 (Type-species: S(iu(im(isinulus chiliensis Nunberg, original desig- nation). New synonymy When the holotype of Squamasinuhis chi- liensis Nunberg and several allied species March 1980 Wood: American Bark Beetles 97 from South American were examined, no characters could be found that distinguish this genus from Xylechinus Chapuis. As nearly as can be determined at the present time, the genus Xylechinus consists of 14 Central and South American, 2 North Ameri- can, 5 Asian, and 1 European species. Schedl has referred four New Guinean and Austra- lian species to this genus, all of which appar- ently should be transferred to Acrantus. One African species placed in Xijlechinus by Schedl apparently belongs elsewhere. Literature Cited Blandford, W. F. H. 1894. The Rhynochophorous Co- leoptera of Japan. Part III. Scolytidae. Trans. Ent. Soc. London, pp. 53-14L Eggers, H. 1929. Zvir Synonymic der Borkeiikafer (Ipidae, Col.). Wiener Ent. Zeit. 46:41-55. Sc;hedl, K. E. 1940. Zur Einteilung und Synonvmie der Cryphalinae (Col. Scoivt.). Mitt. Miichner Ent. Ges. .30:583-591. 1957. Bark- and timber-beetles from South .Af- rica. Ann. Mag. Nat. Hi.st. (12)10:149-159. 1963. Zur Synonymie der Borkenkafer IX. Ent. Abh. Mus. Tierk. Dresden 28:257-268. 1964a. On some Coleoptera of economic impor- tance from New Guinea and Australia. Pacific In- sects 6:211-214. 1964b. Zur Synonymie der Borkenkafer XV. Rei- chenbachia 3:303-317. 1966. Neotropi.sche Scolytoidea VIII. Ent. Arb. Mus. Frey 17:74-128. Wood, S. L. 1978. A reclassification of the subfamihes and tribes of Scolytidae (Coleoptera). Ann. Soc. Ent. France (N.S.).' 14:95-122. THE BACTERIUM THIOPLOCA INGRICA ON WET WALLS IN ZION NATIONAL PARK, UTAH Samuel R. Riishforth', Sheril D. Burton", Jeffrey R. Johansen', and Judith A. Grimes' .\bstract.— Hanging gardens and wet wall habitats have been studied for the past several years in many arid regions of the Intermountain West. One unusual large wet wall habitat in Zion National Park was found to be cov- ered with a mucilaginous red-colored growth of the filamentous gliding bacterium Thioploca ingrica Visloukh. This is the only habitat we have examined where the predominant matrix-forming organism was a bacterium rather than an alga. Hanging gardens are unusual habitats found in several areas around the world, par- ticularly in western North America. Such habitats form when water percolates vertical- ly through permeable rock facies (generally sandstone) imtil it reaches an impervious lay- er. The water then moves laterally imtil it exits the rock formation often on a vertical wall or cliff. Such exit springs often occur along fairly long horizontal lines to form lin- ear seep walls or wet walls. Such habitats rapidly become colonized by a variety of mesic plant species, some of which are en- demic to such gardens. Seep walls become weakened through time, particularly in the massive mesozoic sandstone formations typi- cal of areas of southern Utah and northern Arizona. When this occurs, large portions of the wall slough away to form grottos that are .shaded from the sim and are cooler and more humid than .surrounding areas. Such hanging garden habitats have been imder study for several years (Rushforth et al. 1976, Clark and Rushforth, in press, Welsh and Moore 1965a, 1965b). We have been particularly interested in the algal species that colonize hanging garden walls. The algal floras of such habitats are variable depending upon several factors, primarily the amount of water available. The moister walls are almost always covered with heavy growth of green or blue green algae that secrete copious mucilage. Such algal mats are in turn colo- nized by dozens of other algal species, in- cluding blue green algae, green algae, eu- glenophytes, golden algae, and diatoms. Algal diversity is really quite high in many of these gardens, and the species are often imusual in morphology and/or distribution. We have found that wet wall faces that are exposed to the sun are often colonized by al- gae with dominant nonchlorophylous pig- ments. Thus, on certain walls, the green alga Palmella miniata Leiblein is common. This organism is often a deep red color due to hematochrome pigments, and in turn the gar- den walls colonized by these organisms are often a beautiful brick-red color. Likewise, Scytonema myochrous (Dillw.) C. A. Agardh and Scytoneina alatiim (Carm.) Borzi are prevalent on some walls. These organisms are large, prominently ensheathed blue green al- gae. The sheath and the cell wall of these al- gae become colored as the organisms mature to form yellow or yellow brown filaments which also color their walls of colonization. Recently, we have been studying algae and lichens of Zion National Park. One promi- nent wet wall in the park is Weeping Rock (Fig. 1), a famous tourist attraction. This wall is very large and is usually very moist. It is heavily colonized by many algal species. In several places on the wall, bright orange red patches and streaks are evident. We collected specimens from these areas expecting to find Palmella miniata. Upon returning to the lab- 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. ■Department of Microbiology, Brigham Young University, Provo, Utah 84602. 98 March 1980 RUSHFORTH ET AL.: HANGING GaRDENS BaCTERIUM 99 Fig. 1-4. 1. Habit of Tliioplora and algal communities of Weeping Rock, Zion National Park; 2, "Braid" of Thio- ploca im^rica filaments in a common sheath; .3, Thioplocd with Chamae.siphon species attached; 4, Low magnification photograph showing heavy epiphytic growth of Chamaesiphon species. 100 Great Basin Naturalist Vol. 40, No. 1 oratory, we examined the organisms using a Zeiss RA Research microscope with No- marski differential phase contrast accessories. We were surprised to find that 'the red color- ation was not due to Palmella but to a fila- mentous prokaryote. Careful study elimi- nated known blue green algae as the causal organism. Further examination showed the organism to be a bacterium in the family Beggiatoaceae Migula. These bacteria are gram-negative filaments that are motile by gliding motion. Three genera are presently placed in this family: Beggiatoa Trevisan, Vitreoscilh Pringsheim, and Thioploca Lau- terbom. In addition, the three genera Bactos- cilla Pringsheim, Flexoscilla Pringsheim, and Thiospirillopsis Uphof are possible members of the family (Leadbetter 1974). The organisms we collected from Weeping Rock in Zion National Park may be placed in the genus Thioploca, based upon the presence of more than one filament in the sheaths (Figs. 2-4). Furthermore, the filaments ranged between the width of 2 and 7 um, placing them in Thioploca ingrica Visloukh. The filaments often served as a substrate for epiphytic algae, particularly Cluimaesiphon species (Fig. 3). This observation of T. ingrica is interesting for several reasons. First, it represents the only example in the several gardens we have studied where the predominant mucilaginous, matrix-forming organism was a bacterium rather than an alga. Second, it represents the only reported occurrence of Thioploca from intermountain western North America that we are aware of. Third, Thioploca usually ex- hibits a greenish blue color. However, the specimens we have collected produce a bright orange red color on the wall itself and a paler orange color when examined beneath the microscope. The nature of this color is unknown, although Beggiatoaceae are report- ed to not form carotenoid pigments. And fourth, the presence of this organism on the moist walls is itself unusual because all the reports to date that we are aware of chron- icle Thioploca species as inhabiting the upper layers of bottom muds of freshwater and brackish habitats (Maier 1974). Specifically, such organisms have been collected from both oxidizing and reducing environments in such muds (Perfil'ev et al. 1965). Literature Cited Clark, R. L., and S. R. Rushforth. In review. A study of the algal flora of selected hanging gardens of Glen Canyon, Utah. Great Basin Naturalist Mem- oirs. Leadbetter, E. R. 1974. Beggiatoaceae Migula. Pages 112-115 in R. E. Buchanan, N. E. Gibbons, eds. Bergey's manual of determinative bacteriology, 8th ed. Williams and Wilkins, Baltimore. Maier, S. 1974. Thioploca Lauterborn. Pages 113-116 in R. E. Buchanan, N. E. Gibbons, eds. Bergey's manual of determinative bacteriology, 8th ed. Williams and Wilkins, Baltimore. Perfil'ev, B. V., D. R. Gabe, A. M. Galperina, V. A. Rabinovich, a. a. Sapotnitskii, E. E. Sherman, AND E. P. Troshanov. 1964. Applied capillary microscopy: The role of microorganisms in the formation of iron-manganese deposits. Izadatels- tov, Akad Nauk SSSR, Savarenskii Laboratory for Hydrogeological Problems. Moscow. In Russian (Transl. Consultants Bureau Enterprise Inc. 1965 New York). Rushforth, S. R., L. L. St. Clair, T. A. Leslie, K. H. Thorne, and D. S. Anderson. 1976. The algal flora of two hanging gardens in southeastern Utah. Nova Hedwigia 27:231-323. Welsh, S. L., and G. Moore. 1965a. Plants of Canvon- lands National Park. Part I. The Needles Region. Proc. Utah Acad. Sci. 42(1): 160. Welsh, S. L., and G. Moore. 1965li. Plants of Canyon- lands National Park. Part II. The Islands in the Sky Region. Proc. Utah Acad. Sci. 42(1):160-161. NOTICE TO CONTRIBUTORS Original manuscripts in English pertaining to the biological natural history of western North America and intended for publication in the Great Basin Naturalist should be directed to Brigham Young University, Stephen L. Wood, Editor, Great Basin Naturalist, Provo, Utah 84602. Those intended for the Great Basin Naturalist Memoirs should be similarly directed, but these manuscripts are not encumbered by a geographical restriction. Manuscripts. Two copies of manuscripts are required. They should be typewritten, double spaced throughout on one side of the paper, with margins of at least one inch on all sides. Use a recent issue of either journal as a format, and the Council of Biology Editors Style Manual, Third Edition (AIBS 1972) in preparing the manuscript. 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Cottam, A. M. Woodbury, H. H. Frost. $10. *»Jo. 2 Intermountain biogeography: a symposium. By K. T. Harper, J. L. Reveal et al. $15. ^lo. 3 The endangered species: a symposium. $6. TABLE OF CONTENTS Some aspects of succession in the spruce-fir forest zone of northern Utah. David J. Schimpf, Jan A. Henderson, and James A. MacMahon 1 Utah Flora: Malvaceae. Stanley L. Welsh 27 Utah Flora: Miscellaneous families. Stanley L. Welsh 38 The taxonomic status of the rosy boa Lichamira roseofusca (Serpentes: Boidae). John R. Ottley, Robert W. Murphy, and Geoffrey V. Smith 59 Hespewperla lio^uei, a new species of stonefly from California (Plecoptera: Perlidae). Richard W. Baumann and Bill P. Stark 63 Reproduction in three sympatric lizard species from west-central Utah. John B. Andre and James A. MacMahon 68 Haplopappus alpinus (Asteraceae): A new species from Nevada. Loran C. Anderson .. 73 Miscellaneous plant novelties from Alaska, Nevada, and Utah. Stanley L. Welsh and Sherel Goodrich 78 New genera and new generic synonvmv in Scolytidae (Coleoptera). Stephen L. Wood [ '.....'. '. 89 The bacterium Thioplova ingrica on wet walls in Zion National Park, Utah. Samuel R. Rushforth, Sheril D. Burton, Jeffrey R. Johansen, and Judith A. Grimes 98 HE GREAT BASIN NATURALIST ume 40 No. 2 June 30, 1980 Brigham Young University MUS. COMP. ZOOU, NOV GREAT BASIN NATURALIST Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Life Science Museum; Stanley L. Welsh, Botany; Clayton M. White, Zoology. Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agricul- tural Sciences; Ernest L. Olson, Director, Brigham Young University Press, University Editor. The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been published from one to four times a year since then by Brigham Young University, Provo, Utah. 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W'illianis' .\bstr.\ct.— (w/a hoiaxchiit.s is a dwarf species of cvprinid endemic to a thermal lake in southeastern Oregon. Despite a relatively depauperate fauna and flora in the lake, 24 food items were foimd in intestines of G. horaxobins. Ten of the 24 foods, including six insects, were of terrestrial origin. The relative importance of food items fliictuated seasonally. Diatoms, chironomid larvae, microcrustaceans, and dipteran adults were the primary foods during spring. In summer, diatoms decreased and terrestrial insects increased in importance. During autumn important foods were terrestrial insects, chironomid larvae, and diatoms. Diatoms and microcrustaceans increased in importance during winter. Chironomid larvae were of importance in winter, when the importance of terrestrial food items decreased .substantially. Similar food habits were observed between juveniles and adults, except that adults consumed more gastropods and diatoms and juveniles consumed more copepods and terrestrial insects. Gila howxobius feeds op- portunistically with individuals commonly containing mostly one food item. Fish typically feed by picking foods from soft bottom sediments or from rocks. However, fish will feed throughout the water column or on the surface if food is abundant there. Gila boraxohiiis feeds throughout the day, with a peak in feeding activity just after sunset. .\ daily ration of H.l percent boch' weight was calculated for the species during June. .\ comparison of food habits among G. boraxobius and populations of G. alvordensis during the summer shows that all are opportimistic in feed- ing, but that G. boraxobius relies more heavily on terrestrial foods. The Borax Lake chub, Gilo boraxobius, is a rare species of cvprinid fish endemic to a thermal lake in the Alvord Basin of south- eastern Oregon. The restricted habitat of G. boraxobius is threatened bv geothermal energy development in the Alvord. In recog- nition of this problem, the species is listed as "threatened" by the Endangered Species Committee of the American Fi.sheries Society (Deacon et al. 1979) and is currently on the protected list of the Oregon Department of Fish and Wildlife. Gila boraxobius was recently described and has been diagnosed as a dwarf relative of G. alvordensis (Williams and Bond, in press). Typical adult size of Gila boraxobius is 33-50 mm standard length (SL). Individuals longer than 55 mm SL are rare. Becau.se no life history information was known concerning this species, a study was conducted to determine its food habits rela- tive to sea.sons, fish length, diel feeding chro- nology, and food habits of other Alvord Basin fishes of the genus Gila. Methods Feeding ecology of Gila boraxobius was in- vestigated by analyzing intestinal contents of fish collected monthly from March 1978 to June 1979. Fish were collected from the southwest one-quarter of Borax Lake using a 3 X 5 mm me.sh seine approximately 3 m in length. Specimens were preserved in 10 per- cent formalin and transferred to 45 percent isopropanol after one week. Standard length of specimens was measured to the nearest 0.1 mm with dial calipers. After blotting fish dry on paper towels, wet weight was measured to 'Technical Paper 5335, Oregon .Agricultural Experiment Station. ■Department of Fisheries and Wildlife. Oregon State University. Corvallis, Oregon 97331. 101 102 Great Basin Naturalist Vol. 40, No. 2 the nearest 0.01 g. Gila boraxohius, like other cvprinids, has no stomach; therefore, contents of the intestine, from esophagus to anus, were removed and examined under dissecting mi- croscopes. Percent frequency of occurrence, mean number per intestine, mean percent volume, and a value of relative importance are reported for each food item. Percent fre- quency of occurrence is defined as the num- ber of intestine samples in which one or more of a given food item is found expressed as a percentage of all nonemptv intestines exam- ined (Windell and Bo wen 1978). The total number of a given food item observed in the intestines divided by the number of nonemp- tv intestines examined is the mean number per intestine. Mean percent volume is de- fined as the total volume estimates for a giv- en food item divided by the number of non- empty intestines examined. Percent volumes were derived by separating the intestine into three subsamples and visually estimating the percent contribution of a given food item in each sample. The percent contribution of each subsample to the contents of the entire intestine was estimated so that the volume of a <£iven food item relative to all intestinal contents could be made. Percent frequency of occurrence, mean number per intestine, and mean percent volume each contain a bias which limits their usefulness when used sepa- rately (Windell and Bowen 1978). For ex- ample, percent frequency of occurrence overemphasizes the importance of small food items that may be ingested frequently but have a small impact on the volume of food in the intestine. On the other hand, mean per- cent volume overemphasizes the importance of large food items, such as adult insects, that may be consumed infrequently but have a large volume. To offset these biases against each other, an index of relative importance (RI) is reported for each food item. The rela- tive importance index combines the percent frequency of occurrence and mean percent volume for food item a into an absolute im- portance index (AI„) as follows: AT = P^''<-'(-'"t frequency of occurrence + mean percent volume RI„ = 100AI„/SAI„, where n is the number of different food items. The determination of Rio and AI^ are by methods modified from George and Hadley (1979). Invertebrate identification was based on Pennak (1978). Food habits of G. boraxohius are summa- rized as follows: (1) seasonal variation of foods consumed, (2) foods consumed by dif- ferent size classes of fish, (3) diel feeding chronology, and (4) comparison with food habits of two populations of Gila alvordcnsis. Changes in food habits with season and fish size class were determined from monthly col- lections taken from March 1978 through Jan- uary 1979. Monthly collections were grouped into seasons as follows: spring (March-May), summer (June-August), autumn (Septem- ber-November) and winter (Decem- ber-January). To compare changes in food habits with fish size, two size classes of fish were compared, 15.0 mm to 32.9 mm SL (juveniles), and 33.0 mm SL or longer (adults). Gila boraxobius matures at approx- imately 33.0 mm SL (Williams and Bond, in press). Diel feeding chronology was deter- mined from collections made at 3-hour inter- vals during a 24-hour period in June 1979. Fullness of the intestine was determined ac- cording to the formula defined by Hureau (1969 in Berg 1979) as follows: _ weight of ingested food X 100 percent weight of fish (Ir = L'indice de repletion = fullness index). The daily ration of food for G. boraxobius was estimated from diel trajectories of the fullness of the intestine. This estimate was de- rived by the following formula proposed by Bajkov (1935) and modified by Darnell and Meierotto (1962) and Eggers (1977): Rt = 24Sa where Rj is the daily ration, S is the average weight of intestinal contents expressed as percent of body weight during the 24-hour period, and a is the intestinal evacuation rate. An intestinal evacuation rate of 0.2 hr' is assumed (Eggers 1977). Intestinal contents of Gila alvordcnsis col- lected from Thousand Creek, Nevada, on 13 June 1978 and G. alvordensis collected from Serrano Pond, Oregon, on 6 August 1977 were compared to those of G. boraxobius col- June 1980 Williams, Williams: Borax Lake Chub 103 lected during the summer of 1978. Methods of collection and food habits analysis for pop- ulations of G. alvordensis were the same as those used for G. boraxobius. Study Area Fish were collected from Borax ( = Hot) Lake, Serrano Pond, and Thousand Creek, all in the Alvord Basin of southeastern Oregon and northwestern Nevada. The Alvord is an endorheic basin of semiarid climate, sur- rounded by fault-block mountain ranges. Bo- rax Lake'(T37S, R33E, Sec. 14, Harney County, Oregon) is a 4.1 ha natural lake that receives water from several thermal springs in the lake bottom. Water temperature in Borax Lake is typically 29-32 C, with ex- tremes of 35 C or greater near spring sources and 17 C near the lake margin during winter. The lake is relatively shallow, with a soft, sil- tv bottom interspersed with rocks and hard outcroppings. The water is transparent, and aquatic vegetation is limited to a few areas along the lake margin. Sodium is the princi- pal cation, whereas bicarbonate, sulfate, and chloride are the major anions in Borax Lake (Mariner et al. 1974). Specific conductance of the water is 2410. Serrano Pond (T36S, R33E, Sec. 1, Harney County, Oregon) is a 0.1 ha reservoir that receives water from a cool spring approximately 60 m distant. Wa- ter temperature is usually 16-21 C during the summer. The bottom of this shallow pond is primarily silt. The water is turbid, and aquat- ic vegetation is abundant. Thousand Creek (collection site at T46N, R28E, Sec. 34, Humboldt County, Nevada) is a small, shal- low stream rarely exceeding 2 m in width. The creek often becomes intermittent during summer months. Water temperature varies from 15-27 C during the summer, fluctuating rather closely with air temperature. The bot- tom is a silt and gravel mix. The water is tur- bid, and aquatic vegetation is moderately abundant. Results Seasonal Variation in Foods Consumed Twenty-four different food items were found in intestines of Gila boraxobius during this study (Tables 1-4). Ten of the 24 food items were encountered in all seasons. Many food items fluctuated seasonally in occur- rence; however, some insects, especially chi- ronomid larvae, diatoms, and micro- crustaceans, were of importance throughout the year. During the spring, algae, chirono- mid larvae, copepods, dipteran adults, and ostracods were the predominant food items (Table 1). Algae, which was composed almost wholly of diatoms, was the most frequently ingested food during the spring, occurring in over one-half of the intestines. Some of the diatoms were secondarily ingested with mi- crocmstaceans; however, in some individuals diatoms accounted for 70-80 percent of the intestinal contents by volume. The high vol- ume suggests that diatoms are not exclusively the result of secondary ingestion but are a preferred food item for most fish. The most common diatom observed in intestines was a benthic species, Denticida thermalis. Nov- iciila sp., Synedra sp. Achnanthes lanceolata, and A. minutissima were observed in lesser numbers. Both Achnanthes species are ben- thic, whereas the Navicula and Synedra spe- cies could be benthic or planktonic forms. Adult dipterans accounted for the highest mean percent volume, nearly 20 percent, of all food items during spring. Several fish fed on dipteran adults exclusive of other foods. During May 1978 dipteran adults were heav- ily utilized, appearing in 19 of 23 intestines examined. During the summer, diatoms were less frequently encountered in intestines and comprised a smaller mean percent volume than in spring. Chironomid larvae, copepods, dipteran adults, and gastropods were the most important food items in summer (Table 2). Gila boraxobius utilized more terrestrial insects and spiders and fewer micro- crustaceans and diatoms in summer than in spring. Terrestrial insects and spiders ac- counted for approximately 31 percent mean volume of foods consumed during summer compared to approximately 21 percent in spring. During autumn terrestrial insects, chi- ronomid larvae, and diatoms were the princi- pal food items (Table 3). In winter Gila boraxobius relied more heavily on autoch- thonous food items, utilizing primarily dia- toms, ostracods, copepods, chironomid larvae and cladocerans (Table 4). Terrestrial insects. 104 Great Basin Naturalist Vol. 40, No. 2 which were of importance in spring, summer, and autumn, seldom appeared in intestines of G. boraxobius during winter, when they con- tributed only 2 percent mean volume. Food items consumed in autumn and winter were less diverse than in other seasons. Sixteen food items were observed in fish collected in autumn and winter, 20 food items during spring, and 21 during summer. Aquatic in- sects were important foods throughout the year, comprising mean volumes of approx- imately 19 percent, 23 percent, 16 percent, and 13 percent in spring, summer, autumn, and winter, respectively. The primary con- tributor to these high values were chirono- mid larvae, which consistently exhibited a high relative importance. Chironomid pupae and Odonata nymphs were also consumed throughout the year but were of much less importance. Coleoptera larvae and aquatic Coleoptera adults were utilized to lesser de- grees seasonally. The increased consumption of copepods, ostracods, and cladocerans in the winter was dramatic. These micro- CRistaceans comprised approximately 35 per- cent mean volume of intestines during win- ter, but only 16.5 percent, 12 percent and 4.5 percent in spring, summer, and autumn, re- spectively. Large amounts of inorganic debris were found in intestines throughout the year. Tliis was probably ingested accidently while the fish were feeding on bottom organisms. Many important foods in Borax Lake, such as insect larvae, gastropods, diatoms, and prob- ably many small invertebrates, are benthic. Observations in Borax Lake and in aquaria show that G. boraxobius feeds primarily by rooting around in bottom material and pick- Table 1. Contents of 71 intestines oi Cihi boraxobius collected during the spring of 1978. ND = no data. Item ingested Percent frequency of occurrence Mean number per intestine Mean percent volume RI .\lgae' C'.astropods- C.astropod eggs Haplotaxid oligochaetes Harpacticoid copepods Ostracods Cladocerans Plant seeds Higher plants Fish scales Araneae Insect eggs Unidentified insects' Tekrestrial i\sec;ts CoUemhola Thysanoptera adults Hemiptera adults Coleoptera adults Hymcnoptera adults Diptera adults Asiatic: insects Chironomid larvae C;hironomid pupae Odonata nvmphs Klmid larvae C'oleoptera adults In()Iu;amc: dehhis 56.14 ND 12.67 15.70 22.81 1.07 6.69 6.73 0.00 0.00 0.00 ().(M) 1.75 0.11 0.32 0.47 49.12 44.95 10.38 13..57 40.35 8.39 5.91 10..55 15.79 0.21 0.17 3.64 5.26 0.11 0.06 1.21 12.28 ND 1.72 3.19 3.51 0.11 0.04 0.81 3.51 0.04 0.34 0.88 10.53 12.26 2.15 2.89 26.32 ND 2.50 6..57 0.00 ().(H) 0.(X) 0.00 3.51 0.04 0..39 0.89 0.00 0.00 0.00 0.(K) 1.75 0.02 0.21 0.45 3.51 0.04 0.49 0.91 35.09 1.93 19.49 12.45 47.37 12.30 13.71 13.93 7.02 0.14 2.32 2.13 3.51 0.04 0.91 1.01 7.02 0,39 1.76 2.(X) 0.00 {).()() ().(K) 0.(X) 61.40 ND 17.48 - Total 99.71 'Mostly diatoms 'Mostly PUinorhulla, rarely Phy.m 'Mostly terrestrial forms June 1980 Williams, Williams: Borax Lake Chub 105 ing up food items. However, if Ijenthic food items are scarce, or if other foods are abun- dant, C. homxohius will readilv feed on ma- terials drifting through water column or on the surface. Thus, during the summer some G. boraxohius readilv switched to ingestion of terrestrial invertebrates. This resulted in the lowest mean percent volume of inorganic debris ingested for any season. The shoreline of Borax Lake provides habitat for many ter- restrial invertebrates that can enter Borax Lake. Terrestrial invertebrates are scarce during winter, reducing the likelihood of their being a primary food source at that time. Gila boraxohius is often a highly exploitive omnivore, feeding almost entirelv on one food source. For example, examination of in- testines of fish collected during May 1978 disclosed the following (percent volumes of Tahle 2. C'ontents of 70 intestines of Cila boraxob'uts colh the food item are given in parentheses): one individual contained 32 gastropods (84 per- cent volume), a second fish contained 14 adult dipterans (98 percent volume), a third contained 775 copepods (79 percent volume), a fourth contained 340 first instar chironomid larvae (69 percent volume), and a fifth con- tained 485 insect eggs (64 percent volume)! Although the preceding is somewhat unusual, many fish were found with one food item dominating their intestinal contents. Foods Consumed by Different Size Classes of Fish To study the effect of fish size and age on foods consumed, we compared intestinal con- tents of juvenile and adult Gila boraxohius. Overall food habits of juveniles and adults were similar. Both consumed large amounts :ted di tile summer of 1978. ND = no data. Item ingested Percent frequency of occurrence Mean number per intestine Mean percent volinne RI ■\lgae' Gastropods- Gastropod eggs Haplotaxid oligochaetes Harpacticoid copepods Ostracods Cladocerans Plant seeds Higher plants Fish scales Araneae Insect eggs Unidentified insects' Terrestrial insects Collembola Thvsanoptera adults Hemiptera adults Coleoptera adults H\ nienoptera adults Diptera adults Asiatic: insects C!hironoinid larvae (Ihironomid pupae Odonata nvmphs Elmid larvae Coleoptera adults Inorganic debris 31.15 ND 6.42 7.42 32.79 1.07 9.04 8.26 1.64 0.66 1.56 0.63 1.64 0.08 0..30 0.38 47.54 33.30 8.45 11.06 32.79 2.87 3.03 7.08 8.20 0.38 0.31 1.68 13.11 0.25 0.49 2.69 0.00 0.00 0.00 0.(X) ().(X) 0.00 0.00 0.(X) 14.75 0.25 3.21 3..55 3.28 0.69 0.15 0.68 37.70 ND 4.50 8.34 6.56 0.16 0.23 1.34 9.84 0.13 0.45 2.03 0.00 0.00 0.00 0.00 26.23 0.70 5.94 6.36 18.03 0.21 2.27 4.01 39.34 1.62 14.31 10.60 .57.38 4.10 13.79 14.06 9.84 0.11 1.12 2.17 1.64 0.03 0.49 0.42 11.48 0.23 0.85 2.44 18.03 0.31 6.29 4.80 .54.10 ND 15..30 - Total 98.50 'Mostly diatoms 'Mostly Planorhulla. rarely Phy.sa 'Mostly terrestrial forms 106 Great Basin Naturalist Vol. 40, No. 2 of terrestrial insects in spring, summer, and autumn, and algae (mostly diatoms) and mi- crocm-staceans in winter (Fig. 1). Aquatic in- sects, primarily chironomid larvae, were im- portant to juveniles and adults throughout the year. Despite the overall similarity of food habits between juveniles and adults, some differences were noted. Elmid (Coleop- tera) larvae were consumed almost exclusi- vely by adults. Elmid larvae were found in intestines of 15 adults but only one juvenile. More terrestrial insects were consumed by juveniles than adults except during winter, when terrestrial insects were relatively unim- portant to both groups. The relatively large size of many terrestrial insects, such as the commonly consumed muscoid fly adults, did not deter their ingestion by juvenile Gila horaxohius. Adults consumed more gastro- pods than did juveniles during all seasons. In- testines of adults averaged 8.5 percent mean volume of gastropods during the year, where- as intestines of juveniles averaged 2.3 per- cent. Adults also consumed more diatoms than did juveniles. Increased relative con- sumption of diatoms by adults was primarily evident in summer and autumn, when adults consumed 9.3 percent and 6.9 percent mean volume of diatoms, respectively; however, juveniles consumed 0.1 percent and 1.3 per- cent mean volume, respectively. The small volume of gastropods and diatoms ingested by juveniles was compensated for by inge- stion of large numbers of copepods. Intestines of juveniles averaged 13.4 percent mean vol- ume of copepods during the year, whereas adults averaged 4.2 percent mean volume of copepods. Table 3. Contents of 57 intestines of C.iUi horaxohius collected during the autumn of 1978. ND = no data. Item ingested Percent frequency of occurrence Mean number per intestine Mean percent volume RI Algae' Gastropods- Gastropod eggs Haplotaxid oligochaetes Harpacticoid copepods Ostracods Cladocerans Plant seeds Higher plants Fish scales Araneae Insect eggs Unidentified insects' Terrestrial insects Collembola Thysanoptera adults Hemiptcra adults Coleoptera adults Hymenoptera adults Diptera adults AgUATIC INSECTS Chironomid larvae Chironomid pupae Odonata nymphs Elmid larvae Coleoptera adults I.NORGANIC DEBRIS 50.00 ND 5.03 12.63 10.87 0.50 4.04 3.42 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 34.78 25.46 4.22 8.95 10.87 0.50 0.15 2.53 0.00 0.00 0.00 0.00 39.13 1,57 0..59 9.12 2.17 ND 0.07 0.51 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 2.17 0.04 0.04 0.51 67.39 ND 19.54 19.95 0.00 0.00 0.00 0.00 21.74 0.46 1.69 5.38 13.04 0.22 2.55 3.58 4.35 0.04 0.22 1.05 8.70 0.11 l.,54 2.35 19.56 0.22 3..52 5.30 71.74 5.43 12.00 19.22 13.04 0.65 3.15 3.72 6.52 0.07 1.29 1.79 0.00 0.(K) 0.(X) 0.00 ().(X) 0.(X) 0.(K) 0.00 86.96 ND 40.34 - Total 99.94 'Mostly diatoms 'Mostly Planorhulla, rarely Physa 'Mostly terrestrial forms June 1980 Williams, Williams: Borax Lake Chub 10' Diel Feeding Chronology Feeding chronologv and daily ration were determined bv the relative weight of material ingested bv CUla honixohiiis collected during a 24-hour period in June 1979 (Fig. 2). The average weight of fish was 1.21 g. Gila horax- ohius fed throughout the day with peak feed- ing activity shortly after sunset. Minimal feeding activities occurred after sunrise. An increase in feeding activity after sunset has been observed in Gila hicolor (Snyder 1917). The average weight of ingested material in intestines, as determined from 1800-1500 hours, was 2.32 percent of body weight. This average weight of ingested material (S) was used to determine the daily ration (Rj) as fol- lows: Rt = 24Sa = 24(2.32)(.2) = 11.14 By the above method we calculated that G. horaxohius ingested 11.14 percent of their body weight daily. This estimate is larger than most reported by researchers for other species. Brett (1979) summarized in- vestigations made by various researchers who calculated daily rations that were typically 2-5 percent of body weight. Several studies have noted increased relative ration with in- creased temperature (e.g., Brett et al. 1969, Kinne 1960, Stauffer 1973) and with smaller fish size (e.g., Brett 1971, Brett and Shelbourn 1975, Elliott 1975). Brett (1979) reported that temperature and fish size were of greatest importance in determining ration size. The dwarf size of G. horaxohius and its Table 4. Contents of 62 intestines oi Gila hoiaxobius collected during the winter of 1978-79. ND = no data. Item ingested Percent frequency of occurrence Mean nmuber per intestine Mean percent volume RI Algae' Gastropods- Gastropod eggs Haplotaxid oligochaetes Harpacticoid copepods Ostracods Cladocerans Plant seeds Higher plants Fish scales Araneae Insect eggs Unidentified insects' Terrestrial i\sec:ts C'ollembola Thvsanoptera adults Ileniiptera adults Goleoptera adults Hynienoptera adults Diptera adults AyiATic: insects ('hironomid larvae (;hir()nomid pupae Odouata nymphs Elmid larvae Goleoptera adults I.N()R{;ank: debris 94.44 ND 18.81 22.05 18.52 0.54 3.88 4.36 0.00 0.00 0.00 0.(K) 5.56 0.91 2.21 1,51 75.93 16.44 9.96 16.72 77.78 21.65 20.58 19.15 55.56 6.52 4.07 11.61 3.70 0.04 0.02 0.72 5.56 ND 0.35 1.15 1.85 0.04 0.03 0.37 0.00 0.00 ().(X) (),(K) 0.00 0.00 (),()() 0.00 9.26 ND 1.49 2.09 1.85 0.04 0.33 0.42 0.00 0.00 0.00 0.(X) 0.00 0.00 0.(K) 0.00 0.00 0.00 0.(K) 0.(K) 0.00 0.00 ().(K) 0.00 1.85 0.02 0.15 0.39 70.37 5.15 10.94 15.83 1.85 0.02 0.22 0.40 5.56 0.07 0.77 1.23 9.26 0.11 0.84 1.97 0.00 0.00 ().()() ().(M) 90.74 ND 25.37 - Total 100.02 'Mostly diatoms ■Mostly PkmorhuUa. rarely Physa Mostly terrestrial forms 108 Great Basin Naturalist Vol. 40, No. 2 gastropod gastroDod copepod algae est racod copepod algae debris ostracod other debris other terrestrial insect JUVEN I LES ADULTS SPRING gastropod 2.5 0.1 algae copepod ostracod other gastropod ebris aquatic insect other teirest rial i nsec t JUVENI LES ADULTS SUMMER Fig. 1. Comparison of food habits between 128 juvenile and 1.32 adult Cila Inmixohitis. Mean percent volume of food items are given in circle. June 1980 Williams, Williams: Borax Lake Chub 109 copepod terrestrial insect debris copepod gastropod istracod 0.2 debris 0.9 other JUVENILES ADULTS AUTUMN 0.9 gastropod 1 copepo alqae est racod gast ropod copepo ost racod debris ther aquatic insect i 1.0 terrestrial insect debris f other 2-^ terrestrial insect JUVENI LES ADULTS Wl NTER Fig. 1 continued. 110 Great Basin Naturalist Vol. 40, No. 2 habitation in thermal spring waters contrib- uted to the large daily ration found in this species. Many researchers determined daily ration using prepared foods that were per- haps more nutritious than material ingested by Gila boraxobiiis, which included such un- digestible items as insect exoskeletons, gastro- pod shells, and inorganic debris. The pres- ence of large amounts of undigestible material increases S because the fish must in- gest a larger volume of material to get enough calories. This results in a larger daily ration than would be calculated if fish con- sumed only digestible foods. Food Habits of Gila ahordensis Ten food items were foimd in intestines of Gila alvordensis collected from Thousand Creek, Nevada, in June 1978 (Table 5). Of the ten food items, chironomid larvae, cla- docerans, copepods, and ostracods were of greatest importance. Chironomid larvae oc- curred in all intestines examined and ac- counted for approximately 26 percent mean volume of intestines. Microcrustaceans com- prised almost 45 percent mean volume of in- testines. Diatoms accounted for 5 percent mean volume of intestines. No terrestrial in- sects were ingested by G. alvordensis from Thousand Creek. Eleven food items were found in intestines of Gila alvordensis collected from Serrano Pond, Oregon, in August 1977 (Table 6). Of the eleven food items, chironomid larvae, di- atoms, cladocerans, and ostracods were of greatest importance. Chironomid larvae oc- curred in over three-quarters of the intestines examined and accounted for approximately 50 percent mean volume of all intestines. Di- atoms occurred in one-half of the intestines and accounted for almost 23 percent mean volume of all intestines. Microcrustaceans comprised approximately 17 percent mean volume of intestines. No intestines contained terrestrial insects. Gila alvordensis from Ser- rano Pond were highly opportimistic feeders. Eighty-nine percent of the fish from Serrano Pond with food in their intestines contained one item that accounted for more than 50 percent of their intestinal volume. Thirty- nine percent of fish contained one food item that comprised 90 percent or more of in- testinal volume. This exploitive feeding was focused on chironomid larvae, cladocerans, or algae. One intestine was exclusively filled with 2570 cladocerans. Such exploitive feed- ing was not noted in Gila alvordensis from Thousand Creek. A comparison of foods of Gila alvordensis collected during June and August with foods of Gila boraxobiiis collected during the sum- mer shows several differences. Terrestrial in- sects were important foods for Gila borax- obiiis during the summer but were absent from intestines of Gila alvordensis. Gila boraxobiiis also consumed larger quantities of other terrestrial foods, such as spiders and in- sect eggs, than did G. alvordensis. Intestines of G. alvordensis from Thousand Creek, and to a lesser extent those from Serrano Pond, contained much larger amounts of micro- ciTistacea than did intestines of G. boraxobiiis during the summer. Diatoms were a major food item of fish in Serrano Pond during the simimer, but not of fish in Thousand Creek or of Gc boraxobiiis. Gila boraxobiiis consumed a larger number of food items than either population of G. alvordensis. This is due to the greater opportimism, including the use of terrestrial foods, exhibited by G. boraxobiiis. A larger sample size may also contribute to the greater diversity of foods utilized by G. boraxobiiis. Discussion Gila boraxobiiis and G. alvordensis have tentatively been included with G. bicolor in the subgenus Siphateles (Hubbs and Miller 1972, Williams and Bond, in press). Although no life history information has previously been published for G. boraxobiiis or G. alvor- densis, several researchers have examined food habits of G. bicolor and concluded that they are primarily opportunistic omnivores (Bird 1975, Cooper 1978, La Rivers 1962). However, differences in food habits between the coarse gill raker form, G. b. obesa, and the form with numerous, fine gill rakers, G. b. pectinifer, have been noted. In describing habits of the form with coarse gill rakers from Lake Tahoe, Miller (1951) foimd them to be primarily benthic feeders, with a diet June 1980 Williams, Williams: Borax Lake Chub 111 composed of 89 percent bottom organisms. Snyder (1917) noted that the form with coarse gill rakers collected from the littoral zone of Lake Tahoe fed on algae, other plant material, and insects. Gila b. pectinifer from Lake Tahoe, with its numerous gill rakers, fed almost exclusively on midwater micro- crustacea (Miller 1951). La Rivers (1962) also reported that G. b. pectinifer contained many midwater foods in their intestines, primarilv consuming diatoms and microcrustaceans. Cooper (1978) reported that the form with numerous gill rakers (although he referred it to G. h. obesa) in Walker Lake, Nevada, fed mostly on zooplankton and filamentous al- gae. A population complex of G. bicolor in Eagle Lake, California, that included forms with both coarse and fine gill rakers fed on a variety of foods, including zooplankton, plant material, insect larvae, and surface insects (Kimsey 1954). There appears to be a definite correlation between gill raker morphology Table 5. Contents of 21 intestines of Gila uhordctisis collected 13 June 1978 from Thousand Creek, Nevada. ND = no data. Item ingested Percent frequency of occurrence Mean number per intestine Mean percent volume RI Diatoms Gastropods Harpacticoid copepods Ostracods Cladocerans Plant seeds Araneae AgiL-VTIC INSECTS Chironomid larvae Chironomid pupae Coleoptera adults I.NORCANIC: DEBRIS 36.36 ND 5.00 7.91 9.09 0.09 0.36 1.81 81.82 6.82 6.82 16.94 72.73 15.82 13.73 16.53 90.91 27.73 24.23 22.01 27.27 0.27 0.45 5.30 9.09 0.09 0.55 1.84 10().(K) 16.73 25.78 24.04 9.09 0.09 0.09 1.75 9.09 0.09 0.68 1.87 lOO.(K) ND 20.95 - Total 98.64 Table 6. Contents of 20 intestines of Gila akordensis collected 6 August 1977 from Serrano Pond, Oregon. ND = no data. Item ingested Percent frequency of occurrence Mean number per intestine Mean percent volume RI Diatoms Harpacticoid copepods Ostracods Cladocerans .Araneae Insect eggs Unidentified insects .\quatic insects Chironomid larvae Chironomid pupae Odonata nymphs Ephemeroptera larvae Inorganic debris 50.00 ND 22.94 18.16 .33.33 1.67 0.36 8.39 38.89 5.94 1.70 10.11 55.56 253.28 15.34 17.65 5.56 0.06 0.22 1.44 5..56 0.06 0.05 1.40 5.56 ND 0.05 1.40 77.78 20..33 50.41 31.92 22.22 0.33 2.96 6.27 5.56 0.06 0.67 1.55 5.56 0.06 1.33 1.72 55..56 ND 3.96 - Total 99.99 112 Great Basin Naturalist Vol. 40, No. 2 and food habits, tho.se with coarse gill rakers ingesting more benthic food organisms and those with fine gill rakers ingesting more zooplankton. Gila boraxobius and G. alvor- densis possess approximately 16 and 20 short gill rakers, respectively, agreeing closely with the gill raker morphology of G. bicolor obesa form. Although G. boraxobius typically feeds on benthic organisms, large amounts of dia- toms, microcnistaceans, and terrestrial insects are ingested sea.sonally. The ingestion of terrestrial insects by Gila is not common. However, several researchers have found that terrestrial insects comprised a small part of the diet of Gila (Cross 1978, Kimsey 1954, Moyle 1976, Sigler and Miller 1963). Terrestrial insects were the primary foods of G. robusta and G. elegaas longer than 200 mm SL collected from the Green River (Vanicek and Kramer 1969). Smaller G. robusta and G. elegans contained pre- dominantly aquatic insect larvae. Juvenile and adult G. boraxobius consumed large quantities of terrestrial insects. Several re- searchers (Kimsey 1954, Miller 1951) have noted that as Gila grow, they switch to larger food items; however, at least one study (Gra- ham 1961) found foods of different sized groups of CAla to be nearly identical. We found foods of juvenile and adult G. borax- obius to be very similar, except that adults exhibited a greater consumption of gastro- pods and diatoms, and juveniles consumed more copepods and terrestrial insects. The hard shells and, to a lesser extent, the rela- tively large size of gastropods probably con- tributed to juveniles avoiding them as a food source. Larger Gila ingest more algae than do smaller fish in studies by Moyle (1976) and Vanicek and Kramer (1969). Age II Gila coe- rulea feed predominantly on filamentous al- gae, whereas algae were entirely or prac- tically absent from age I fish (Moyle 1976). Large adult Gila from the Green River con- sumed more algae than did smaller fish (Van- icek and Kramer 1969). Juvenile G. borax- Tl ME OF DAY Fig. 2. Feeding throiiologv^ of Cila boraxobius on 5-6 June 1979. N = 72. June 1980 Williams, Williams: Borax Lake Chub 113 obitis consumed more terrestrial insects than did the adults, except in winter, when small amoiuits of terrestrial insects were ingested by both groups. The reason for juveniles con- suming large volumes of terrestrial insects is unknown; apparently the relatively large size of this food item is not a deterrent. Both juvenile and adult G. horaxoJyiiis in- creased consumption of diatoms and micro- crustaceans in winter. This probably in- dicates a scarcity of food items during the winter, as is reflected by finding 24 percent fewer food items in intestines during winter than in summer. A factor contributing to the winter scarcity of food items is a decrease in the availability of terrestrial foods, causing concentrated feeding on remaining food items. For example, the Devil's Hole pupfish, Cyprinodon diabolis, which inliabits a small thermal spring, dramatically increases in- gestion of diatoms during winter due to a scarcity of preferred foods (Minckley and Deacon 1975). Although the amount of nutri- tion derived from consuming diatoms is un- known, we suspect that the large con- sumption of diatoms at certain times of the year by G. bomxobius would indicate that some nutritive value is gained. Arnold (1971) found that species of Ci/prinodon derived oil droplets from ingested diatoms, thus extract- ing nutritive value. A similar mechanism could operate in Gila bomxobius. Examination of summer foods of both G. ah'ordensis populations showed differences from the summer foods of G. boraxobius. During summer Gilo boraxobius relied heavi- ly on terrestrial food items, whereas popu- lations of G. alvordensis consumed prac- tically no terrestrial foods. Forty-three percent of food items consumed by G. bora- xobius during the summer were of terrestrial origin. Surveys of potential food items in Bo- rax Lake conducted at various times of the year found that all potential food items were utilized by G. boraxobius except some adult hemipteran and coleopteran insects that were probably too large to be ingested. Also, many hemipterans possess scent glands that render them impalatable to predators. Acknowledgments The authors are indebted to Carl E. Bond for his guidance during the course of this study and for his review of the manuscript. Stanley V. Gregory provided identification of diatoms and information on their habitats, the Oregon Department of Fish and Wildlife provided collecting permits, and James J. Long, Kevin M. Howe, and Glen DeMott as- sisted with field collections. Kevin M. Howe reviewed the manuscript and provided field notes of the Serrano Pond area. 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Determi- nation of feeding chronology in fishes. Trans. .\in. Fish. Soc. 91:313-320. Deacon, J. E., G. Kobetich, J. D. Willlams, S. Contreras, and other members of the Endangered Species Committee of the 114 Great Basin Naturalist Vol. 40, No. 2 American Fisheries Society. 1979. Fishes of North America endangered, threatened, or of special concern: 1979. Fisheries. 4:29-44. EcGERS, D. M. 1977. Factors in interpreting data obtain- ed bv die] sampling of fish stomachs. J. Fish. Res. Bd. Can. 34:290-294. Elliott, J. M. 1975. Number of meals in a day, max- imum weight of food consumed in a day and maximum rate of feeding for brown trout, Salmo trtitta L. Freshwater Biol. 5:287-303. George, E. L., and W. F. Hadley. 1979. Food and habi- tat partitioning between rock bass (Ambloplites rupestris) and smallmouth bass (Micropteriis do- lomieui) yoimg of the year. Trans. Am. Fish. Soc. 180:253-261. Graham, R. J. 1961. Biology of the Utah chub in Heb- gen Lake, Montana. Trans. Am. Fish Soc. 90:269-276. HuBBS, C. L., AND R. R. Miller. 1972. Diagnoses of new cyprinid fishes of isolated waters in the Great Ba- sin of western North America. Trans. San Diego Soc. Nat. Hi.st. 17:101-106. HuREAU, J. C. 1969. Biologie comparee de quelques poissons antarctiques (Nothotheniidae). Bull. In.st. Oceanogr. Monaco. 68:1-44. KiMSEY, J. B. 1954. The life hi.story of the tui chub, Siph- ateles bicolor (Girard), from Eagle Lake, Califor- nia. Calif. Fish Game. 40:.395-410. KiNNE, O. 1960. Growth, food intake, and food con- version in a euryplastic fish exposed to different temperatures and salinities. Physiol. Zool. 33:288-317. La Rivers, 1. 1962. Fishes and fisheries of Nevada. Ne- vada State Fish Game Comm. 782 pp. Mariner, R. H., J. B. Rapp, L. M. Willey, and T. S. Presser. 1974. The chemical composition and es- timated minimum thermal reservoir temper- atures of selected hot springs in Oregon. U.S. Geol. Surv. Open-File Report. Menlo Park, Calif. Miller, R. G. 1951. The natural history of Lake Tahoe fishes. Unpublished dissertation. Stanford Univ. 160 pp. MiNCKLEY, C. O., AND J. E. Deacon. 1975. Foods of the Devil's Hole pupfish, Cijprinodon diaboUs (Cy- prinodontidae). Southwestern Nat. 20:105-111. MoYLE, P. B. 1976. Inland fishes of California. Univ. California Press, Berkeley. 405 pp. Pennak, R. W. 1978. Freshwater invertebrates of the United States, 2d ed. Wiley-lnterscience, New York. 803 pp. SiGLER, W. F., AND R. R. MiLLER. 1963. Fishes of Utah. Utah State Dept. Fish Game, Salt Lake City. 203 pp. Snyder, J. O. 1917. The fishes of the Lahontan system of Nevada and northeastern California. U.S. Bureau Fisheries Bull. .35 (for 1915-16):3.3-86. Stauffer, G. D. 1973. A growth model for salmonids reared in hatchery environments. Unpublished dissertation. Univ. Washington. Vanicek, C. D., and R. H. Kramer. 1969. Life history of the Colorado squawfish, Ptijclwcheilus luciiis, and the Colorado chub, Gila robusta, in the Green River in Dinosaur National Monument, 1964-1966. Trans. Am. Fish. Soc. 98:19.3-208. Williams, J. E., and C. E. Bond. Gila boraxobius, a new species of cyprinid fish from southeastern Oregon with a comparison to G. alvordensis Hubbs and Miller. Proc. Biol. Soc. Wash. Vol. 93. Windell, J. T., and S. H. Bowen. 1978. Methods for study of fish diets based on analysis of stomach contents. Pages 219-226 in T. Bagenal, ed.. Methods for assessment of fish production in fresh waters, .3d ed. Blackwell Scientific Pub- lications, Oxford. FIRST RECORD OF THE PALLID BAT (ANTROZOUS PALLIDUS) FROM MONTANA Jeff Shryer' and Dennis L. Flath' .\bstract.— a pallid bat {Antrozous pallidiis) was taken 20 km SE Warren, Carbon County, Montana. This repre- sents a 410 km range extension and a new record for Montana. The pallid bat {Antrozous pallidiis) has not been previously reported from Montana. On the basis of specimens from Grangeville and Pocatello, Idaho, Hoffman and Pattie (1968) suggested that the species may occur in southwestern Montana. Fichter (1964) report- ed A. pallidus from Boise, Idaho. On 25 August 1978 we mist-netted an adult female A. pallidus (Montana Fish, Wildlife, and Parks Collection NG 748) at an isolated spring approximately 20 km SE Warren, Carbon County, Montana. This rec- ord extends the known range of A. pallidas approximately 410 km northeast of Pocatello, Idaho, and approximately 460 km north of Dinosaur Quarry, Utah, where Knitzsch and Heppenstall (1955) obtained a specimen. The new locality is in an alluvial plain south of the Pryor Mountains, at 1370 m ele- vation, and lies within the 25-30 cm precipi- tation zone (U.S. Soil Conservation Service 1977). The area is characterized by scattered outcrops of sandstone and .shale, with a vege- tative community dominated by western wheatgrass {Agropyron smitliii), sagebrush {Artemisia spp.), and saltbush (Atriplcx spp.). External measurements of the specimen are as follows: total length, 110 mm; length of tail, 37 mm; length of hind foot, 13 mm; length of ear, 29 mm; and length of forearm, 59 mm. The dentition displaved substantial wear, indicating the bat was old. This speci- men is similar to specimens in the University of Montana Zoological Museum from New Mexico (MSUZ 10419) and Arizona (MSUZ 12996) in that it is of a similar color— a very pale fawn. Allen (1864) described A. p. pal- lidas as having two varieties of color, fawn and yellowish brown. Bailey (1936) described A. p. cantwelli as darker than pallidus, with dark brown ears and membranes. The mem- branes of our specimen are medium brown. Other species captured concurrently in- clude: Myotis lucifugus and M. leihii. We thank John Ciralli for field a.ssistance and P. L. Wright for assistance in the identi- fication of the specimen. This paper is a con- tribution of the U.S. Bureau of Land Manage- ment, Montana Nongame Program, and the Montana Department of Fish, Wildlife, and Parks, Nongame Program, State Project 5853. Literature Cited .\LLEN, H. 1864. Monograph of the bats of North Ameri- ca. Smithsonian Misc. Coll. 165:68-69. .\.\o.NYMOUS. 1977. Average annual precipitation for Montana based on 1941-1970 base. USDA, Soil Cons. Serv., Portland, Ore. 13pp. B.\iLEY, V. 1936. The mammals and life zones of Oregon. North American Fauna 55:390-.393. Fichter, E. 1964. The pallid bat in Idaho. Tebiwa 7:23-27. Hoffman, R. S., a.nd D. L. Pattie. 1968. .-^ guide to Montana mammals: Identification, habitat, distri- bution and abundance. Univ. Montana Print. Serv. 1.33 pp. Krutzsch, p. H., and C. A. Heppenstall. 1955. Addi- tional distributional records of bats in Utah. J. Manmial. .36:126-127. 'U.S. Bureau of Land Management, Lewistown District Office. Lewistown, Montana 594.57. 'Montana Department of Fish, Wildlife, and Parks, Montana State University, Box 5, Bozeman, Montana 59717 115 A CHIRACANTHIUM SPIDER BITE Dorald M. Allied' .\bstr.\ct.— a bite bv Cliimcanthium mildci L. Koch is described. In May 1979, Mr. Lee Carson of Provo, Utah, brought a spider to me which had bit- ten him on the right index finger. It was sub- sequently identified by Dr. Wilhs J. Gertsch as a female Chiracanthium niildei L. Koch. Mr. Carson was in the process of placing a pair of Ribbers over his shoes to work in his garden. The mbbers were kept in his closed garage, and had been worn a few days pre- viously. As he inserted his fingers into one of the rubbers, he felt a sudden pain at the tip of his finger. Examination disclosed the spi- der with a web and cocoon in a rubber. Within 3 to 4 seconds after the bite, the finger began to ache severely. The pain soon extended to his upper arm where it remained for 2 to 3 hours, although it was most con- centrated in the finger. He described the sen- sation as a painful "tingling" similar to what one experiences when his elbow "crazy- bone" is bumped. No nausea, headache. swelling, or numbness was experienced. No evidence of inflammation or necrosis oc- curred at the site of the bite, and the cheli- ceral punctures healed rapidly. Pain in the finger and arm disappeared after about 4 hours. Dr. Gertsch kindly supplied information on the spider and its ecology. The species appar- ently was introduced into the United States, became a typical house spider in the New York and Boston area, and subsequently spread to other areas of the U.S. As early as 1930 Dr. Gertsch collected it at Salina, Utah. It is known from southern California, and Kaston (1972, "How to Know the Spiders, p. 221) indicated its distribution as New Eng- land, New York, New Jersey, Alabama, Mis- souri, and Utah. Gertsch further stated that the genus is reputed to have a venom of in- termediate potence, and that of the Eu- ropean species is said to be next to their Lat- rodectus in severity. 'Department of Zoology, Brigham Young University, Provo, Utah 84602. 116 IDENTITY OF NARROW-LEAVED CHRYSOTHAMNUS VISCIDIFLOR US (ASTERACEAE)' Loraii ('. Aiidersoir .\bstr.\ct.— Two ijroups of glabrous, narrow-leaved Chiysotliaiiinits visciclifloius (Asteraceae) are perceived, and appropriate taxononiie combinations are made, i.e., C. v. ssp. vmidiflorus var. stcnofjlttjlhis and C. v. ssp. axillaris. The two are fairl\' distinct ideograph icallv, and thev can be separated l)v floral niorpliologv. .\ key to all species of section Chrtjsothanntus (to which C. liscidiflorus belongs) is given. Classification of intra.specific variants of Chrysothamnus liscidiflorus has been prob- lematic, in part, becau.se floral features seem- ingly lacked sufficient distinctions. Hence, vegetative aspects such as stature, vesture, and leaf dimensions have been used. Experi- mental -Studies (Anderson 1964) have demon- strated that in many in.stances plant height, leaf twisting, and leaf width are expressions of differing edaphic conditions, droughtiness, or other environmental parameters and thereby complicate taxonomic resolution. Study of interpopulational variation and the distribution of the narrow-leaved, yellow rabbitbmsh (C. viscidiflorus ssp. stenophyUus) reveals that there are two different taxa rep- resented. One is found sporadically through the northern latitudes of the western United States and into southern California (open cir- cles in Fig. 1). The other taxon (stars. Fig. 1) is found further .south and is more generally distributed, i.e., has a more pronounced "range integrity." In south-central Nevada, Beatley (1976) reports it is the common Chrysothamnus of basin floors and foothills, especially in volcanic areas and on disturbed sites, usually below 5500 ft. Examination of populations at possible type locality sites (type specimens labeled either West Hum- boldt Mountains or Huntington Valley) for ssp. StenophyUus shows that the narrow- leaved plants represent extremes of the broader-leaved ssp. viscidiflorus. I have concluded from field observations, garden culture, and herbarium studies that the northern elements, which include the tvpe collection of C. i". ssp. stenophyUus, are actually environmentally induced variants of ssp. liscidiflorus. Although quadrinomials are cumbersome, the following nomenclatural combination more appropriately identifies the relationship of these plants: Chrysothamnus viscidiflorus (Hook.) Nutt. ssp. viscidiflorus var. stenophyUus (Gray) L. C. Anderson, comb. nov. Basionym: Bigelovia douglasii Gray var. stenophylla Gray. Proc. Am. Acad. Sci. 8:646, 1873. W. ' Humboldt Mtns, Nevada, Watson 566 (GH, holotype; NY, US, iso- types). Synonymy: Chrysothamnus pumilus Nutt. var. varus A. Nels. Bot. Gaz. 28: 375, 1899. Centennial Valley, Wyoming, Nelson 1847 (RM, holotype; GH, NY, i.sotypes). Tlie .southern elements that had previously been referred to ssp. stenophyUus warrant subspecific recognition. Their narrow-leaved characteristic is independent of environmen- tal conditions. These plants are diploids; broad-leaved forms of ssp. viscidiflorus that grow in the same region are tetraploids or hexaploids (Anderson, 1966, 1971). The only available name for these narrow-leaved plants is C. axillaris. In 1964, I noted that C. axillaris was not specifically distinct from C. viscidiflorus, and the name was synonomized luider .ssp. stenophyUus. Munz (1968), in re- 'This shidy was supported by .National Science Foundation Grant DEB 76-10768. Ken VVonilile helped with ijraphics. •Department of Biological Science, Florida Stale University, Tallahassee, Florida 32306. 117 118 Great Basin Naturalist Vol. 40, No. 2 ferring to my studies, stated I had made C. uxillaris a suljspecies of C. viscidiflonis. The statement was inaccurate at that time— but prophetic. The combination is now made: Chrysothamnus viscidiflonis (Hook.) Nutt. ssp. axillaris (Keck) L. C. Anderson, comb. & Stat. nov. Basionvm: Chri/sothamntis axiUaris Keck. Ahso 4:103, 1958.' Deep Springs Valley, Cali- fornia, Ferris 6924 (NY, holotype; DS, LL, POM, isotypes). Keck (1958), in describing C. axiUaris, re- lated it to C. albidus and more distantly to C. greenei. Actually, ssp. axillaris is not close to C. albidus in relationship, but it is to C. greenei. In fact, the type collection of ssp. ax- illaris with very acute phyllaries suggests some intergradation with C. greenei. Keck stated that the latter was known only from eastern Nevada and eastward, but it does ex- tend through southern Nevada into Inyo County, California. At its western limit in California and also in northeastern Arizona, C. greenei intergrades somewhat with C. vis- cidiflorus. The feature of vertically aligned phyllaries noted by Keck (1958) is not con- sistent for ssp. axillaris (Anderson 1964). Although ssp. axillaris and var. steno- plnjllus are fairly distinctive habitally, they could not be "keyed out" easily vinless refer- ence was made to geographic distribution (Fig. 1). Floral morphology was studied in search of additional distinguishing features. Methods are those used earlier (Anderson 1964). Detailed floral data and a list of speci- mens examined are on file at FSU. Significant comparisons are graphically illustrated in Figure 2. Involucral width in Chrysothamnus is gen- erally strongly correlated with flower num- ber because more flowers per head require a broader receptacle. The pattern is evident in general range for C. v. sep. viscidiflorus C. V. ssp. viscidiflorus var. stenophyllus C. V. ssp. axillaris Fig. 1. Range of Clirysolhamnu.^ viscidiflonis ssp. viscidiflonis and ssp. axillahs. Range for ssp. viscidiflonis nearly equals that of the species; ssp. tanceolatus extends into southern British Columbia and north-central New Mexico. Distribution of ssp. (ixillaiis (stars) is fairly general through southern parts of Utah and Nevada and adjacent regions, whereas that of .ssp. viscidiflonis var. stenophijllus (open circles) is sporadic, but mostly north of ssp. axillaris. June 1980 Anderson: Chrysothamnus 119 C. viscidiflorus ssp. viscidiflorus, wherein plants averaging 10.8 (with up to 12) flowers per head have involucral widths over 50 per- cent of involucral length (Fig. 2); these plants represent an altitudinal record for the genus of 12,800 ft in the White Mountains of Cali- fornia. Previous descriptions of C. t/,s- cidiflonis listed flower number as about 5 (Hall and Clements 1923). Nearly all plants of ssp. viscidiflorus with high flower number come from altitudes over 10,000 ft. Most populations of the subspecies, including var. stenophyUus, have 4-6(7) flowers per head with proportionately narrower involucres. Heads of ssp. axillaris depart from the bas- ic correlation of flower number— involucral width /length ratio. Although they average fewer than 5 flowers per head, the width /length ratio for the involucre is high (Fig. 2). Tlius, ssp. axilluris and var. steno- phijllus can be distinguished by floral fea- tures as well as geographically. Taxonomic interpretation of Chryso- thamnus section Chrysothamnus (to which C. viscidiflorus belongs) has been altered consid- erably since Hall's monograph (Hall and Cle- ments 1923). Chrysothamnus vaseyi and C. molestus (C. viscidiflorus var. tnolestus) have been transferred to section Fulchelli (Ander- son 1970) and C. gramineus to Petradoria (Anderson 1963). Additional species have been recognized in the section. A key to sec- tion Chrysothamnus as currently understood is presented here. -/At- Fig. 2. Correlation of involucral shape and flower number in heads of Cltrysothamntis viscidiflorus ssp. vis- cidiflorus (closed circles, var. viscidiflorus; open circles, var. stcnoplnjilus) and ssp. axillaris (stars). Vertical axis represents the involucral width /length relationship ex- pressed as percent; the horizontal axis is average flower number per head. Note that ssp. axillaris departs from the general correlation in ssp. viscidiflorus and other members of the genus. 1. Flowers white; leaves terete, strongly punctate C. albidus (Jones) Greene — Flowers yellow; leaves planate or involute, not punctate 2 2(1). Flowers 2-3(4); style branches included in erect corolla lobes; plants mostly less than 1.5 dm tall C. hurnilis Greene — Flowers (3)4-6; styles exerted beyond spreading corolla lobes; plants often over 2 dm tall 3 3(2). Style appendages long (40-70 percent of style branch); leaves never twisted or involute; tall shRibs 4 — Style appendages short (30-45 percent of style branch); leaves frequently twisted or involute 5 4(3). Leaves lanceolate; achenes densely pubescent C. linifolius Greene — Leaves spatulate to oblanceolate; achenes sparsely pubescent C. spathulatus L.C.Anders. 5(3). Phyllaries acuminate-cuspidate; leaves 1-2 mm wide C. greenei (Gray) Greene — Phyllaries obtuse to acute; leaves 1-10 mm wide (C. viscidiflorus) 6 120 6(5). 7(6). 8(7). 9(7). 10(9). Great Basin Naturalist Vol. 40, No. 2 Leaves planate, glabrous; flowers 3.5-4(4.5) mm long C. V. ssp. planifolius L.C.Anders. Leaves twisted or pubescent or flowers longer 7 Upper stems, frequently leaves, hairy 8 Stems glabrous; leaves only ciliate 9 Stems and leaves pubenilent; leaves 1-2(4) mm wide C. V. ssp. puberiilus (D.C.Eat.) H.&C. Stems hispid near inflorescence; leaves over 2 mm wide, hirsute or glabrous C. t;. ssp. lanceolatiis (Nutt.) H.&C. Leaves ± 1 mm wide; flowers 3-4(5); involucres somewhat turbinate C. r. ssp. axillaris (Keck) L.C.Anders. Leaves 1-10 mm wide; if 1 mm, flowers 4 or more and involucres narrowly cylindric 10 Leaves more than 1.5 mm wide; plants up to 1 m tall C. V. ssp. viscidiflorus var. viscidiflorus Leaves 1-1.5 mm wide; plants mostly less than 3 dm tall C. V. ssp. viscidiflorus var. stenophyllus (Gray) L.C.Anders. Names often applied in C. viscidiflorus, but not representative as distinct subspecies, include: (1) elegans, usually misapplied to certain forms of ssp. puberulus with bracts with enlarged green tips, but the type speci- men does not have such bracts and is clearly part of ssp. lanceolatus; (2) pumilus, low form that is part of ssp. viscidiflorus; (3) torti- folius, environmental variant with strongly twisted leaves, part of ssp. viscidiflorus; and (4) latifolius, wide-leaved plants from north- ern Nevada that could be considered a varie- ty of ssp. viscidiflorus; however, not all wide- leaved plants of the subspecies would belong to that variety. A conceptual distinction between sub- species and variety exists in Chnjsothamnus for me. Subspecies is applied to groups of populations with pronounced geographical and fairly distinct morphological limits. The variety can be applied in two ways. It may represent a sporadic but rather distinctive morphotype within a given subspecies, such as ssp. viscidiflorus var. stenophijllus, and it could possibly be applied to such variation in species where subspecies are not recognized, such as C. grcenci var. filifolius for the nar- row-leaved variant. The second application (my preferred usage) of variety would be for elements of a subspecies that have some fair- ly consistent morphological distinction and also have relatively sharp geographic limits. but are clearly subordinate to the subspecies. An example would be ssp. viscidiflorus var. latifolius— a the combination were made. Publication of additional quadrinomials as needed to clarify relationships in the genus will be part of my upcoming monograph. Literature Cited Anderson, L. C. 1963. Studies on Pctradoria (Conipos- itae): Anatoniv, cvtologv, taxonoiiiv. Trans. Kans. Acad. Sci. 66:632-684."' 1964. Taxononiic notes on the CIinjsotlKininus viscicliflonis complex (Astereae, Conipositae). Madrono 17:222-227. 1966. Cytotaxonomic studies in Clinisothiimnus (Astereae, Conipositae). Amer. J. Bot. 53:204-212. 1970. Floral anatomy of Chrysotli(iiuniis (As- tereae, Conipositae). Sida 3:466-503. 1971. Additional chromosome counts in Chnjso- tlunnntts (Asteraceae). Bull. Torrev Bot. Club 98:222-225. Be.\tley, J. C. 1976. Vascular plants of the Nevada Test Site and central-southern Nevada. Energy Res. & Dev. ,\dmin., Nat'l Tech. Inf. Service, Springfield, Va. Hall, H. M., .\nu F. E. Clements. 1923. The phyloge- netic method in taxonomy: The North American species of Artemisia, Chnjsoihamnus, and Atri- plcx. Carnegie Inst. Publ. 326:l-.355. K.ec:k, D. D. 1958. Taxononiic notes on the California flora. Aliso 4:101-114. Munz, p. a. 1968. Supplement to A California Flora. I'niv. Calif. Press, Berkeley. RIBULOSE DIPHOSPHATE CARBOXYLASE ACTIVITIES IN COLD-RESISTANT COMMON MALLOW, MALVA NEGLECTA WALLR. AND A COLD-SENSITIVE TOMATO, LYCOPERSICON ESCULENTUM L., ACE 55 VAR. W illiam H. AiidfiSL-ii' and Jack D. Brotlicrson' Abstfl^ct.— Coiiinioii mallow (Malta nc^hcta W'allr. ) and tomato (Li/copcisicon csciiUnttim L. var. Ace 55) were compared as to certain characteristics: C02 fixation properties, ribulose diphosphate carboxyl activities, (RiiDPCase) photosynthesis, respiration, and compensation points. Significant differences in these factors were observed in all cases except dark respiration. Mallow enzyme (RuDPCase) activities were higher per unit of enzyme than those of tomato. The Mallow RuDPCase exhibited slightly higher activity at 5 to 25 C. Mallow leaves retained their capacity for photosynthesis and respiration after long periods of exposure to subfreezing temperature. The cold adapted mal- low had a higher C(% compensation point, suggesting a lower efficiency for CO2 fixation. The results suggest that cold acclimation in common mallow affects photosynthesis but has little effect on respiration. Several physiological factors are associated with the development of resistance to winter injury in plants. Qualitative and quantitative changes in protein, carbohydrate, and lipid contents have been observed during cold ac- climation (Roberts 1969, Cerloff et" al. 1967, Hochachka and Somaro 1968, Zeller 1951). However, the in vivo features of observed biochemical and physiological alterations as- sociated with cold acclimation in specific in- stances are not clear. In particular, very little is known about the intracellular mechanisms of freezing resistance in broad-leaved plants that remain conspicuously green and meta- bolically active throughout the winter months of cool temperature regions. Common mallow, Malva neglecta Wallr., is an example of a broad-leaved plant that of- ten remains green and succulent throughout the winter in north-temperature regions. Its green leaves can tolerate subfreezing temper- atures without visible evidence of injury. It appears that common mallow is capable of surviving winter cold by some mechanism other than dormancy, because the plant re- tains the capacity for photosynthesis and rel- atively high respiration rates when favorable conditions are present. These observations have prompted an in- vestigation of certain photosynthetic charac- teristics and CO2 fixation properties in win- ter-hardened mallow. This paper reports the activity of purified ribulose diphosphate car- boxylase (RuDPCa.se) and the capacity of whole leaves to fix COo from cold-accli- mated, field-grown mallow and from green- house-grown mallow and tomato. Materials and Methods Plant nuitcriah: Common mallow is a per- ennial weed characteristic of cultivated ground, gardens, yards, and waste places throughout the United States. Introduced from Eiuope, the weed belongs to the same plant family as cotton, hollyhocks, rose of sharon, and the weeds known as velvet-leaf and flower-of-the-hour. This family (Mal- vaceae) has flowers which contain a tube of stamens siurounding the pistil and a ring of seeds centered in persistent floral parts remi- niscent of a small flat cheese (thus one of the plant's common names, "cheese weed"). The plant's long tap root and its wide distribution in relation to habitat occupation indicates a wide ecological amplitude in regard to envi- ronmental stress factors. The garden tomato, Lycopersicon esculen- tiim L., variety Ace 55, cannot tolerate sub- freezing temperatiues. Tomatoes are warm 'Department of Botany and Range Science, Bni;liam Young University. Provo. Utah 84602. 121 122 Great Basin Naturalist Vol. 40, No. 2 season plants; the Ace 55 variety yields very well under high day and cool night temper- ature regimes. Method of sampling plants for measure- ment of photosynthesis and respiration rates: Plants of mallow and tomato were grown in the greenhouse at 25 C (76 F) day and 20 C (68 F) night temperatures. Mallow plants were also grown in the field near Provo, where they were exposed to subfreezing tem- peratures. Plant samples were taken from greenhouse and field areas during January. When harvested, plants were collected whole, petiole ends were cut under water, and then they were placed in controlled envi- ronment chambers with the cut ends remain- ing immersed in water. Only deep green suc- culent growth was harvested. Photosynthetic and respiration measurements were then made repeatedly as described below. COo fixation methods: Rates of net photo- synthesis (APS), dark respiration (DR), and CP2 compensation points (CP) were deter- mined in mid-January on excised shoots cut under water. Analysis was made utilizing a Beckman IR-215 infrared gas analyzer and a plexiglas controlled-environment chamber. Apparent photosynthesis (APS) was deter- mined by the time required for the closed system's CO2 content to decrease from 315 to 275 jul per 1 of air. Dark respiration (DR) was determined by the time for the closed sys- tem's CO2 content to return from 275 to 315 jtxl per 1. The CO2 compensation point (CP) was determined in a closed system by allow- ing the plants to fix CO2 until no further change in CO2 concentration occurred in the atmosphere of the lighted plexiglas chamber surrounding the plant. The assimilation chamber was housed inside a large growth chamber with lighting provided by 8 cool- white inforescent tubes, 8 grolux (Sylvania) inforescent tubes, and 10 25-watt in- candescent globes. The light was filtered through 4 cm of water and provided an in- tensity of 6.05 X 104 ergs per cm per minute at leaf height. This light intensity has been indicated to be saturating for tomato at 315 jul CO2 per 1 of air. Chamber parameters were: temperature -23 ±.6 C; relative hu- midity— 65 ±10 percent; wind speed— 4.0 dm per minute (3 chamber volumes per minute) (Brewster 1971). Preparation of extracts from leaf homoge- nates and enzyme purification: Fully ex- panded leaves were washed and their midribs removed and blotted dry. From this step on, all procedures were carried out at 5 C. Ap- proximately 3.0 gm samples of leaf tissue were groimd manually with cold mortar and pestle for 5 minutes, using 5 ml of 0.1 M HEPES (N-2-hvdroxvethvlpiperzaine-N-2- ethanesulfonic acid) buffer pH 8.00, 0.001 M EDTA, 0.0001 M DTT, 0.01 M MgClai, 0.025 mM NaHC03, per gram fresh weight leaf tis- sue. The homgenates were centrifuged for 10 minutes at 20,000 rpm in a Sorvall model RC-2B centrifuge with the S-34 rotor. The supernatant was decanted and used as the crude enzyme extract. The crude enzyme ex- tract from the low speed centrifugation was further clarified by centrifugation at 40,000 rpm in a Spinco model L3-50 for 5 minutes. The supernatant fraction was collected. The RuDPCase enzyme was purified further by sedimentation of the extract into a sucrose step gradient consisting of 2 ml of 5 percent, 2 ml 30 percent and 3 ml 50 percent sucrose solution in HEPES buffer layered in a centri- fuge tube. The rapidly sedimenting RuD- PCase accumvilated in the 50 percent sucrose layer after sedimentation for 12 hours at 25,000 rpm in a Spinco SW-25 rotor. The su- crose-enzyme solution was then passed through a 10 X 1.0 cm Sephadex G-25 col- umn for further purification. Measurement of enzyme activities: Deter- mination of RuDPCase activity was based upon fixation of ^^COa. into acid stable prod- ucts. The assay mixture contained 0.01 M HEPES-S04 buffer (pH 8.00) 0.01 M MgC12, 0.001 M DTT, 0.02 M NaHi^COa, and ribu- lose-1, 5-bisphosphate in 200 [i\. The enzyme (30jul) was added to initiate the reaction and was allowed to proceed for 10 minutes. The reaction rates were linear over this time peri- od. The enzyme reaction was stopped by the addition of 50 ju.1 of glacial acetic acid. A 100 jul aliquot of the reaction mixture was spotted onto a strip of Whatman No. 1 filter paper and dried under the hood. The sample was counted in a Packard Tri-Carb (Model 3320) liquid scintillation counter in toluene scintil- lation fluid. The counted samples were cor- rected for machine efficiency and quenching June 1980 Andersen, Brotherson: Cold Acclimation 123 and the values converted to disintegrations per minute (dpm). Determination of specific activity of RuD- PCase: RuDPCase has been identified with a large, rapidly moving boundary observed in the Spinco Model E ultracentrifuge known as fraction I protein. The area of the Schlieren boundary curve corresponding to fraction I protein provides a means to determine the relative concentration of RuDPCase present. An estimate of specific activity per unit of enzyme can then be calculated by comparing enzvme activitv in a given extract with the area of the corresponding fraction I protein peak (Andersen et al. 1970). Results Net photosynthesis, dark respiration, CO2 compensation point: Table 1 shows that cold- adapted, field-grown mallow exliibited lower rates of APS than greenliouse-grown mallow or tomato. However, rates of DR were sim- ilar in all cases. This resulted in an APS to DR ratio for field-grown mallow of one-half that for the greenhouse-grown plants. The CP shows a significant increase for field- grown Mallow over values for the green- house-grown plants. Reaction velocities for carbonate and ri- hu\osedip}\osphate substrates: The com- parative reaction velocities for purified "cold-adapted" mallow and tomato RuD- PCase at different carbonate substrate con- centrations are depicted in Figure 1. The ri- bulosediphosphate (RuDP) substrate was maintained at maximum concentration for both enzymes. The mallow RuDPCase exliib- ited higher catalytic capacity per unit of en- zyme than tomato RuDPCase. The turnover number at V.^^, (4 u moles of carbonate sub- strate per 200 ul of reaction mixture) for to- mato (RuDPCase) was calculated at 1036 moles carbonate fixed per mole of enzvme per minute. The turnover number for puri- 2 V 10 i 8 < 6 O •MALLOW •■TOMATO Ofl 1£ 2.4 32 40 4B 56 64 CONCENTRATION HCO," (>i MOLES) Fig. 1. Dependence of RuDPCase activity in pmified extracts upon HC0.3 concentrations. RuDPCase was pu- rified from common mallow and tomato ACE 55 var. The enzyme activities are based upon total amount of RuDPCase enzvme present in the reaction mixture as calculated from the Schlieren curve of the sedimenting boundaries in a model E ultracentrifin'e. T.\BLE 1. Rates of net photosynthesis, dark respiration, and the CO2 compensation point of excised plant shoots. See text for description of plant treatments. Species Location .\pparent photosynthesis (ugCo2'dnr^*min"^ ) Mallow Mallow Tomato (ACE 55) Greenhouse Field Greenhouse 3.2 1.5 2.6 Dark respiration (ugCo2'dnr^Mnin"') L7 1.7 1.3 CC^ compensation point (uI.LM 64 86 68 *.\pparent photosynthesis {.\?S) was measured bv determining time required to lower closed system COj concentration from 320 ul/I to 280 ul/1 air-1. ^'Dark respiration' (DR) was measured by determining the time required for a darkened closed system to return COj concentration from 280 to 320 ul/1 air. '^Compensation point was measured by allowing photosynthesizing plants to fix CO2 from a closed atmosphere until no further change in CO2 concentra- tion could be obser\ ed. 124 Great Basin Naturalist Vol. 40, No. 2 fied mallow RuDPCase was calculated at 1400 moles of carbonate fixed per mole en- zyme per minute. The differences in V,„^^ val- ues for the two enzymes were judged to be highly significant, based on the student-t test for measuring differences between paired variates. The calculated t value exceeded the 0.001 level of significance. Only slight differ- ences between the corresponding Km values could be observed. The reaction velocities of mallow and to- mato RuDPCase for different ribulosediphos- phate substrate concentrations are graphed in Figure 2. The shapes of the reaction velocity curves for tomato and Mallow RuDPCse are similar. The mallow enzyme exhibited a sig- nificantly higher V^,^, value. Both enzymes showed substrate inhibition at RuDP substate concentrations higher than 8 u moles per 200 jLil of reaction mixture. The significance of the differences between the two reaction ve- locity cmves in Figure 2 was measured by the student-t test for paired variates. The cal- culated t value exceeded the 0.01 level of sig- nificance. The Km values for substrate con- centration at half maximal velocity were slightly higher for tomato. Effect of temperature on reaction velocity with purified RuDPCase enzyme: Purified to- mato and mallow RuDPCase enzyme extracts were compared for catalytic velocities at re- action temperatures ranging from 4 C to 63 C (Figure 3). Purified mallow RuDPCase had significantly higher catalytic activity per unit of enzyme under the temperatine range of 4 to 25 C. The calculated student-t value for differences between paired variates exceeded the 0001 level of significance. On the other hand, purified tomato enzyme showed signifi- cantly higher catalytic capacity in the 38 to 53 C temperature range. The corresponding calculated student-t value for paired reaction rates in the 38 to 53 C temperature range ex- ceeded the 0.01 significance level. The gener- al shapes of the temperature curves for the purified mallow and tomato RuDPCase were quite similar, however, with heat denatura- tion for both enzymes occurring near 53 C. Discussion Our studies indicate that respiration and photosynthesis are differentially effected by COMCCNTRATION OF RuOP (ji MOLES) Fig. 2. Dependence of RuDPCase activity in purified extracts from mallow and tomato ACE 55 var. upon RuDP concentrations. Enzyme activities are normalized for equal concentrations of RuDPCase. 20 25 30 35 40 45 50 55 60 65 TEMPERATURE °C Fig. 3. Dependence of RuDPCase activity in purified extracts from mallow and tomato ACE 55 var. upon temperatine. Enzyme activities are normalized for equal concentrations of RuDPCase. June 1980 Andersen, Brotherson: Cold Acclimation 125 cold acclimation in mallow. The measured rates of apparent photosynthesis values in greenhouse-thrown mallow were approx- imately twice the measured rate of apparent photosynthesis values of cold-acclimated, field-grown mallow. No differences were ob- served for rates of dark respiration (Table 1). Higher COo compensation points were ob- .served for field-grown, cold-adapted mallow, which suggested a depressed efficiency for CO2 fi.xation. However, because respiration and photosynthesis measurements were made at 25 C in the laboratory, it is possible that relative efficiencies of carbohydrate accumu- lation would change at lower temperatures. The cold-acclimated mallow might under such circumstances become relatively more efficient. Present evidence, however, in- dicates that cold acclimation in field-grown mallow is a matter of maintaining a steady state of metabolic activity rather than the rapid acciunulation of carbohydrate reserves. Further studies are underway to assess inter- action of lower temperatures and carbohy- drate accumulation in cold-acclimated mal- low. Purified RuDPCase from cold-adapted Malva neglecta had the highest catalytic ca- pacity per unit of enzyme. The Vmax values for carbonate and ribulosediphosphate sub- strates were highest for mallow RuDPCase (Figs. 2 and 3). The Km values for tomato were only slightly higher. The Km and Vmax values for tomato RuDPCase agree in general with corresponding reported values (Ander- sen et al. 1970). Although we cannot yet compare in vitro RuDPCase activity to in vivo CO2 fixation without some misgivings, our studies suggest that mallow RuDPCa.se would promote slightly more rapid CO2 fix- ation per unit of enzyme in vivo. The lower Km value for CO2 substrate of RuDPCase from the cold-adapted mallow would suggest a higher photosynthetic efficiency for the in- tact plant. Yet the higher compensation point of these cold-adapted plants indicates that photosynthetic efficiency is depressed in the intact leaf. Recent work by several in- vestigators clearly implicates RuDPCa.se as a major contributing factor to the high com- pensation points of the C3 species (Ogren and Hunt 1978). Our study would indicate that the higher compensation point in the cold-adapted mallow is due to some other factor in the photosynthetic carbon cycle than RuDPCase. On the other hand, since we are using purified enzyme for our studies, it is likely that control molecules that may af- fect Km for CO2 fixation of RuDPCa.se could be removed in our purification process. In any case, if RuDPCase has a higher Km for CO2 in vitro, which would result in a higher compensation point in the cold-adapted mal- low, the effect does not persist through puri- fication of the enzyme. Therefore, at least a change has not been detected on the purified enzyme that would affect the compensation point and thus be a basis for lower photo- synthetic efficiency durmg cold acclimation. The mallow RuDPCase enzyme showed higher catalytic capacity than tomato RuD- PCase under temperature ranges of 0-25 C, and the tomato RuDPCase enzyme exhibited higher activity under temperature ranges 40-60 C. This may be indicative of Mallow's lower-temperature environmental adaptation and its COo-fixing enzymes. Other studies have shown that RuDPCase extracts from plants of different climatic regions exliibit correspondingly different temperature reac- tions (Trihame and Cooper 1969). Also, we have observed that the RuDPCase activities in ciiide extracts from Mallow were con- sistently higher than RuDPCase activities in tomato crude extracts, (on a per-gram fresh weight basis). These results, along with the distribution patterns of these two species, suggest that the temperature interaction of the enzyme might be related in some way to the different seasonal adaptations of the two species. The results also suggest that the process of cold acclimation in mallow affects photo- synthesis and dark respiration differently. Respiration was not seriously affected, but photosynthetic capacity per unit of leaf area and photosynthetic efficiency were signifi- cantly reduced (Table 1). It may be hypoth- esized then that the processes of cold accli- mation in mallow either depresses the in vivo activity of RuDPCase or alters in some way other chloroplast functions which affect the plant's capacity for photosynthesis. 126 Great Basin Naturalist Vol. 40, No. 2 Acknowledgment This research was supported in part by the National Institute of Health, Grant GM 17868-02. Literature Cited Andersen, W. R., G. F. Wildner, and R. S. Griddle. 1970. Ribiilose diphosphate carboxylase from mu- tant tomato plants. .\rch. Biochem. Biophys. 137:84-90. Brewster, S. F. 1971. The physiological vitality of scar- let globemallow, Sphaeralcea gwssuhiriaefolia (Hook. & .\RN.) Rydberg, under drought. Unpub- lished dissertation, Brigham Young University, Prove, Utah. Gerloff, E. D., M. a. Stahmann, and D. Smith. 1967. Soluble proteins in alfalfa roots as related to cold hardiness. Plant Physiol. 42:895-899. Hochachka, P. W., and G. N. Somero. 1968. Adapta- tions of enzymes to temperatures. Comp. Bio- chem. Physiol. 27:659-668. Ogren, W. L., and L. D. Hunt. 1978. Gomparative bio- chemistry of ribulose-bisphosphate carbo.xylase n higher plants. Pages 127-1.39 in Siegelman and Hind, eds., Photosynthetic carbon assimilation. Basic Life Sciences 11. Roberts, D. W. A. 1969. A comparison of isozymes of wheat plants grown at 6 G and 20 G. Ganadian J. Bot. 47:263-265. Treharne, K. J., AND J. P. GooPER. 1969. Effect of tem- perature on the activity of carboxylase in tropical and temperature graminae. J. Exp. Bot. 20:170-175. Zeller, O. 1951. Respiration at low temperatures in cold hardened cereals. Planta. .39:.500. RECOVERY OF GAMBEL OAK AFTER FIRE IN CENTRAL UTAH L. M. Kiinzler' and K. T. Harper' .\bstr.\ct.— The height of oak (Qucrcus gdinbelii Nutt.) stems was measured on several fire scars within the Uinta National Forest and vicinity and compared with the height of oak stems on adjacent, nonhumed areas. A significant relationship exists between the recovery rate of oak after fire and elevation, with the recovery rate being greatest at low elevations. \ trend also exists showing that recoverv tends to be greater on south to westerly exposvires than on north to easterly exposures. Gainbel oak {Qucrcus gambelii Nutt.) is an important species of the deer winter ranges of central Utah, providing both food and cov- er for deer (Alhiian 1952, Smith 1949). How- ever, because of its growth habit, it often forms impenetrable thickets (Allman 1952, Baker 1949, Dills 1970, Marquiss 1972, McKell 1950). This, coupled with its height, places most of the available browse out of reach of big game (Plummer et al. 1966, 1970). By treating these dense stands of oak with chemical herbicides, fire, or machinery to break them up, the stands can be opened up and made available to browsing animals (Anon. 1966, Dills 1970, Hallisey et al. 1976, Marquiss 1971, 1972, Plummer et al. 1966, 1970, Price 1938). Because oak stands recov- er rather rapidly after these treatments, it is necessary to determine a rotation period for treatment to maintain an optimum amount of browse for wildlife (McKell 1950, Plummer et al. 1966, 1970). Literature Review In central Utah, Gambel oak has been re- ported to occur in almost pure stands from 5000 (1525 m) to 8000 (2440 m) feet eleva- tion along the Wasatch Range (Allman 1952, Baker 1949, McKell 1950). This area con- stitutes a large portion of the deer winter range in the area (Allman 1952, Anon. 1966, Dills 1970, Hallisey et al. 1976, Plummer et al. 1966, 1970, Smith 1949). Treatments of oak using fire, herbicides, or machinery to destroy the oak canopy result in prolific sprouting, with several stems re- placing each preexisting stem. Impenetrable thickets often result from such treatments (Allman 1952, Baker 1949, Dills 1970, Mar- quiss 1972, McKell 1950). Yet, treatments can be effective in improving deer range. When a follow-up program is used, such as seeding with competitive herbs and grasses, the ben- efits of the treatment can be prolonged for over 15 years (Anon. 1966, Dills 1970, Hal- lisey et al. 1976, Marquiss 1971, 1972, Plum- mer et al. 1966, 1970, Price 1938). By treat- ing oak, deer use can be increased up to four times, but deer use declines as the time from treatment increases (Anon. 1966, Hallisey et al. 1976, Price 1938). Methods The height of oak stems was measured on several stands within the Uinta National For- est and vicinity. One half of these stands were located in oak stands that had burned 3 to 15 years ago. The other half of the stands were located in nonburned areas adjacent to each burned stand considered. Unburned stands were selected so as to have the same slope, exposure, and elevation as the burned stand that each was paired with. Mea.sure- ments were taken along a 100-foot transect at 16-foot intervals, with the oak stem that was nearest to the point being measured. Slope varied from 20 to 70 percent and elevation ranged from 5100 feet (1555 m) to 6800 feet 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 127 128 Great Basin Naturalist Vol. 40, No. 2 (2070 m) in elevation among the stand pairs considered. Exposure also varied, with one- half of the stands having a south to west ex- posure and the other half having a north to east exposure. A percent recovery value was calculated for each pair by dividing the aver- age height of the oak in the fire scar by the average height of the oak in the nonburned area. A recovery rate was then calculated by dividing the percent recovery value by the age of the fire scar (Table 1). To minimize the variation among recovery rates caused by the nuisance factors of age, elevation, slope, and site, recovery rates were also calculated on a uniform hill within the Wallsburg Burn (Table 2). Three transects were placed in the Ijumed area at 500-foot (152 m) intervals in elevation. Results In comparing the recovery rates from the stands throughout the Uinta National Forest and vicinity, a significant correlation (power equation) exists between elevation and oak recovery rates (r = -.85, .01 < P < .05, Fig. 1). Using data collected from the Wallsburg Burn, a similar analysis confirms that a signif- icant correlation (linear equation) exists be- tween elevation and the recoverv rates of oak stems (r = .99, P <.01, Fig. 2)'. Both sets of data show that, as elevation is increased, the recovery rates decrease. Although there is in- dication that stands on south to west expo- sures have faster recovery rates, the differ- ence between the recovery rates of these stands and those with north to east exposures is not significant (Fig. 3). Discussion Comparing the results of the two sets of data, it seems clear that elevation has a strong influence on oak's rate of recovery fol- lowing fire. The variation between recoverv rates in Figure 2 is primarily due to elevation Table 1. Annual recoverv rates in percent for oak on burns scattered throuij;hout the Uinta National Forest and nearby areas. Average Average height of Age of height of oak in non- Recoverv Stand Elevation burn oak on burn burned area Percent rate # (m) Exposure (years) (cm) (cm) recovery (percent) 1 1,740 East 3 77 579 13.3 4.4 2 1,800 North 15 140 320 43.7 2.9 3 1,560 South 3 145 274 52.8 17.6 4 1,770 South 4 83 343 24.2 6,1 5 1,650 East 3 84 290 28.9 9.6 6 1,680 West 3 61 135 45.0 15.0 7 1,680 East 3 83 290 28.5 9.5 8 1.770 West 4 88 399 22.1 5.5 9 1,860 East 6 116 533 21.7 3.6 10 2,070 West 4 37 320 11.4 2.9 T.\BLE 2. Percent recovery values for oak on a unih)rni hill in the Wallsburg Burn. This stand had burned four years earlier. Transect 1 1-a 2 2-a 3 3-a Elevation Status Average height Percent recovery 1,770 m Burned 88.4 cm 22.2 1,770 m Unburned 398.8 cm 1,920 m Burned 59.9 cm 16.8 1,920 m Unburned 356.6 cm 2.070 m Burned ■36.6 cm 11.4 2,070 m Unburned 320.0 cm June 1980 KuNZLER, Harper: (jAmbel Oak 129 alone. Even with the nuisance variables men- tioned earlier, it will he noted that elevation is a significant factor for recovery rates of oak (Fig. 1). Possible reasons for the change in oak re- coverv rates with elevation include the fol- lowing: (1) the species is approaching its up- per elevational limit on some of the burns and, because of this, its growth may be slow- er; (2) more moisture and nutrients may be available to plants at the bottom of slopes be- cause of precipitation's surface nuioff and at- tendant erosion, nutrient transport, and re- sultant differences in soil depth at the top and bottom of the slope; and (3) the shorter growing season at the higher elevations gives less time for growth there. There may be oth- er reasons or a combination of reasons for this phenomenon. In anv event, the relation- ship is strong and has management implica- tions. In the winter, deer in the Uinta National Forest and nearby areas primarily use south- and west-facing slopes at lower elevations (Bruce Giunta and Jordon Pederson, Utah Di- vision of Wildlife Resources, and Juan Spil- lett, Uinta National Forest, pers. comm.). Such areas coincide with situations where oak recovery is most rapid. Thus, manage- ment programs to regenerate oakbrush on deer winter ranges in our area may be short- lived. If Gambel oak is to be manipulated to improve deer range using conventional meth- ods in this area, a follow-up program that will retard oak recovery should be used. The 20-1 15 10- .85 1,530m 1,740 m 1,950 m Elevation 2,130m Fig. 1. The relationship between recovery rates of oak and elevation on various burns throughout the Uinta National Forest and nearbv areas. significant increase in time between major treatments will thus minimize management costs. Conclusions Elevation is a significant factor in affecting the recovery rate of oak after fire, with high- er elevation stands recovering more slowly. Recovery takes from 6 to .35 years in this area, with a modal recovery time of about 15 years. 25H 20- 15- 10 r = .9998 1,770m 1,920m Elevation 2,070 m Fig. 2. The relationship between recovery rate and elevation for oak in the Wallsburg Burn area. 10-1 0) 4-1 CD P^ >, U > o o 0) Pi > <: 5- s-w n -e Exposure Fig. .3. Histograph of the average recovery rate of oak stands with south to west exposures and north to east ex- posures on burns throughout the Uinta National Forest and nearby areas. 130 Great Basin Naturalist Vol. 40, No. 2 Acknowledgments The authors wish to thank the Uinta Na- tional Forest and the Utah Division of Wild- life Resources for permitting access to the study areas. Funding for this study was pro- vided by the Uinta National Forest (Supple- ment to Cooperative Agreement 12-11-204- 31). The authors also express thanks to Karl McKnight for help in the statistical analysis of the data. Literature Cited Allman, v. p. 1952. A preliminary study of the vegeta- tion in an exclosure in the chaparral of the Wasatch Mountains, Utah. Unpublished thesis, Brigham Young University, Provo, Utah. 236 pp. Anon. 1966. Conversion of thicket covered areas to pro- ductive grazing lands. USDA, Forest Service, Int. Reg. Range Imp. Notes 11(4): 1-4. Baker, W. L. 1949. Soil changes associated with recov- ery of scrub oak (Querctis gambelii) after fire. Un- published thesis, Univ. of Utah, Salt Lake City, 65 pp. Brown, H. E. 1958. Gambel oak in west-central Colo- rado. Ecology .39:317-327. Dills, G. G. 1970. Effects of prescribed burning on deer browse. J. Wildl. Manage. 34:540-545. Eastmond, R. J. 1968. Vegetational changes in a moun- tain brush commimity of Utah during 18 years. Unpublished thesis, Brigham Yoiuig Univ., Provo, Utah, 64 pp. Hallisey, D. M., and G. W. Wood. 1976. Prescribed fire in scrub oak habitat in central Pennsylvania. J. Wildl. Manage. 40:507-516. Marquiss, R. W. 1971. Controlling Gambel oak on rangelands of southwest Colorado. Colo. .'Vgri. Exp. Stat. Progress Report #PR71-9. 2 pp. 1972. Soil moisture, forage, and beef production from Gambel oak control in Southwest Colorado. J. Range Manage. 25:146-150. McKell, C. M. 1950. A study of plant succession in the oakbnish (Querctis gambelii) zone after fire. Un- published thesis, Univ. of Utah, Salt Lake City. 74 pp. Plummer, a. p., D. R. Christensen, and S. B. Monsen. 1966. Highlights, results, and accomplishments of game range restoration studies. Utah State Dept. of Fish and Game. Publ. No. 67-4:7-9. Plummer, A. P., R. Stevens, and K. R. Jorgensen. 1970. Highlights, results, and accomplishments of game range restoration studies. Utah State Dept. of Fish and Game. Publ. No. 70-3:26-30. Price, R. 1938. Artificial reseeding on oakbrush range in Central Utah. USDA Circ. No. 458. 19 pp. Smith, J. G. 1949. Deer forage observations in Utah. J. Wildl. Manage. 13:314-315. RELATIONSHIPS AMONG TOTAL DISSOLVED SOLIDS, CONDUCTIVITY, AND OSMOSITY FOR FIVE ARTEMIA HABITATS (ANOSTRACA: ARTEMIIDAE) Nicholas C. Collins' and Gray Stirling' .\bstract.— Graphs allowing interconversion between various physical chemical parameters are presented for five Aiicmia habitats in the western USA. Both the mean osmosity and its typical yearly range differ greatly among habi- tats. Consequently, Arteiiiici populations provide an interesting opportimity to study physiological and life history adaptations to differing degrees of habitat stability. Populations of Aiiemia, the brine shrimp, exist in isolated hypersaline environments throughout most of the world (McCarraher 1972). Their source waters span the entire natural spectrum of ion ratios (Cole and Brown 1967) and range from the massive and relatively permanent Great Salt Lake to tem- porary ponds 50 m to diameter (e.g. Broch, 1969; Khalaf et al. 1977). Not surprisingly, the individual populations exhibit morpholo- gical, physiological, developmental, and gen- etic differences that indicate they are locally adapted (e.g. D'Agostino 1965, Clark and Bowen 1976, Glaus et al. 1977, Collins 1977). Because the resting cysts of these popu- lations are easy to collect, transport, store, and hatch, Artemia populations are excellent subjects for comparative studies of genetics (Barigozzi 1974, Clark and Bowen 1976), physiology of ion regulation (e.g., Geddes 1975a, b, c), and life history tactics (Collins 1977, Glaus et al. 1977). Many such studies have involved comparisons of the perform- ance of various strains grown in a common medium, usually diluted or concentrated sea- water. An alternate approach, which accom- modates some strains that will not grow well in sea water, involves growing them each in their own source water, but at a common os- motic pressure. This alternative requires measurement of the osmotic characteristics of various dilutions of water from each source lake, a time-consuming process requir- ing osmometers that are both expensive and uncommon. To reduce the necessity for fu- ture osmometric measurements for studies of Artemia populations in the western United States, this paper presents relationships be- tween osmosity and more easily measured pa- rameters for source waters of five popu- lations. Data on pH changes with source concentration and information on the natural range of concentrations for each source are also presented. Methods Source waters from Arizona, New Mexico, Nebraska, and Washington, collected during 1976 and 1977, were filtered and diluted or concentrated by evaporation. Locations for each source are specified in the references in Table 1. Total dissolved solids (TDS) concen- trations were determined by evaporating five or ten ml samples to a constant weight at 100 C. Salt scale had to be repeatedly broken to insure completion of the drying process. Drying at temperatures higher than 100 G resulted in steam explosions within salt mas- ses that scattered the salt and biased the de- terminations. Conductivity meter readings were con- verted to specific conductance at 20 G using an NaCl calibration curve based on Wolf, Brown, and Prentiss (1975). Specific gravity at 20 C was measured gravimetrically using individually calibrated 50 ml volumetric flasks. Each determination 'Department of Zoolog)' and Erindale College, University of Toronto, 3359 Mississauga Road, Mississauga, Ontario L5L 1C6. 131 132 Great Basin Naturalist Vol. 40, No. 2 is the average of duplicate measurements. Osmosity, the molar concentration of NaCl having the same freezing point or osmotic pressure as the measured solution, was deter- mined with a Wescor vapor pressure os- mometer. The microvoltmeter output was calibrated with a series of NaCl solutions whose osmotic properties were assumed to correspond with Wolf et al. (1975). Two to five determinations were averaged for each data point in the figures. Results For three of the five sources (Figs. 1-3) os- mosity is closely, linearly related to TDS. Where Y is osmosity, X is TDS, n is the num- ber of measurements, and r is the product- moment correlation coefficient, the relation- ships for Penley Lake (Washington), Green Pond (Arizona), and Lily Lake (Nebraska) are respectively: E u (/) o E E TOTAL DISSOLVED SOLIDS Fig. 1. Relationships among TDS, conductivity (open circles), and osinositv (closed circles) for Penley Lake, near Oniak, Washington (Broch 1969). Numbers below upper margin are specific gravity measurements and those above lower axis are pH values for indicated TDS levels. June 1980 Collins, Stirling: Artemia Habitats 133 Y = .0081 X + .0176 n = 23 r = .995 Y = .0160 X + .0141 n = 19 r = .995 Y = .0103 X + .0439 n = 30 r = .998 The TDS-osmosity relationships for the other two lakes (Figs. 4, 5) did not appear to be lin- ear, and were drawn by eye. Similarly, the curvihncar TDS-conductivity relationships lor all lakes were drawn by eye. E a «/) o "I E >- > a o u 160 40 80 120 160 1 i 1 II 1 ' 1 ' 1 1 1 - 1.0390- 1.0465 • 1 CD CO eo yv - 140 - / / "~ - o/ ° / - 120 - GREEN POND o / /o / — ■ jPo / 100 f /• 80 - /• - 60 - • — 40 - — 20 - [ — -/ 9.8 1 1 ll 1 9.4 ll 1 1 1 1 — 3.20 - 2.80 - 2.40 - 2.00 _. o E -11.60 r" - 1.20 - .80 CO o C/) o 40 80 120 160 200 TOTAL DISSOLVED SOLIDS ( g/l ) Fig. 2. Relationships among physical-chemical parameters for Green Pond, near St. Johns, .\rizona (Cole and Whiteside 196.5). Legend as in Figure 1. 134 Great Basin Naturalist Vol. 40, No. 2 Discussion Generally, conductivity is the most con- venient indicator of osmosity or TDS. The natural concentrations of these soiuce lakes fall well above the highest scale of most exist- ing conductivity meters, but Figs. 1-5 will al- low rough predictions of TDS or osmosity from conductivity measurements of samples diluted to within the range of such meters. In such cases the errors in any prediction based on the graphs will be multiplied by the dilu- tion factor. At the highest source concentra- tions conductivity is not a precise predictor of TDS or osmosity (Figs. 4, 5). For such solu- tions an accurate dilution can be made for conductivity measurement, or TDS can be measured directly. Inaccuracies in TDS mea- O E, >- CO o C/3 O 40 80 120 160 200 TOTAL DISSOLVED SOLIDS (g/l) 240 Fit;. 3. Kelationships among pliysical-eheniical paraiufteis ior Lilv Lake, near Alliance, Nebraska (McCarraher 1970). Legend as in Figure L June 1980 Collins, Stirlinc;: Arteml\ Habitats 135 siirements from water inclusions in the crys- talizing salt mass can be eliminated by using a micrometer syringe for applying precisely measured small volumes (< 1 ml) to a pre- weighed filter paper circle. Such a technique will also allow much faster determinations tlian the one we used. Hydrometer measurements of specific gravity are often reported as a measure of concentration. Although this method appears to be quick, straightforward, and suitable for field measurements, our experience indicates even marginally accurate results require careful control of water temperature, wind, and cleanliness of the hydrometer that to- gether preclude most field measurements. In 160 140 ^ 120 E u o E E 100 80 60 40 20 - 50 100 150 200 250 300 350 T — ^ — T] — rpi — I 1 1 1 1 1 1 1 — I — r 2.0 4.0 50 100 150 200 250 300 350 400 TOTAL DISSOLVED SOLIDS (g/l) CO o CO Fig. 4. Relationships among phvsical-chemical parameters of Jesse Lake, near .Alliance, Nebraska (McCarraher 1970). Legend as in Figure 1. 136 Great Basin Naturalist Vol. 40, No. 2 the laboratory, TDS measurements or gravi- metric density determinations are almost as easy as hydrometer readings, if a con- ductivity meter is not available. A summary of natural osmosity and TDS levels from western Artemia habitats (Table 1) indicates that different populations expe- rience not only very different ion ratios and mean osmosities, but also very different sea- sonal ranges in source osmosity. Western North American Aiiemiu populations there- fore provide an interesting opportunity to E o o "I E >- > I— Q O 50 100 150 200 250 300 225 1 1.0827-= 1 52 / 52-^ 1 1 -75 8- 200 ZUNI SALT LAKE / / : 175 ■ 1 ' ^ 150 / • / - 125 / / - 100 - / / - 75 ■ /y - 50 ' 1/ - 25 r - • 8.7 \ ).5 8.1 7.9 1 1 ill 1. . J- 1 1 1 1 li 1 1 4.5 - 4.0 3.5 -13.0 =; o H2,5 ^ - 2.0 - 1.5 - 1.0 - .5 CO o C/9 O 50 100 150 200 250 300 350 TOTAL DISSOLVED SOLIDS (g/l) Fig. 5. Relationships amonn pliysical-cliemital parameters of Zuiii Salt Lake, near Queniado, New Mexico (Brad- bury 1971). Legend as in Figure 1. June 1980 Collins, Stirling: Artemia Habitats 137 identity the genetic, physiological, and life history characteristics that evolve in response to variability of environmental conditions, and in response to substantial differences in length of the growing season. Note added in proof: Recent talks with Nebraska residents revealed that the lake re- ferred to in this paper and in Collins (1977) as Lily Lake of McCarraher (1970, 1972) is actually an imnained smaller lake slightly northwest of the tnie Lily Lake. The range of osmosity for this lake is unknown; therefore the figures for it in Table 1 should be dis- regarded. Acknowledgments The National Geographic Society, the Nat- ural Sciences and Engineering Research Council of Canada, and the University of To- ronto supported this work financially. 1 thank Mr. Drew Piatt for permission to sample Green Pond, Gerald Cole, J. P. Bradbiuy, and Sarane Bowen for information on sampling sites, and R. W. Cummins and J. Svoboda for access to the o.smometer. LeeAnne Wilson and John Stoneman provided excellent tech- nical assistance. Liter.\ture Cited Baricozzi. C. 1974. Aitcinui: a survey of its significance in genetic prolileins. Evolutionary Biol. 7: 221-252. Bhadburv, J. P. 1971. l.iniiiology of Zuni Salt Lake, New Mexico. Geol. Soc. Anier. Bull. 82: .379-398. Broch. E. S. 1969. The osmotic adaptation of the fairy slirinip Brcinrhinccta campestris Lynch to salaine astatic waters. Linmol. Oceanogr. 14: 48.5-492. Clark, L. S., and S. T. Bowe.n. 1976. The genetics of Artemia xalina. VIL Reproductive isolation. J. Hered. 67: 38.5-.388. Claus, C, F. Benijts, a.nd P. Sorgeloos. 1977. Com- parative study of different geographical strains of the brine shrimp, Artemia salimi, pp. 91-105. In: Jaspers, E. (ed.). European Mariculture Society special publication No. 2. Bredene, Belgium. Cole, G. \., a.nd M. C. Whiteside. 1965. Kiatuth- lanna— a limnological appraisal. II. Chemical fac- tors and biota. PJateau .38: .36-48. Cole, G. \. and R. J. Brown. 1967. The chemistry of Artemia habitats. Ecology' 48: 8.58-861. Collins, N. C. 1977. Ecological studies of terminal lakes— their relevance to problems in limnology and population biology, pp. 411-420. In: D. C. Greer (ed.). Desertic terminal lakes. Utah Water Research Laboratory, Logan, Utah. D".\(;osTiNo, \. 1965. Comparative studies of Artemia ■salina (development and physiology). Unpub- lished dissertation. New York Univ. Gedde.s, M. C. 197.5a. Studies on an .Australian brine shrimp, Parartemia zietziana Sayce (Crustacea: .\nostraca)— I. Salinity tolerance. Comp. Bio- chem. Physiol. 51,\: .5.53-5.59. Table 1. Comparisons of physical and chemical characteristics of six Artemia .sources in the western USA. Water body Major salt Great Salt Lake, Utah NaCl Jesse Lake. Nebraska NaCOj Lily Lake. .Nebraska NaC()3-Cl Green Pond, NaC03-Cl .\rizona Penley Lake, Na2S()4 Washington Zuni Salt Lake, New Mexico NaCl Observed^ TDS range (g/1) "Typical" annual osmosity range^ (moles/1) References 130-300 .52-87 12-69 61-112 33-230 17.5-.350 2.-2.25 ..57-.90 .16-. 76 .98-1.8 .24-dryness Handv and Hahl 1966 ' McC:arraher 1970. 1972 .McCarraher 1970, 1972 Cole and Whiteside 1965 Broch 1969 ..35-5.3-1- Bradbury 1971 Range of values recorded in literature and personal observations, .\nnual range is usually smaller. IncKides only the period during which active Artemia are present. For some lakes, only one year's data are available, so nothing Ls known about be- tween-vear differences. 138 Great Basin Naturalist Vol. 40, No. 2 1975b. Shidies on an Australian brine shrimp, Parartemia zietziana Sayce (Crustacea: Anost- raca)— II. Osmotic and ionic rei;iilation. Comp. Biochem. Physiol. Sl.A: 561-571. 1975c. Studies on an Australian brine shrimp, Parartemia zietziana Sayce (Cnistaceae: .\nost- raca)— III. The mechanisms of osmotic and ionic regulation. Comp. Biochem. Physiol. 51A: 573-578. Handy, A. H., a.nd D. C. Hahl. 1966. Chemistry of the water in the Great Salt Lake, pp. 135-151. In: Stokes, VV. L. (ed.). The Great Salt Lake, Guide- book 20. Utah Geol. Soc., Salt Lake City. Khal.\f, a. N., M. a. Lattif, H. H. Mangalo, and M. Salih. 1977. A bioecological study on the brine .shrimp Artemia salina L. (Anostraca: Branchiop- oda) in two inland brine temporary ponds in Iraq. Bull. Biol. Res. Center, Bagdhad. 8: .37-48. McCarraher, D. B. 1970. Some ecological relations of fairy shrimps in alkaline habitats of Nebraska. .\mer. Midi. Nat. 84: .59-68. 1972. A preliminary bibliography and lake index of the inland mineral waters of the world. F.\0 Fisheries Circular No. 146. .33 pp. Wolf, A. V., M. G. Brown, and P. G. Pre.ntiss. 197.5. Concentratiye properties of aqueous solutions: conversion tables, pp. D218-D273. In: Weast, R. C. (ed.). Handbook of chemistry and physics, 56th ed. SPAWNING OF THE LEAST CHUB [lOTICHTHYS PHLEGETHONTIS) riioiiKis M. Baugh' .\bstract.— The least tluib, loticlitlujs phlf^clhonlis (Cope), a relict fisli in I'tali, spawned suecesslully under lab- oratory conditions. The least chub, lotichthys pJiIcgethontis (Cope), is a small (ca 5 cm) relict fish (Hubbs and Miller 1948) found only in a few local- ities in the western desert region of Utah. In 1973 the Utah Division of Wildlife Resources classified this species as endangered. There is little literature on /. phlcgethontis (Crawford 1979, Hubbs and Miller 1948, Pendleton and Smart 1954, Sigler and Miller 1963), and there are no reports of this species spawning under artificial light in closed-svstem aquaria. On 26 May 1979, I obtained five male and five female least chub from an open raceway at the Utah Division of Wildlife Resources fa- cility, Logan, Utah. The fish had originally been collected from Leland Harris Spring and the associated marsh between May 1977 and February 1978 by Crawford (1979). I placed the fish in a 61 X 41 X 31 cm (72.5 liter) aquarium with commercial aquarium gravel placed over a subgravel filter to a depth of about 5 cm. The water was con- stantly aerated and the tank was densely planted with simulated, broad-leafed plants. The fish were fed a mix of TetraMin^ Staple Food and Tetra KrillflakesR at 0630 and fro- zen San Francisco Bay Brand'^ brine shrimp at 1630 each day. Once each week, for a two-hour period, the water was filtered through a Vortex Dia- tom'^ filter. Also once each week, 15 percent of the aquarium water was drawn off and re- placed with an equal amount of aged tap wa- ter. One oimce of Instead Ocean*^ marine salt mix in solution was added each week. On 15 October 1979, I added two 29 cm long strips of Living World^ spawning grass to the aquarium. This spawning meditun was examined daily, and on 26 October one length of the medium contained five mildly adhesive eggs. This piece of medium was re- moved from the aquarium and placed in a 3.6 liter glass jar containing water from the aqua- rium. The water in the jar was mildly agi- tated with air. On 27 October one and on 28 October three additional eggs were removed from the aquarium to the jar. The water in the jar was maintained at the same temper- ature as that in the aquarium. Free swimming larvae were first noted on 31 October, and by 2 November all nine eggs had hatched. The larvae were able to adliere to glass and plastic. The mechanism of at- tachment was not studied. The following conditions existed at the time of spawning. During the 10 days prior to the day of last-noted egg deposition, the water temperature ranged from 17.7 to 18.8 C and averaged 18.2 C. The photoperiod was 14 hours of daylight and 10 hours of darkness. Water conditions were: pH 7.6, total alka- linity 84.1, total hardness 186, CI- 683, Ca 48.4, an Mg 45.1. Due to an equipment mal- fimction, dissolved oxygen was not measured. Two additional spawnings took place on 5 Noveiuber 1979 and 8 November 1979. Fif- teen eggs were gathered from the former and eight eggs from the latter spawning. In addi- tion, several other spawnings from these fish occurred from which the eggs were not taken. From the above, it appears that /. pfilc^e- thontis is amenable to culture in closed-sys- tem aquaria under artificial light. '1020 Custer .\ venue, Ogden, Utah 84404. 139 140 Great Basin Naturalist Vol. 40, No. 2 Acknowledgments I thank the Utah Division of Wildhfe Man- agement, especially Donald Andriano, for granting Permit s'CC-SL-919, which made this work possible; and Kent Miller, Utah Di- vision of Wildlife Resources, for reviewing this note. Literature Cited Cr.\\vford, M. 1979. Reproductive modes of the least chub [lotichtltijs pJilcgetJiontis Cope). Unpub- hshed thesis. Utah State Univ. 79 pp. HuBBs, C. L. .\ND R. R. Miller. 1948. Correlation be- tween fish distribution and hvdrographic historv in the desert basins of the western United States. Bull. Univ. Utah, Biol. Ser. 19(7): 17-166. Pe.ndleton, R. C, and E. W. Sm.\rt. 1954. A stud\ of the food relations of the least chub, lotichthijs phlegcthontis (Cope), using radioactive phos- phonis. J. Wildlife Manag. 12(2): 226-228. SiGLER. W. F.. .\ND R. R. Miller. 1963. Fishes of Utah. Pages 82-84 in Utah Div. ^\■ildlife Resour. Salt Lake Citv. TRANSFERRIN POLYMORPHISM IN BIGHORN SHEEP, OVIS CAXADEXSIS, IN COLORADO Patrick W, Roberts . Donald J. Nash . and Robert E. Keiss' .\bstr\ct.— Senmi transferrins were analyzed by polyacrylamide gel electrophoresis in four populations of Colo- rado bighorn sheep. Oti.s canadensis canadensis. Transferrin was found to be polymorphic, with two alleles. Tf D and Tf E. being represented in each of the four populations. Within herds the phenot) pic ratios confonned to values predicted h\- the Hardv- Weinberg equilibrium, .\niong populations, significant differences were seen with respect to phenotvpic frequencies. Transferrin polymorphisms have been de- scribed in a number of breeds of domestic sheep and in different species of wild sheep including Ovis canadensis, O. dalli, and O. mouflon (Nadler et al. 1971). They reported three transferrin alleles, Tf B+, Tf D, and Tf E, in two subspecies of bighorn sheep, O. c. canadensis and O. c. mexicana. In 14 speci- mens of O. c. canadensis from Montana, 13 had the Tf DE phenotype and one sheep was B^D. Two specimens of O. c. mexicana from .\rizona were of the EE phenotvpe. In Colorado, bighorn sheep historically ranged over much of the central and western parts of the state, but the distribution has been fragmented in recent times (Armstrong 1972), and there are now more tlian 30 dis- junct bands occurring in the less accessible parts of the higher mountains. A study was undertaken to characterize electro- phoretically demonstrable genetic variation in several sennu proteins and in hemoglobin of several disjimct herds to determine the de- gree of genetic similarity or dissimilarity among and within the bands sampled. The present stud\ is a report of transferrins ob- served in several herds. Blood samples were collected from four different herds in Colorado. The designation of the herds and their centers of distribution are as follows: (1) Poudre— north slope of Poudre Canvon. Larimer Co.. (2) Tarrvall— Tarryall and Kenosha Mts., Park Co., (3) Chalk Creek— Chafee Co., and (4) Gunnison- Gunnison Co. Transferrins were analyzed by poly- acrvlamide disc gel electrophoresis using the techniques described by Smith (1968). Gels were prepared at 7 percent (w/v) concentra- tion. Senuu samples were prepared by mak- ing serum with 50 percent sucrose containing 0.25 percent brom phenol blue as a tracking dve. Electrophoresis was carried out in tris- givcine buffer at pH 9.5. Twelve senmi sam- ples were electrophoresed for 26 minutes at 3 milliamps per gel at 10 C. Samples of domestic sheep blood of known transferrin tvpe were obtained from the Sero- log\" Laboratorv of Dr. Stormont of the Uni- versitv of California at Davis and were used as reference sera. All populations were polymorphic for transferrin phenotypes (Table I). Two herds. Chalk Creek and Poudre. had three pheno- t\pes and two herds. Tarr\ all and Gunnison, each had two phenot>pes. The phenotypes Table 1. Transferrin phenotypic frequencies of big- horn sheep, Ovis canadensis canadensis, in Colorado (uiunbers of observations in parentheses). Herd Tf DD TfDE TfEE Gunnison (7) 0.000 0.857 0.143 Chalk Creek (16) 0.313 0.374 0.313 Tarrvall (26) 0.577 0.423 0.000 Poudre (18) 0.316 0.526 0.158 'Department of Zoologj- and Entomolog\-, Colorado State University . Fort Collins, Colorado 80523. -Colorado Di\ision of Wildlife, Fort Collins, Colorado 80521. 141 142 Great Basin Naturalist Vol. 40, No. 2 were determined to correspond to those pro- duced by two alleles, Tf D and Tf E. Allelic frequencies ranged from 0.43 to 0.79 for Tf D and from 0.21 to 0.57 for Tf E. Significant differences among herds were observed for the distribution of phenotypes. Within herds the transferrin frequencies followed a Hardy- Weinberg distribution. The proportion of heterozygotes was high in all populations, with the lowest value of 0.375 being ob- served in the Tarryall herd. Although surveys of isozymes in natural populations of small mammals have indicated considerable genetic variability, relatively few biochemical studies have been done on large mammals. Bonnell and Selander (1974) found no polymorphisms in 24 presumptive loci in northern elephant seals. Hetero- zygosites of 0.04 have been reported for elk (Cameron and Vyse 1978), 0.04 for moose (Ryman et al. 1977), and 0.32 for white-tailed deer (Manlove et al. 1976). These species were monomorphic at the transferrin locus except for white-tailed deer, which had 23 percent heterozygosity. It is of interest that the bands of bighorn sheep in Colorado retain such a high degree of polymorphism, at least at the transferrin locus, although the populations have been relativelv isolated and have had relatively small population numbers. Some recent esti- mates of population size include Poudre, 65-75, Tarryall, 100, and Chalk Creek, 90-100. Results at the transferrin locus in- dicate that inbreeding within the herds may not be a major problem, although surveys of additional genetic loci should be undertaken. Literature Cited Armstrong, D. 1972. Distribution of mammals in Colo- rado. University Kansas Mas. Nat. Hist., Monogr. 3:1-415. Bonnell, M. L.. and R. K. Selander. 1974. Elephant seals: Genetic variation and near extinction. Sci- ence 184: 908-909. Cameron, D. C, .a.nd E. R. Vyse. 1978. Heterozygosity in Yellowstone Park elk, Cervus canadensis. Bio- cheni. Genet. 16:651-657. Manlove, M. N., J. C. Avise, H. O. Hillestad, P. R. Ramsey, M. H. Smith, and D. O. Straney. 1975. Starch gel electrophoresis for the study of popu- lation genetics in white-tailed deer. Proc. 19th Ann. Conf. S. E.. Game and Fish Comm. 29:.392-403. Xadler, C. F., .\. WooLF, AND K. E. Harris. 1971. The transferrins and hemoglobins of bighorn sheep {Otis canadensis), Dall sheep (Oris dalli) and mouflon (Otis niusimon). Comp. Biochem. Phvs- iol. 40B: 567-570. Ryma.n, N., G. Beckman, G. Briun-Petersen, and C. Reuterwall. 1977. Variability of red cell en- zymes and genetic implications of management policies in Scandinavian moose (Alces alces). He- reditas 85:157-162. Smith, I. (Ed.). 1968. Chromatographic and electro- phoretic techniques. Vol. 2. Zone— electro- phoresis. Wiley, New York, 524 pp. THE GENUS ERIOGONUM MICHX. (POLYGONACEAE) AND MICHEL GANDOGER James L. Reveal' Abstract.— Michel Gandoger, a notorious "splitter," proposed several new entities in the plant genus Eriogonum (Polvgonaceae) in a 1906 paper published in Belgium. Because he used the term species at two different ranks, in violation of the International Code of Botanical Nomenclature, manv of his names are invalid. Unlike his papers published in France, this one was apparentl\- edited so that some names were validly published and some are invalid. A review of the 1906 Eriogonum paper shows that a majority of both specific and infraspecific entities proposed are valid, but some names, long in use and assinned to be valid, are, in fact, invalid. Even so, most of his names are sviionyms. Each name proposed by Gandoger is reviewed and a nomenclatural and taxonomic disposition made. Two new combinations are made within E. hitcolum Greene, var. caninum (Greene) Reveal and var. pedunrulatum (S. Stokes) Reveal. Michel Gandoger (1850-1926), in the words of Keck (1958), was a "French abbe; author of 'Flora Europae' (27 vols.); volumi- nous writer; amasser of a huge herbarium now at Lyon; a 'splitter' who named thou- sands of unacceptable species." Gandoger is also mentioned in the International Code of Botanical Nomenclature (Stafleu et al. 1978) as an example of Art. 33.4. This article deals with the problem of misplaced ranks such as the use of the term species as a rank within a species. As noted in the code, Gandoger ap- plied the term species and used binary no- menclature for two categories of taxa of con- secutive rank, the higher rank being equivalent to that of species in contemporary literature, while he misapplied the same term to a lower rank. These latter terms are not validly published. In his 1906 paper, Gando- ger used three sets of ranks, the species, spe- cies of the second order ("speciebus secundi ordinis"), and a rank which has come to be identified with the rank of variety (Heller 1907), although Gandoger causally refers to this latter rank as forms ("formis," "formas secernendas, " "formae memorabiles," "mem- orantur sequentes formae," "varieas formas," "modo formae," etc.). He used the terms "varians" and "variabilis" to allude to the same category. As for the species of the sec- ond order, in addition to terming them "spe- ciebus secimdi ordinis," he also used the term siibspeciebus or subspecies to refer to entities he then proceeded to treat as binomials and to designate by the expression sp. n. The following year. Heller (1907) present- ed a "compilation" of the Gandoger paper, noting that he was presenting only "the new forms described in this paper." Heller himself had some difficulties with the names. He mentioned, after Eriogonum aspalathoides (as it was called bv Heller), that this name was "perhaps intended as a variety of E. fasci- ciilatiim, but the way the name is printed .should indicate a species." Only once does Heller include Gandoger's terminology allud- ing to species of a lower rank, this being for the phrases published under E. sphoerocepha- him, "Inter formas varias duae sequentes, ut subspecies, praecipue distingui possunt." It must be noted that Heller was merely presenting Gandoger's results, and he cannot be assumed to have validated any of the oth- erwise invalid Gandoger names. Normally, Gandoger published his papers in the Bulletin de la Societe Botaniqiie de France, but for Eriogonum his choice was the Bulletin de la Societe Roijale de Botanique de Belgique, and, because of this, some of the names may be validly described. Possibly 'Department of Botany, University ot Maryland. College Park. Maryland 20742, and National Museum of Natural History. Smitfisonian Institution, Wash- ington, D.C. 20560. Research supported by National Science Foundation Grant BMS75-13063. This is Scientific Article .\2714. Contribution No. 5761 of the Maryland .\gricultural Experiment Station, Department of Botany. 143 144 Great Basin Naturalist Vol. 40, No. 2 they may because of the way the editor had the paper set, thereby removing from the original manuscript some of Gandoger's ec- centricities. In reviewing Gandoger's papers in the French journal, it is clear that the ma- jority of his names are invalidly published. In reviewing a few of his articles containing species from the United States, I failed to note a single instance when a species name was validly published. This little known fact, at least in the United States and perhaps else- where, means that all the Gandoger names must be carefully checked before they can be accepted. This is particularly true for floristic workers. Finally, it is sometimes difficult to know exactly where the various sections of Gando- ger's paper start and end so that one can de- termine if the name is validly published or, in fact, is not because of a sentence presented several pages before. I have tried to follow the intent of Gandoger's paper, and hope I have interpreted each segment correctly. The following treatment indicates which names are valid, which are not, and the tax- onomic status of each name. The full author citation of each name is given, and the num- ber following the name is the page on which it was proposed by Gandoger. Erio^onum abertianum Torr. in Emory var. rubenimiim Gandoger, 185, valid, a synonym of var. abertianum. Eriogonum aberiianum var. neomexicanum Gandoger, 185, valid, a synonym of var. abertianum. Eriogonum arizonicum Gandoger, non Stokes ex Jones, 186, valid, a synonym of E. pharnaceoides Torr. in Emory var. pharna- ceoides. Eriogonum alattim Torr. in Sitgr. var. mac- douglasii (Tandoger, 186, valid, a synonym of var. mogollonense Stokes ex Jones. Eriogonum alatum var. brevifolium Gando- ger, 186, valid, a synonym of var. alatum. Eriogonum anemophiUim (as anemophijl- luni) Greene var. cusickii Gandoger, 186, valid, a synonym of E. cusickii M. E. Jones. Eriogonum angulosum Benth. var. rectipes Gandoger, 186, valid, a synonym of E. ma- cukitum Heller. Eriogonum angulosum var. patens Gando- ger, 187, valid, a synonvm of E. maculatum Heller. Eriogonum angulosum var. pauciflorum Gandoger, 187, valid, a synonym of E. ma- culatum Heller. Eriogonum angulosum var. flabellatum Gandoger, 187, valid, a synonym of E. ma- culatum Heller. Eriogonum annuum Nutt. var. pauciflorum Gandoger, 187, valid, a synonym of E. an- nuum. Eriogonum hitchcockii Gandoger, 187, valid, a synonym of E. annuum. Eriogonum juncinellum Gandoger, 187, valid, a synonym of E. davidsonii Greene. Eriogonum salicorniodes Gandoger, 187, valid, a good species restricted to clay slopes in southwestern Idaho and adjacent south- western Oregon. It is most closely related to E. collinum Stokes ex Jones but most often confused with E. baileyi S. Wats. Synonyms of this species include E. demissum S. Stokes (1936) and the var. romanum S. Stokes. Eriogonum caespitosum Nutt. var. alijs- soides Gandoger, 188, valid, a synonym of E. caespitosum. Eriogonum nevadense Gandoger, 188, valid, a synonym of E. ochrocephalum S. Wats. var. ochrocephalum. Eriogonum elatum Dougl. ex Benth. var. limonifolium Gandoger, 188, valid, a syn- onym of £. elatum var. elatum. Eriogonum elatum var. erianthum Gando- ger, 188, valid, a synonym of var. elatum. Eriogonum fasciculatum Benth. var. oleifo- lium Gandoger, 189, valid, a synonym of E. fasciculatum var. fasciculatum. Eriogonum fasciculatum Benth. var. as- palathoides Gandoger, 189, valid, a synonym of var. fasciculatum. As noted by Heller (1907), there is some problem with this name; that is, there is an E. prior to the epithet. However, based upon the rest of the text, this is clearly as printer's error and should be list- ed as a variety and not as a species, as was done by Heller in his "compilation. " Eriogonum flavum Nutt. in Fras. var. folia- tum Gandoger, 189, valid, a synonym of E. jamesii Benth. in DC. va.r. flavescens S. Wats. Eriogoni4m flavum var. linguifolium Gan- doger, 189, valid, a synonym of var. flavum. Eriogonum leucocladum Gandoger, 189, valid, a synonym of E. baileyi S. Wats. var. divaricatum (Gandoger) Reveal. ErioEonum heracleoides Nutt. var. micran- June 1980 Reveal: Eriogonum Nomenclature 145 thum Gandoger, 189, valid, a synonym of £. herocleoides var. angustifolium (Niitt.) Torr. & Gray. Eriogonum hcracleoides var. viride Gando- ger, 190, valid, a synonym of £. umhellatum Torr. var. nevadense Gandoger. Eriogonum hcracleoides var. multiccps Gandoger, 190, valid, a synonym of var. hcra- cleoides. Eriogonum hcracleoides var. utahensis Gandoger, 190, valid, a synonvm of var. hcra- cleoides. Eriogonum hcracleoides var. rydbergii Gan- doger, 190, valid, a synonym of var. hcra- cleoides. Eriogonum jamesii Benth. in DC. var. sim- plex Gandoger, 190, valid, a good variety of- ten included within var. jamesii; this phase of the species is restricted to southwestern Kan- sas. Eriogonum jamesii var. neomexicanum Gandoger, 190, valid, a synonym of var. jamesii. Eriogonum longifolium Nutt. var. long- idcns Gandoger, 190, valid, a synonym of E. longifolium var. gnaphalifolium Gandoger. Eriogonum longifolium var. gnaphalifo- lium Gandoger, 190, valid, that phase of the species restricted to Florida; often called E. floridanum Small. Eriogonum longifolium var. floridanum (as floridana) Gandoger, 190, valid, a synonym of £. longifolium var. gnaphalifolium Gando- ger. Eriogonum longifolium var. lindheimeri Gandoger, 190, valid, the western phase of the species now best considered a synonvm of var. longifolium. Eriogonum longifolium var. caput-fclis Gandoger, 190, valid, a synonym of var. lon- gifolium. Under the heading, Eriogonum micro- thccum Nutt., Gandoger states "inter quas se- quentes altem pro subspeciebus habueris" and refers to the following six entities all published with binary names. Eriogonum macdougalii Gandoger, 191, in- valid, a synonym of E. microthecum Nutt. var. foliosum (Torr. & Gray) Reveal. The name was used by Stokes (1936) at the varia- tal rank within E. microthecum, where she validated the name as E. microthecum var. macdougalii S. Stokes. Eriogonum myrianthum Gandoger, 191, in- valid, a synonym of E. cffusum Nutt. var. cf- fusum. Eriogonum sputhulare Gandoger, 191, in- valid, a synonym of E. microthecum Nutt. var. laxiflorum Hook. The name was used by Stokes (1936) at the variatal rank within E. microthecum, where she validated the name as E. microthecum var. spathulure S. Stokes. Eriogonum intricatum Gandoger, 191, in- valid, a synonym of E. microthecum Nutt. var. laxiflorum Hook. Eriogonum Iwlichrysoides Gandoger, 192, invalid, a synonym of E. cffusum Nutt. var. rosmarinioides Benth. in DC. The name was validly published by Rydberg (1931) and the citation of the name should be E. helichry- soides Rydb., Brittonia 1: 87. 1931, without reference to Gandoger. Rydberg was in- correct that Gandoger had proposed a varie- ty, so the combination attributed to Gando- ger by Rydberg in synonymy, E. nucrothecum var. helichrysoides, must be attributed to Rydberg as well. Eriogonum sarothriforme Gandoger, 192, invalid. The type of this name, collected by Osterhout at Glenwood Springs, Garfield Co., Colorado, may represent a distinct taxon. It belongs to the E. brevicaule Nutt. complex and is seemingly a part of the polymorphic species E. lonchopln/llum Torr. & Gray, a taxon that morphologically bridges the E. brevicaule complex, a group of herbaceous perennials, with those species typified bv E. corymbosum Benth. in DC, a series of shrubs or subshnibs. The Garfield Co. plants tend to be more slender than typical E. lonchophyl- lum of southern Colorado and adjacent northern New Mexico. These plants also tend to resemble some of the more robust, but yel- low-flowered, forms of E. brevicaule found in Rio Blanco Co., Colorado. Although well known to me, I am still uncertain what to do with the Glenwood Springs plants. Eriogonum niveum Dougl. ex Benth. var. suksdorfii Gandoger, 192, valid, a synonym of E. niveum. Eriogonum niveuni var. candelabrum Gan- doger, 192, valid, a svnonvm of E. niveum. Eriogonum ochrolcucum Small var. macro- podum Gandoger, 192, valid, the basionym for E. ovalifolium Nutt. var. macropodum (Gandoger) Reveal. 146 Great Basin Naturalist Vol. 40. No. 2 Eriogonum ochroleucum var. decahans Gandoger, 192, valid a synonym of E. ocali- folium Nutt. var. macropodum (Gandoger) Reveal. Under the heading of Eriogonum ovalifo- lium Nutt., Gandoger proposed a series of names in two ranks, all of which he refers to by "formas quarum non paucas saltern pro speciebus secimdi ordinis haberi possimt," which I believe he wished to apply only to those names he indicated by the designation oi sp. n. Eriogonum flavissitnum Gandoger, 193, in- valid, a svTionym of E. ovalifolium Nutt. var. anserinum (Greene) R. J. Davis. The name was used by Stokes (1936) as a subspecies of E. ovalifoliwn, where the name was validated as E. ovalifolium Nutt. ssp. anserinum S. Stokes. Eriogonum cusickii Gandoger, non M. E. Jones (1903), 193, invalid, a synonym of E. strictum var. proliferum (Torr. & Gray) Re- veal. The name was used at the variatal rank by Stokes (1936) within E. strictum, where the name was validated as E. strictum Benth. var. cusickii S. Stokes. Eriogonum cusickii Gandoger var. califor- nicum Gandoger, 193, invalid, a synonvm of E. strictum var. proliferum (Torr. 6c Grav) Reveal. Eriogonum ovalifolium Nutt. var. neva- dense Gandoger, 193, valid, a good varietv of the species, this being the yellow-flowered, early flowering expression which I have called var. multiscapum Gandoger (see be- low). Eriogonum ovalifolium var. deltoideum Gandoger, valid, 193, a synonym of £. ovali- folium var. nevadense Gandoger. Eriogonum dichroanthum Gandoger, 193, invalid, a synonym of E. ovalifolium Nutt. var. nevadense Gandoger (see discussion un- der var. multiscapum below). Eriogonum ovalifolium Nutt. var. utahense Gandoger, 194, valid, a synonym of var. ova- lifolium. Eriogonum ovalifolium Nutt. var. multi- scapum Gandoger, 194, valid, a synonym of var. ovalifolium. For several years I have misapplied this name to the yellow-flowered phase of the species (Reveal & Munz 196S, Reveal 1973, 1976). Gandoger based this name on plants gathered by Nelson (4658) at Cokeville, Uinta Co., Wyoming, upon which he also based the name E. dicJiroanthum. An examination of this collection b\ me in 1966, and then by my brother, Jon A. Reveal, in 1973, was inconclusive in that it could not be fully determined if the flowers of var. multi- scapum were truly yellow. Gandoger divided the collection and moimted each on a sepa- rate sheet, with E. dichroanthum having "flores fructusque flavissimi." and the flowers of var. midtiscapum, to him. were "'ochro- leuci." The same collection at KSC is strictly yellow flowered, but the specimen at RM is a mixture of a bright-yellowed specimens matching E. dichroanthum and a whitish or. at best, pale yellow-flowered specimen that matches var. multiscapuryi. I now believe that the yellow-flowered, early spring flowering phase of the species should be called var. nevadense and the type of var. lyiultiscapum assigned to var. ovalifolium. Eriogonum ovalifolium Nutt. var. cijclo- phijllum Gandoger, 194, valid, a synonym of var. macropodum (Gandoger) Reveal. A reex- amination of the types of var. cyclophyllum and var. cerastoides (see below), as part of this study, clearly shows that these names must be referred to what I (Reveal 1968) had earlier called var. macropodum. making a new combination for this name by transfer- ring it from E. ochroleucum Small to E. ovali- folium. Previously, these two names have been referred to var. ovalifolium (Hitchcock 1964). The var. cijclophyUum is close to var. ovalifolium and may represent one of the many intermediate expressions between the two varieties. I retain the usage of var. macropodum. Eriogonum ovalifolium Nutt. var. ceras- toides Gandoger, 194, valid, a synonym of var. macropodum (Gandoger) Reveal (see the discussion above). Eriogonum ruhidum Gandoger. 194, in- valid, a synonym of E. ovalifolium Nutt. var. depressimi Blankinship. Eriogonum ruhidum var. frigidum Gando- ger, 194, invalid, a synonym of E. ovalifolium Nutt. var. depressum Blankinship. Eriogonum roseiflorum Gandoger. 194, in- valid, a svnonvm of E. ovalifolium Nutt. var. ovalifolium. Eriogonum piperi Greene var. ochrocepha- lum Gandoger, 195, valid, a synonym of E. June 1980 Reveal: Eriogonum Nomenclature 14' flaviim \utt. in Fras. var. piperi (Greene) M. E. Jones. Eriogonum piperi var. longifloruni Gando- ger, 195, valid, a synonym of E. flavum Xutt. in Fras. var. piperi (Greene) M. E. Jones. Under Eriogonum pohjanthum Benth., now better known as E. umbellatum Torr. var. pohjanthum (Benth. in DC.) M. E. Jones, Gandoger introduces two validly described species simply stating that "species duae se- quentes huic sunt affines." Eriogonum marginale Gandoger, 195, valid, a synonym of E. umbellatum Torr. var. aureum (Gandoger) Reveal. Eriogonum glaherrimum Gandoger, 195. valid, the basionym of E. umbellatum Torr. var. glaberrimum (Gandoger) Reveal. Eriogonum glaberrimum var. aureum Gan- doger, 195, valid, the basionym of E. um- bellatum Torr. var. aureum (Gandoger) Re- veal. Eriogonum pohjcladon Benth. var. mexi- canum Gandoger, 196, valid, a synonym of E. pohjcladon. Eriogonum polycladon Benth. var. crispum Gandoger, 196, valid, a synonym of E. poly- cladon. Eriogonum racemosum Xutt. var. sagitta- tum Gandoger, 196, valid, a synonym of E. racemosum. Eriogonum racemosum Xutt. var. cordi- folium Gandoger, 196, valid, a svnonvm of E. racemosum. Eriogonum reniforme Torr. & Frem. var. asarifolium Gandoger, 196, valid, a svnonvm of E. pusillum Torr. 6f Gray. Eriogonum praebens Gandoger, 196, valid, a s\nonym of E. bailcyi S. Wats. var. diva- ricatum (Gandoger) Reveal. Eriogonum praebens var. diiaricatum Gan- doger, 196, valid, the basionym of E. baileyi S. Wats. var. divaricatum (Gandoger) Reveal. Under the heading Eriogonum sphaero- cephalum Dougl. ex Benth.. Gandoger states "inter formas varias duae sequentes. ut sub- species, praecipue distingui possunt." Eriogonum cupreum Gandoger, 196. in- valid, a synonym of E. umbellatum Torr. var. umbellatum. Eriogonum halimioides Gandoger, 197, in- valid. This name was validated by Stokes (1936) as E. sphaerocephalum var. hali- mioides S. Stokes without reference to Gan- doger. Eriogonuin subalpinum Greene var. arach- noideum Gandoger, 197. valid, a synonym of E. umbellatum Torr. var. dichrocephalum Gandoger. Eriogonum subalpinum var. vulcanicum Gandoger, 197, valid, a synonymy of £. um- bellatuin Torr. var. majus Hook. Eriogonum subalpinum var. stenophyllum Gandoger, 197, valid, a synonym of E. um- bellatum Torr. var. majus Hook. Eriogonum subalpinum var. subnivale Gandoger, 197, valid, a synonym of E. um- bellatum Torr. var. majus Hook. Eriogonum tenellum Torr. var. grandi- florum Gandoger, 197, valid, a synonym of E. microthecum Nutt. var. laxiflorum Hook. Eriogonum tenelhw} var. sessiiflorum Gan- doger, 198, valid, a synonym of E. micro- thecum Xutt. var. laxiflorum Hook. Eriogonum tenellum var. erianthum Gan- doger, 198, valid, a synonym of E. micro- thecum Xutt. var. ambiguum (M. E. Jones) Reveal in Mimz. Eriogonum thurberi Torr. var. parishii Gan- doger. 198, valid, a svnonvm of £. thurberi. Eriogonum thurberi var. acutangulum Gan- doger, 198. valid, a synonvm of E. macula- turn Heller. Eriogonum tlnjmoides Benth. in DC. var. pallens Gandoger, 198, valid, a synonym of £. thymoides. Eriogonum umbellatum Torr. var. cran- dallii Gandoger, 198, valid, a synonym of var. umbellatum. Eriogonum umbellatum var. chrysanthum Gandoger, 198, valid, a synonym of E. um- bellatum var. stellatum (Benth.) M. E. Jones. Eriogonum umbellatum var. nevadense Gandoger, 198, valid, a good variety referr- ing to that phase of the species found in the Sierra Xevada of California northward into Oregon and eastward into uestern Xevada that has been routinelv called var. umbella- tum (Reveal & Munz 1968; see Howell 1976). Eriogonum umbellatum var. cladophorum Gandoger, 198, valid, a synonym of var. um- bellatum. Eriogonum umbellatum var. dichro- cephalum Gandoger, 199, valid, a good varie- ty applied to that phase previously called E. 148 Great Basin Naturalist Vol. 40, No. 2 umbeUatum var. aridum (Greene) C. L. Hitchc. (Hitchcock 1964). Eriogonwn umhellatum var. californicuni Gandoger, 199, valid, a synonym of E. um- beUatum var. nevadense Gandoger. Eriogonum vimineiim Dougl. ex Benth. var. rigescens Gandoger, 199, valid, a synon- ym of E. vimeneum. Eriogonum vimineum Dougl. ex Benth. var. califomicum Gandoger, 199, valid, a syn- onym of E. luteohim Greene var. caninum (Greene) Reveal, comb, nov., based on E. vi- mineum var. caninum Greene, Fl. Francisc. 150. 1891. The type of var. califomicum is somewhat intermediate between var. luteo- lum and var. caninum, being closer to the latter than the former. I have long recog- nized the caninum phase (Reveal & Munz 1968) as distinct from E. vimineum, but J. T. Howell, who has considered the expression only as a variant, has pointed out in our con- versation that he had observed a large series of intermediate populations, as had I, which held the Mt. Tamalpais plant, var. caninum, well within the boundaries of what he called E. vimineum. This latter expression, however, proved to be E. luteohim rather than E. vimi- neum, and I am now following Howell's (1970) taxonomic disposition of this local en- demic. In addition to this variant of E. luteo- lum, the Sierra Nevada plant I recognized as a distinct species previously (Reveal 1970) should be included within this species as well. Thus, I propose E. luteohim var. peduncula- tum. (S. Stokes) Reveal, stat. & comb, nov., based on E. pedunculatum S. Stokes, Leafl. W. Bot. 2: 48. 1937. Eriogonum vimineum var. oregonense Gan- doger, 199, valid, a synonym of E. vimineum. Eriogonum restioioidcs Gandoger, 199, valid, a synonym of E. baileyi S. Wats. var. bailey i. Literature Cited Gandoger, M. 1906. Le genre Eriogonum (Poly- gonaceae). Bull. Soc. Roy. Bot. Belgique 42: 18.3-200. Heller, A. A. 1907. Compilations. Miihlenbergia 3: a3-96. Hitchcock, C. L. 1964. Eriogonum. Univ. Wash. Publ. Biol. 17(2): 104-138. Howell, J. T. 1970. Marin flora. 2d ed. Berkeley: Univ. California Press. 366 pp. 1976. Eriogonum notes VII. Mentzelia 1: 17-22. Keck, D. D. 1959. "Abbreviations of authors' names." Pages 1551-1576 in P. A. Munz, & D. D. Keck, eds. A California flora. Berkeley: Univ. California Press. Reveal, J. L. 1968. Some nomenclatural changes in Eriogonum (Polygonaceae). Taxon 17: 531-533. 1973. Eriogonum (Polvgonaceae) of Utah. Phvto- logia 25: 169-217. 1976. Eriogonum (Polvgonaceae) of Arizona and New Mexico. Phytologia 34: 409-484. Reveal, J. L., and P. A. Munz. 1968. "Eriogonum." Pages 33-72 in P. A. Munz, ed. Supplement to A California flora. Berkeley: Univ. California Press. Rydberg, p. a. 19.31. Taxonomic notes on the flora of the prairies and plains of central North .\merica. Brittonia 1: 79-104. Stafleu, F. a., et al. 1978. International code of bot- anical nomenclatme. Regnum Veg. 97: 1-457. Stokes, S. G. 1936. The genus Eriogonum, a preliminary studv based on geographic distribution. San Fran- cisco: J. B. Nebiett. 124 pp. PARASITES FROM TWO SPECIES OF SUCKERS (CATOSTOMIDAE) FROM SOUTHERN UTAH J. Craig Biieiiholt' and Richard A. Heckmann .\bstr\c:t.— Twenty Ciitostoiims latipiiiuis and 50 Catostomiis lUscohohs from La Verkin Creek and the Fremont River in southern Utah were collected and surveyed tor parasites. Data from the survey indicated that 83 percent of the fish were infected with at least one parasite, with the fish from La Verkin Creek harboring more parasites. Twelve genera and 12 species of parasites were identified from these fish. .\ monogenetic trematode, G\jwd(trtiilus dedans, which was found in 90 percent of the fish, was the most common parasite. Comments are included on habi- tat and host variations for the parasitofauna from suckers taken from the two locations. A survey of the parasites of the cato,sto- mids, Catostomiis latipinnis and C. discoboUs, was conducted at La Verkin Creek, southern Utah, and the Fremont River near Hanks- ville, Utah. The objectives of this survey were to provide a hst of parasites for C. discoboUs and C. latipinnis in La Verkin Creek and Fremont River and to correlate water param- eters and benthos from these streams with parasite loads. Both streams contain well-e.s- tablished populations of the listed suckers. An exhaustive survey of parasites can explain the source or reservoir of serious pathogens for endangered species and commercially impor- tant fish. Catostomids are found exclusively in North America, excluding two or three Asiatic spe- cies (Pflieger 1975). Catostomiis discoboUs is found in Idaho, Utah, and Nevada in the fol- lowing drainages: Colorado River above the Grand Canyon, upper Snake River, Bear Riv- er, and Weber Lake outflows. Catostomiis latipinnis is unique to the Colorado River drainage (Eddy 1959). Information con- cerning the life history of these two .species is limited. Catostomid levels in both study areas for this project are maintained by resident sucker populations. Both of the streams selected in this survey are unstable desert streams. Much of the sub- strate is sand which shifts and prevents deep pools from forming. Flash floods can disrupt and completely change the nature of the streams and change the macroinvertebrate population. Because of this, fish species, e.g., salmonids and centrarchids, that cannot with- stand the instability of the stream and the consequent change in macroinvertebrate food source are not found extensively in these two streams. The ichthyofauna found in the study area of the Fremont River are: Longnose dace, RJiinichthys cataractae; speckled dace, R. osciiliis; leatherside chub, Gila copei; blue- head sucker, C. discoboUs; and flannelmouth sucker, C. latipinnis (Heckmann 1976). The speckled dace and leatherside chub are omni- vores that feed on aquatic plants, insects, and cmstaceans. The flannelmouth suckers are herbivores which feed on algae, diatoms, parts of higher plants, and seeds. The blue- head sucker is a bottom feeder which scrapes algae and other organisms from rocks (Sigler and Miller 1963). The largest fish found in the Fremont River is the flannelmouth sucker and the smallest is the speckled dace. None of the fish found in the sample area are pisci- vorous. All fish feed either on aquatic in- vertebrates or plant material. Fi,sh predators mav include birds and small mammals. Fish .species inhabiting the survey site at La Verkin Creek are: speckled dace, R. os- cuUis; Virgin River spinedace, Lcpidomeda mollispinis; red shiner, \otropiis liitrcnsis; woundfin minnow, Plagopterus argentissimus; 'Department of Zoolog\', Brigham Young University, Provo, Utah 84602. 149 150 Great Basln Naturalist Vol. 40, No. 2 flannelmouth sucker, C. kitipinnis; bluehead sucker, C. discoholis or desert sucker, C. clarki; and rainbow trout, Salmo gairdneri (Winget and Baumann 1977). La Verkin Creek, in comparison to the Fremont River, is more stable and less turbid, resulting in the presence of riffles, pools, and some holes 1-1.5 m deep. Because of the difference in stream conditions, rainbow trout are planted by the Utah Division of Wildlife Resources in small numbers. Two species of fish, L. moUis- pinis and P. argentissimus, are considered en- dangered. The woimdfin minnow, rainbow trout, and Virgin River spinedace are consid- ered carnivores feeding mainly on in- vertebrates. Catostoriius discoholis and N. lut- rensis are considered bottom-dredging detritovores. Catostoinus kitipinnis and R. os- culus are selective omnivores (Winget and Baumann, 1977). The top carnivore in a tro- phic scheme would be S. gairdneri because it may feed on smaller fish. Direct competition "s virtually eliminated because those species vith similar feeding habits have different habitat preferences or specific food prefer- ences (Winget and Baumann 1977). Parasites of catostomids other than C. lati- pinnis and C. discoholis have been studied by researchers in the United States and Canada. Hoffman (1967) lists known parasites for 12 species of catostomids. Other surveys have been conducted bv Voth and Larson (1968), Amin (1969), Threlfall and Hanek (1970), Amin (1974), White (1974), Mackiewicz (1963), Price and Arai (1967), Dechtiar (1969), Daly and De Giusti (1971), Clifford and Facciani (1972), Hatha wav and Herlev- ich (1973), Schell (1974), andHayunga and Grey (1976). The most widely studied ca- tostomids are white suckers, C. cornmersoni, and longnose suckers, C. catostomiis. These surveys deal primarily with metazoan para- sites, and little information concerning the protozoan parasites is included. M.\TERIALS AND METHODS Through the use of electrofishing, 18 flannelmouth suckers and 40 i)luehead suck- ers were collected from La Verkin Creek near the Toquerville cemeterv, southern Utah. Two flannel-mouth suckers and 10 bluehead suckers were collected from the Fremont River one mile west of Capitol Reef National Park, near Hanksville, Utah. The fish were transported to Brigham Young Uni- versity in iced holding tanks. Limited num- bers of fish were obtained due to collecting restrictions. Each fish was checked for parasites. The suckers were euthanized by a blow to the head before being weighed and measured (Table 1). Following macroscopic exam- ination, scrapings of the surface, gills, medial area of the opercula, and eyes, were exam- ined for parasites. Because the blood vessels were niptiued, gill scrapings were used to check for hemoflagellates. Intestine, liver, and gall bladder were excised and examined for endoparasites. The presence of metacer- cariae inhabiting the liver was checked by pressing a piece of the organ between two glass slides and examining it without magnifi- cation. Protozoans were either air dried or pre- served in 10 percent formalin. Permanent preparations of monogenetic trematodes were made with Turtox mounting and stain- ing medium (nonresinous stain mountant CMC-S). Leeches were also fixed in formalin and all were identified tlirough the use of keys listed in Hoffman (1967). Cestodes were placed directly into AFA fixative to prevent total relaxation. Digenetic trematodes were placed in 95 C water to promote relaxation and then placed in AFA fixative. Cestodes and digenetic trematodes were stained with Semichon's carmine for 12 hours and then destained in changes of acid alcohol to improve color contrast. After des- taining, the specimens were dehydrated in 95 percent and 100 percent ethyl alcohol for one hoiu' each. Once dehydrated, specimens were cleared in xylene and then mounted with Permount on glass slides. Morphological characteristics given in Hoffman (1967) were used for identification of trematodes and nematodes. Preliminary identification of the caryophyllid tapeworms was confirmed by John S. Mackiewicz (State University of New York at Albany). Water chemistry and macroinvertebrate data were obtained from studies by Heck- mann (1976), Winget and Reichert (1976), and Winget and Baumann (1977). June 1980 Breinholt, Heckmann: Fish Parasites 151 Results Data from the examination of 40 bluehead and 18 flannelmouth suckers from La Verkin Creek in southern Utah indicated that 55 suckers harbored at least one species of para- site. Thirty-seven of 40 bluehead suckers were infected and all 18 flannelmouth suck- ers harbored parasites (Table 2). Postmortem examination of 12 suckers from the Fremont River revealed one of 10 bluehead suckers and 2 of 2 flannelmouth suckers were para- sitized (Table 2). Twelve genera and 12 species of parasites were identified from fish from La Verkin Creek. The most frequently encountered par- asite was Gyrodactylus elegans, a mon- ogenetic trematode which was found in 52 Table 1. Weights and measurements of fish exam- ined. Species Weight Length No. offish Stream (gm) (TL) (cm) 1 C. discobolis La Verkin Creek 45 17 2 43 14 3 38 16 4 33 15 5 33 14 6 55 18 7 65 19 8 50 16 9 40 16 10 25 13 11 35 16 12 50 16 13 60 18 14 85 21 15 90 21 16 45 17 17 20 13 18 40 16 19 40 16 20 40 17 21 15 11 22 45 17 23 30 14 24 35 15 25 25 13 26 55 17 27 20 13 28 35 Ifi 29 30 Hi 30 45 17 31 20 13 .32 25 14 33 25 14 34 30 15 35 25 15 (90 percent) of the fish. Other monogenetic trematodes recovered were Octomacrwn lan- ceatinn, found in one (2 percent) fish, and PcUucidhaptor alahauius, foimd in six (10 percent) of the fish. Metacercariae of two digenetic trematodes were also recovered. Neascus sp. was found in 25 (43 percent) fish and Clinostomum marginatum was found in two (3 percent) fish. Cystidicola sp., a nema- tode, was found in one (2 percent) fish and Monobothrium hunteri and Isoglaridacris hex- acotyle, both caryophyllid cestodes, were found in 29 (50 percent) and 28 (48 percent) fish, respectively. Three protozoans were re- covered from the fish. Myxosoma sp. was found in 11 (19 percent) fish, Myxidiiim sp. in four (7 percent), and Tridiondina sp. was found in 20 (34 percent) of the fish examined (Table 3). Table 1 continued. No. Species of fish Weight Stream (gm) Length (TL) (cm) 36 C. discobolis 37 38 .39 40 41 C. latipinnis 42 43 44 45 46 47 48 49 .50 51 52 53 54 55 56 57 " 58 59 C discobolis 60 61 62 63 64 65 66 67 68 69 C. latipinni.s 70 La Verkin Creek 30 15 45 17 15 11 20 13 20 13 115 24 180 27 185 27 230 30 205 29 260 32 210 30 125 25 185 29 260 31 225 24 110 24 195 23 130 25 170 28 125 24 105 24 85 22 Fremont Hiver 20 14 10 11 35 16 35 17 20 14 30 16 5 10 25 15 10 11 30 15 5.30 40 350 37 152 Great Basin Naturalist Vol. 40, No. 2 Differences in protozoan parasite load from the two species of fish taken from La Verkin Creek are as follows: Myxosoma sp.. 25 percent bluehead suckers and 6 percent flannelmouth suckers; Myxidium sp., per- cent bluehead suckers and 22 percent flannelmouth suckers; Trichodina sp., 80 per- cent bluehead suckers and 22 percent flannelmouth suckers were infected. For the metazoan parasites, G. elegans was fomid in 90 percent of the bluehead suckers and 89 percent of the flannelmouth suckers, Octoma- cntm lanceatum and P. alahamus were found exclusively on bluehead suckers, 3 percent and 6 percent, respectively. Neascus sp. was found on 28 percent of the bluehead suckers and 78 percent of the flannelmouth suckers, but C. marginatum was found only in 11 per- cent of the flannelmouth suckers. The flannelmouth suckers have a higher incidence of both species of caryophyllid cestodes. MonobotJirium hunteri was found in 38 per- cent of the bluehead suckers and 78 percent of the flannelmouth suckers, and /. hexacotyJe was found in 33 percent of the bluehead suckers and 83 percent of the flannelmouth suckers. The nematode Cystidicola sp. and the leech Piscicola sp. were symbiotic exclusi- vely to the bluehead suckers. Three percent of the fish were infected with each of these two parasites. One species of parasite was recovered from T.\BLE 2. Number and percentage of bluehead and flannelmouth suckers parasitized from La Verkin Creek and the Fremont River. Host species Total fish sampled Total fish parasitized La Verkin Creek fish parasitized* 37 ( 93%) 18 (100%) .55 ( 95%) Fremont River fish parasitized^ Bluehead sucker Flannelmouth sucker Total 50 20 38 ( 76%) 20 (100%) 58 ( 83%) 1 ( 10%) 2 (100%) 3 ( 2,5%) *58 fish examined from La Verkin Creek (40 bluehead and 18 flannelmouth suckers). ''12 fish examined from the Fremont River 1 10 bluehead and 2 flannelmouth suckers). Table 3. Parasites identified from 58 suckers from La Verkin Creek. Parasite species Number and percentage of Species of fish fish positive Bluehead Flannelmouth 11 (19) 10 (25%) 1 ( 6%) 4( 7) ( 0%) 4 (22%) 20 (34) 16 (80%) 4 (22%) 52 (90) .36 (90%) 16 (89%) 1( 2) 1 ( .3%) ( 0%) 6(10) 6 (15%) 0( 0%) 25 (43) 11 (28%) 14 (78%) 2 ( 3) 0( 0%) 2(11%) 29(50) 15 (38%) 14 (78%) 28 (48) 13 (3.3%) 15 (8,3%) 1( 2) 1 ( 3%) 1 ( 0%) 1( 2) 1 ( 3%) ( 0%) Protozoans Myxosoma sp. Myxiditnn sp. Trichodina sp. Trematodes Gijrodactijlufi elegans Octomacrum lanceatutn Pellucidhaptor alahamus Postodiphstom am mit^ im iim Clinostomiim marginatum Cestodes Monobothritim hunteri Isoglaridacris hexacotyle Nematodes Cystidicola sp. Leeches Piscicola sp. June 1980 Breinholt, Heckmann: Fish Parasites 153 the fish examined from the Fremont River (Table 4). Gyrodactyhis elegans was identi- fied in 25 percent of the fish examined. Ten percent of the bluehead suckers were in- fected with this inonogentic trematode, and 100 percent of the flannelmouth suckers were infected. Table 5 lists the preferred tissue in the host for each parasite. These parasites were found in onlv five areas of the fish. Eight of the par- asite species were fomid on the external sur- face and only four were found in more than one area. Discussion There are habitat and host variations for the parasitofauna from suckers taken from La V'erkin Creek and Fremont River. The ca- tostomids from La V'erkin Creek were more heavily parasitized, both in the number of fish infected and in the number of species en- countered, than were the fish from Fremont River. Explanation for these differences may be attributed to many factors, such as water qualitv and macroinvertebrates. Water chem- istr\ for the two streams was found to be sim- ilar except during spring rvmoff. Oligocheates, which usually act as the in- termediate hosts for caryophyllid tapeworms (Mackiewicz 1972), occur in both streams. Recent studies listed 14,203 (Winget and Baumann 1977) and 882 (Heckmann 1976) oligocheates per m^ in La Verkin Creek and Fremont River, respectivelv, where the fish for this studv were obtained. The number of oligocheates should not have caused the dif- ference in parasite load because infected worms would have been ingested by fish from both streams. Milbrink (1975) correlates Table 4. Parasites identified from 12 suckers from the Fremont River. Number and percentage of fish positive Species offish Parasite species Bluehead suckers Flannelmouth suckers Trematodes Gyrodactyhis elegans 3 (25) 1 (10%) 2 (100%) Table .5. Location in host of parasites found in fish from La Verkin Creek and the Fremont River. Parasite species Surface Gills Operculimi Intestine Gall bladder Protozoans Mijxosotna sp. MyxidiuDi sp. Trichodiud sp. Trematodes Gyrodactyhis ch-gans Octomacrum hmccatiiiu PcUucidJiaptor ahihamus Chnostomuin mart^inatiiui Cestodes Monohothriuin hunteri Isogla ridac ris h cxacoty Ic Nematodes Cystidicola sp. Leeches Piscicola sp. = present o = not present 154 Great Basin Naturalist Vol. 40, No. 2 the caryophyllid worm burden of fish with the number of infective oligocheates con- sumed. If the Fremont River contained in- fected oHgocheates, some of the fish sampled should have been infected. The geographical location of the two streams may have caused the difference in parasite load. Parasites can be found in one area but not in another even though both have similar aquatic characteristics. Myxo- soma cerebralis, a myxosporidan parasite which caused whirling disease in trout, has been reported in eight states (American Fish- eries Society, 1974). Whirling disease has not spread to the other states even though suit- able habitats exist. Diplostomum spathacewn, the eye fluke of fish, has been reported in some areas of Utah but not others (Palmieri, Heckmann, and Evans 1976). Most parasites have some effect on the health of the host (Olsen 1974). The fish sam- pled from the Fremont River were infected with only one species of parasite, Gijro- dactylus elegans, and the incidence of that parasite was low in comparison to infected fish from La Verkin Creek. The most com- mon parasite found on the fish from La Ver- kin Creek is G. elegans. This organism was on the surface and occasionally in gill scrapings. Large numbers of G. elegans can cause dam- age to the fish by physical blockage of the gill surface, thus interfering with the gas ex- change area (Hoffman 1967). Other mon- ogenetic trematodes, Octomacrum lanceatum and Pellitcidhaptor alahamiis, and the pro- tozoan, Trichodina sp., are capable of causing similar problems in the fish. These parasites were not foimd in great enough quantities to pose a threat at the present time. The two myxosporidans, Myxosonia sp. and Myxidium sp., are capable of encysting and destroying tissue. However, no cysts were found. Thus, it is assumed that these myxosporidans are causing little damage to the fish. The fact that no hemoflagellates were found dining the course of this study does not disprove their existence in these fish, because some of these parasites have seasonal fluctua- tion. Only one nematode, Cystidicola sp., was recovered from all fish examined in the sur- vey. This round worm is not detrimental to the fish unless it is found in high numbers (Hoffman 1967). The other intestinal hel- minths, Monobothrium hunteri and Isoglari- dacris hexacotyle, are adult cestodes that usu- ally cause little damage to the definitive host. High numbers (200 plus) result in mechanical blockage or cause nutritional deficiencies (Mackiewicz 1972). The adult worms adhere to the intestinal lining by means of suckers. There is little intestinal damage by individual cestodes because the scolex is unarmed. The metacercariae of Neascus sp. and Cli- nostomwn marginatum were recovered from suckers from La Verkin Creek. For these trematodes, the metacercarial stage is usually encysted in the second intermediate host and does not cause damage through migration (Hoffman 1967). Unless the metacercaria is encysted in vital organs, such as a parasite in the eye lens (D. spathaceum), it does not present a pathogenic health problem to the fish. (Neascus sp. was observed encysted in the fins and gills. The cysts found in the gills were not numerous and did not appear to in- terfere with gas exchange for the fish. The leech, Piscicola sp., did not present a current problem to the fishing that di^ly one specimen was recovered. Leeches are period- ic feeders and should not attach permanently to the host. The major problem with leeches on fish is due to large numbers on one host or tlie transmitting by hemoflagellates (l:)lood parasites) (Hoffman 1967). No blood parasites were found during this survey. None of the fish expired during the journey from their natural habitat, approximately 400 km, to holding tanks. Thus, it is assumed that the effects of all the parasites on the suckers were not evident when the fish were placed under stress of capture and transportation. Limited host specificity is demonstrated by the parasites recovered in this survey. Most of the parasites encountered have been re- ported in other species of fish (Hoffman 1967). Species of Myxidium, Myxosoma, Trichodina, Cystidicola, and Piscicola have all been reported in trout. The parasitic species found in salmonids may be different than those found in the suckers. Gyrodactylus ele- gans and P. minimum have also been report- ed in salmonids. Octonuurum lanceatum has been reported in the catostomids, Catostomus teres, C. commersoni, C. macrocheilus, and Erimyzon secetta, the cyprinids, Mylocheilus June 1980 Breinholt, Heckmann: Fish Pahasites 155 caurinus (peamouth), and Notropis corntitits (common shiner). FclhicitUuiptor dldluiiniis has been reported in Ictiohus htihalus, the smalhnoiith buffalo (Chien and Rogers 1970). The caryophyllid tapeworms. A/, hunteri and /. hcxncoti/lc, common to C. discobolis and C. Uitipinnis, have both been reported from oth- er catostomids (Hoffman 1967). Because the parasites recovered in this study have been reported in other species of fish, it is possible that these parasites may infect game fish or commerciallv cultured fish. In the case of the listed digenetic trematodes, infected birds can fly from one body of water to another and "seed" other streams and ponds. Also, currents can carrv infected fish and other in- termediate hosts downstream to contaminate the lower drainage system. Thus, potential infections of other fish in the same stream could threaten endangered species such as the woundfin minnow and the Virgin River spinedace, which are also found in La Verkin Creek. The identity of one of the hosts from La Verkin Creek is doubtful. Originally, it was classified as a chiselmouth sucker (Sigler and Miller 1963). Later studies considered this ca- tostomid a desert sucker (C. clarki), which is still a valid species (Bailey et al. 1970). Then, with the taxonomic revision of some of the members of the catostomid family, Pan- tosteus delphinius, the bluehead sucker, and P. virescens, the green sucker, were combined to form C. discobolis (Bailey et al. 1970). Af- ter this revision some investigators have con- sidered the fish as C. discobolis. Because of the anatomical similarities between C. dis- cobolis and C. clarki and the activities and feeding habits, the two could be considered similar. It may also be concluded that they could harbor similar parasites even if they are two distinct species. Of the two suckers, C. latipinnis is more selective in its feeding habits than is C. dis- cobolis. Winget and Baumann (1977) report- ed stomach contents of the flannelmouth sucker contained seeds, identifiable plant matter, and dipteran larvae; stomachs of the bluehead sucker contained detritus, uniden- tifiable plant matter, and very few macroin- vertebrates. The difference in feeding habits of these two fish is probably the reason for the difference in resident parasite species. The parasites that these two fish have in common may be due to the ingestion of a common intermediate host. Even though C. latipinnis is more selective in its feeding habits, it still would ingest detritus and other material due to its feeding technique. The parasites identified in this survey are not unexpected. Although no parasitic sur- veys of C. discobolis and C. latipinnis have been reported, Mi/xosorna sp., Myxidiian sp., Trichodina sp., G. elegans, O. lanceatum, P. alaJianius, P. mininiitm, C. marginatum, I. Jiexacotyle, M. hunteri, Cystidicola sp., and Piscicola sp. have all been reported from suckers (Hoffman 1967). Literature Cited American Fisheries Society. 1974. Suggested procedures for the detection of certain infectious diseases of fishes. U.S. Department of the Interior, Washing- ton. D.C. Amin, O. M. 1969. Helminth fauna of suckers (Catosto- midae) of the Gila River system, Arizona. II. Five parasites from Ccito.'itomus spp. Amer. Midi. \at. 82(2): 429-443. 1974. Intestinal helminths of the white sucker, Catostomus coninicrsoni (lacepede) in southeast Wisconsin. Proc. Helminthol. Soc. Wash. 41(l):81-88. Bailey, R. M., J. E. Fitch, E. S. Herald, E. A. Lachner, C. C Lindsey, C. R. Robins, and W. B. Scott. 1970. .\ list of common and scientific names of fishes from the United States and Clan- ada. .\merican Fisheries Society Special Pub- lication No. 6. Washington, D.C. Chien, S. M., and W. Rogers. 1970. Four new species of monogenetic trematodes. genus PelliicicUuiptor. ftoin fishes of the southeast United States. J. Par- asitol. 56(3):480-485. Clifford, T. S., and S. Facciani. 1972. Philoinetra nodulosu in Wvoming white suckers. Prog. Fi.sh- Cult. .34(4):23.5-2.36. ' Daly, J. J., a.nd D. L. De(;ivsti. 1971. Tn/pdnosoiiia ca- tosiomi n. sp. from the white sucker Cdtostoiiiiis cominersoni. J. Protozool. 18(.3):414-417. Dechtiar, .\. O. 1969. Two new species of monogenetic trematodes (Treniatoda: Monogcnca) from nasal cavities of catostomid fishes. J. Fish. Wvs. Board Can. 26(4):86.5-869. Eddy, S. 1969. The freshwater fishes. Win. C Brown C. Puhl., Dubuque, Iowa. IL\THAWAY, R. p., and J. C. Herlevich. 1973. Gyro- dactyhis stahlcri sp. n. with new host and locality records for species of Giircddiliihis. J. Parasitol. .59(2):801-802. Hayunga, E. C, and .\. J. Grey. 1976. Cystobracluis meyeri sp. n. (Hirudinea: Piscicolidae) from Ca- tostomtis commersoni Lacepede in North Ameri- ca. J. Parasitol. 62(4):621-627. 156 Great Basin Naturalist Vol. 40, No. 2 Heckmann. R. a. 1976. Aquatic habitat evaluation of the Fremont, Muddy, and Dirt\ Devil rivers and Caine Springs and Pleasant Creek. Westinghouse Corporation contract .\o. P. O. ESD- 151-76. HoFFM.w. G. L. 1967. Parasites of North American freshwater fishes. University of California Press, Berkeley and Los .\ngeles. Mackiewicz, j. S. 1963. Monobothhum httntcri sp. n. (Cestoidea: Carvoph\llaeidae) from Catustomus commersoni (Lacepede) (Pisces: Catostomidae) in North America. J. Parasitol. 49(.5):723-7.30. 1972. Caryophvllidea (Cestoidea): \ review. Exp. Parasitol. 31(3)':417-512. MiLBRi.NK, G. 1975. Population biolog> of the cestode Canjophyllaeus laticeps (Pallas) in bream. Ahra- mis brama (L.), and the feeding of fish on oligo- cheates. Institute of Freshwater Research Report No. 54:36-51. Olsen, O. W. 1974. .\nimal parasites: Their life cycles and ecologN . University Park Press, Baltimore. Palmieri, J. R., R. .\. Heckm.\.xn. and R. S. Evans. 1976. Life cvcle and incidence of Diplostomum spatho- ceiim Rudolphi (1819) (Trematoda: Diplostoma- tidae) in Utah. Great Basin Nat. 36(l):86-96. Pfliecer. W. L. 1975. The fishes of .Missouri. Missouri Department of Conservation. Jefferson City. Price, C. E., .and H. P. .\r.\i. 1967. The monogenean parasites of Canadian freshwater fishes. Canadian J. Zool. 45(6/2)123.5-1245. ScHELL, S. C. 1974. Two new genera and three new^ spe- cies of allocreadiidae trematodes (Digenea; .\1- locreadiidae) from freshwater fishes. J. Parasitol. 60(2):24.3-246. Sigler, W. F., .a.\d R. R. Miller. 1963. Fishes of Utah. Utah State Department of Fish and Game. Salt Lake City. Threlfall, W., a.nd G. Hanek. 1970. Metazoan para- sites, excluding monogenea, from longnose and white suckers. J. Fish. Res. Board Can. 27(7): 1317-1319. N'oTH, D. R., .\ND O. R. Larson. 1968. Metazoan para- sites of some fi.shes from Goose River, North Da- kota. Amer. Midi. Nat. 79(l):216-224. White. G. E. 1974. Parasites of the common white suck- er (Catastomiis commersoni) from the Kentuckv River drainage. Trans. Amer. .Micros. Soc. 93(2):280-282. WiNGET, R. N., AND R. W. Bal'm.a.nn. 1977. Virgin River, Utah-.\rizona-Nevada, aquatic habitat, fisheries, and macroinvertebrate studies. Center for Health and Environmental Studies, Brigham Young Uni- versitv, L'tah. WiNGET, R. -M., A.ND .M. K. Reichert. 1976. .\quatic habitat inventory in the hot desert eis area, Utah. U.S. Bur. Land Man., Contract No. YA-512-CT6- 77. SOIL WATER WITHDRWVAL AND ROOT DISTRIBUTION UNDER GRUBBED, SPRAYED, AND UNDISTURBED BIG SAGEBRUSH VEGETATION David L. Stiirges' .Vbstract.— Seasonal depletion by vegetation where sagebrush was selecti\elv removed bv gnibbing and where sagebrush was sprayed with 2,4-D was 33 and 12 percent less, respectively, than that for undisturbed big sagebrush vegetation in the surface 122 cm of soil. Differences were located primarilv below 61 cm in vegetation gnibbcd the previous fall and below 91 cm in vegetation sprayed three years previously. Total root weights under gnibbed and sprayed vegetation were 29 and 16 percent less, respectively, than for unclisturbed big sagebrush vegetation. Total herbaceous production by grubbed and sprayed vegetation was 69 and 43 percent less, respectiveh, than production l)\ undisturbed vegetation. Big sagebrush [Artemisia trident at a) is commonly controlled with herbicides, me- chanical methods, or fire to increase livestock forage production. Pheno.xy herbicides such as 2,4-D damage forbs as well as sagebrush, so that the net effect of spraying is to favor grass productivity.- Burning or mechanical sagebrush control techniques, however, do not selectively favor grasses. Herbaceous pro- duction commonly doubles or triples bv the .second or third year after sagebnish removal. The shift from a shrub to a herbaceous- dominated vegetation produces other ecolog- ic and hydrologic changes. This studv was made to quantify differences in the soil water regime and in root biomass between undis- turbed big sagebnish vegetation and (a) her- baceous vegetation three years after spraying with 2,4-D and (b) herbaceous vegetation from which only big sagebnish was removed by mechanical means the previous fall. Infor- mation about herbaceous productivity was also collected. LlTER.\TL RE ReVIEW Ghanges in the soil water regime after sagebrush control are strongly influenced by rooting characteristics of sagebnish and her- baceous species. Roots of basin big sagebnish (A. t. sub. tridcntata) and mountain big sage- brush (A. t. vaseyana) commonly extend about 2 m deep and have a maximum lateral spread from the trunk of 1.5 m (Goodwin 1956, Cook and Lewis 1963, Tabler 1964, Hull and Klomp 1974, Sturges and Trlica 1978). Most roots are in surface soil where maximum spread occurs. About 60 percent of total root length (Tabler 1964) and 85 per- cent of total root system weight were present in the surface 61 cm of soil, with only about 4 percent in soil below 91 cm (Sturges and Trlica 1978). The principal soil water reservoir utilized by isolated mountain big sagebnish plants ex- tended 0.9 m laterally from the trunks and 0.9 m deep (Sturges 1977b). Tlie plants utiliz- ed water from surface soil adjacent to the trunk early in the growing season, but use- zones shifted outward and downward later in the summer as water adjacent to the trunk was depleted. .Appreciable water uptake was detectable until early in .\ugust. Tabler (1968) and Sturges (1977a) found that seasonal soil water withdrawal was re- duced after spraying sagebnish vegetation 'Rocky Mountain Forest and Range Experiment Station, Laramie. Wyoming 82070. Central headquarters is at Fort Collins in cooperation with Colorado State University; research reported here was conducted at the station's Research Work Unit at Laramie, in cooperation with the University of Wyoming. Portions of the research were supported by the Bureau of l^nd Management. U.S. Department of the Interior. This article reports research involving pesticides. It neither contains recommendations for their use nor implies that the uses discussed here have been registered. .\ll uses of pesticides must be registered by appropriate state and/or federal agencies before they can be recommended. Use all pesticides selecti- vely and carefullv, read and follow the directions on the label. 157 158 Great Basin Naturalist Vol. 40, No. 2 with 2,4-D on sites with deep soils that were fully recharged by snowmelt. This reduction was located almost entirely below 91 cm as depletion -of surface soil water by sprayed vegetation sometimes exceeded depletion by untreated vegetation. Water depletion in sur- face soil increased the first few years after treatment as herbaceous vegetation respond- ed to release from sagebrush competition. Other studies also detected an increasing moisture draft from surface soil with time (Hyder and Sneva 1956, Cook and Lewis 1963, Shown et al. 1972). Herbaceous production was measured in most soil moisture studies. Grass production doubled the year after spraying mountain big sagebrush and was 2.6 times higher than un- treated vegetation three years after treat- ment (Sturges 1977a). Shown et al. (1972) found that usable forage production in- creased 300 percent compared to pre- treatment conditions once a planted grass stand became established. Hyder and Sneva (1956) found the increase in grass production to be the same whether big sagebrush was controlled by spraying or by grubbing. Total herbaceous production increased the most where sagebrush was grubbed, because forbs were damaged by the spray. Study Area The study was performed at the Stratton Sagebnish Hydrology Study area 29 km west of Saratoga in south-central Wyoming. The experimental site is at an elevation of 2,225 m and lies on a north-facing slope in a mod- erate snow catchment zone. Annual precipi- tation is about 500 mm, with two-thirds of the total falling as snow. Precipitation be- tween 1 June and 30 September averages 114 mm. Sufficient snow usually accumulates to completely recharge the soil mantle. Soils de- veloped in place from sandstone and belong to the Argic Cryoboroll great soil subgroup. A dense stand of mountain big sagebrush, un- derlain by a productive understory of bunch grasses-primarily Idaho fescue (Festuca ida- Jwensis), bluegrass (Poa spp.) and needle- grasses {Stipa spp.)-was present before study initiation. The site had been grazed by sheep, but no grazing occurred during the study. Methods Work began in 1968 with a study that uti- lized 14 0.4-ha experimental units arranged in seven blocks to determine how the soil wa- ter regime would be affected by spraying big sagebnish (Sturges 1977a). One experimental unit within each block was sprayed with 2,4- D in 1970; the other unit remained un- treated. Experimental units from three of the seven blocks were used in the current study. In October 1972, smaller plots 23 m long and 10 m wide were established on either side of the common border between sprayed and un- sprayed vegetation (Fig. 1). These plots were used to obtain soil cores and to create the grubbed sagebrush vegetative condition. Sagebnish was grubbed from four circular areas 6.1 m in diameter by cutting plants at or slightly below the ground surface. Grubb- ing was done in the fall of 1972, when vege- tation was dormant, thereby minimizing damage to residual herbaceous vegetation and insuring that herbaceous vegetation would be as comparable as possible to that within the undisturbed sagebnish stand when study measurements began the following spring. Soil Water Measurements Soil water content was measured with a neutron-scattering soil moisture meter at four randomly located access tubes on each exper- imental unit. Access tubes within grubbed vegetation were installed at the center of each cleared circle in October 1972. If one assumes that big sagebnish has a maximum lateral root spread of 1.5 m, these tubes were surrounded by a volume of soil at least 1.5 m in radius devoid of live sagebrush roots. Moisture measurements began 31 May 1973 upon completion of snowmelt and con- tinued at biweekly intervals until 19 Septem- ber 1973, when vegetation was dormant. Measurements were taken at eight depths: 15, 30, 46, 61, 76, 91, 107, and 122 cm. The manufacturer-supplied calibration curve re- lating field neutron count (expressed as a per- centage of shield count) to volume moisture content was applied to all data except that collected at 15 cm. Here, a correction was made for escape of neutrons into the atmo- June 1980 Sturges: Big Sagebrush 159 sphere using a polyethylene shield technique similar to Pierpoint's (1966). Root Weights Soil cores for sampling root weight were obtained at four random locations within the small plots that straddled the common border between sprayed and midisturbed sagebrush vegetation. The cores were collected in Sep- tember following the final soil water mea- surement (Fig. 2). Each core was 7.6 cm in diameter and 122 cm long. The cores were collected in 15-cm increments using the de- vice described by Brown and Thilenius (1977). Each sample site in grubbed vegeta- tion was located within 2.4 m of an access tube, a minimum of 0.6 m from the surround- ing sagebrush cover. Soil cores were placed in plastic bags and frozen on the day of col- lection. After thawing, core segments were individually washed in a core-washing ma- chine (Brown and Thilenius 1976) to isolate root matter from soil. Roots were oven dried for 24 hours at 70 C and weighed on an ana- lytical balance. It was not possible to dis- tinguish between live and dead roots, but woody sagebrush roots from cores taken within grubbed vegetation were discarded before samples were weighed. Herbaceous Production Above-ground herbaceous productivity was measured by clipping 12 randomly lo- cated plots within each experimental unit as grasses matured in mid-July. In grubbed veg- etation, three production plots were placed Undisturbed sagebrush vegetation Sprayed sagebrush vegetation 10 m 23nn 64 nn 64 nn X Access tube on 0.4 ha plot fj Circular area where sagebrush grubbed + Access tube in grubbed vegetation Fig. 1. The experimental design for one block showing the 0.4-ha experimental units of undisturbed and sprayed sagebrush vegetation and smaller plots where sagebrush was grubbed. Soil moisture data, soil cores, and herbaceous productivity information for the grubbed treatment were obtained on the small plot. 160 Great Basin Naturalist Vol. 40, No. 2 at random within 2.4 m of access tubes. Veg- etation was harvested to a 1 cm stubble height from plots 30.5 cm wide and 61 cm long. Vegetative matter was separated into grass, forb, or sagebnish components and placed in paper bags when harvested. Only leaves and herbaceous stem material were in- cluded with sagebnish herbage. Vegetation samples were subsequently dried at 105 C for 24 hours and weighed. Selected big sagebrush and productivity characteristics were measvired in 1969 on the 0.4-ha experimental units, the year before spraying (Table 1). No statistically significant differences before treatment were present. Big sagebrush contributed 76 percent of aboveground herbaceous production while grasses contributed 20 percent and forbs 4 percent. About one-third of the area was cov- ered by the live, leafy portion of the sage- bnish canopy. Sagebrush plants had an aver- age height of 34 cm and an average crown area of 7 dm-. Statistical Analysis Soil water withdrawal and root weight dif- ferences among the three vegetative condi- tions were tested for statistical significance by variance analysis utilizing a split-plot de- sign. Experimental units (whole units) were arranged in three randomized 'blocks, and the eight measurement depths served as subunits. Analyses were based on average plot values determined from the four replicated mea- surements on the plot. Variables analyzed were the change in soil water content be- tween successive sampling dates, the seasonal change in soil water content, and root weight. Herbaceous productivity data were analyzed with a randomized block design. Results Soil Water Depletion Soil under undisturbed and sprayed vegeta- tion was completely recharged by snowmelt on the first measurement date, but only to 61 cm under grubbed vegetation (Fig. 3). At the end of summer, water content in the surface 46 cm of .soil was similar for all treatments. Below 46 cm, progressively more water re- mained in soil under grubbed vegetation compared to undisturbed sagebnish vegeta- tion, but appreciable differences between sprayed and undisturbed vegetation were present only below 91 cm. Seasonal water withdrawal by undisturbed, sprayed, and grubbed vegetation was 24.3, 21.4, and 16.2 cm of water, respectively, in the surface 122 cm of soil. These differences Table 1. Characteristics of vegetation on plots as- signed to sprav and undisturbed treatments in 1969, one vear before 2,4-D was applied. Sagebrush Undisturbed Sprayed Height (cm) 30° 37 Canopy area (dm') 6 8 Canopy intercept (%) 31 32 Density (number/ha) 57,(X)0 52,000 Hebbac;eous production (kg /ha) Sagebrush 926 1095 Grass 249 290 Forb 53 60 Total 1228 1445 "Differences between treatment means were not significant for any mea- surement parameter at the 0.05 level of probability. tig. 2. Root samples were obtained using a core sam- pler driven into the soil to a 122-cm depth by 15-cm in- crements. June 1980 Sturges: Big Sagebrush 161 were significant at the 0.01 probability level. Treatment differences did not accrue uni- formly through the soil mantle, but were con- centrated at deeper soil depths (Fig. 4). Be- tween 91 and 122 cm, depletion by grubbed and sprayed vegetation was 31 and 66 per- cent, respectively, of depletion by undis- turbed sagebnish vegetation. Treatment soil water withdrawal differ- ences between consecutive measurement dates were significant (p<0.05) only between 25 June and 10 July. The treatment x depth interaction term was significant during five of the eight measurement intervals, though, indicating that the three vegetative condi- tions were utilizing water differently from within the soil. For example, most of the dif- ference in depletion below 91 cm between 1 20 CL .^ D Q Undisturbed Sprayed Grubbed 76 cm 91cm 46 cm 107cm 61cm 122 cm -J I 1 i_ ilLlJI IlL 15 31 15 30 li .J .1 I nil. I -Id L 5 31 15 31 15 30 15 31 15 30 I III.. I .1 I 1. L 5 31 15 31 15 30 May June July Aug. Sept. May June July Aug. Sept. Fig. .3. Daily precipitation and soil water content in the surface 122 cm of soil for undisturbed, sprayed, and grubbed .sagebnish vegetation in the 197.3 growing season. 162 Great Basin Naturalist Vol. 40, No. 2 sprayed and undisturbed vegetation devel- oped after 25 July. Sagebrush remained phys- iologically active through the summer and flowered about 1 September, so that appre- ciable water usage continued all summer. Most grass and forb species had matured and set seed by early August, thus reducing the need for water by grubbed and sprayed vege- tation. Root Weights The average weight of roots obtained from soil cores extending 122 cm deep was 12.2, 10.2, and 8.7 g under undisturbed, sprayed, and grubbed vegetation, respectively. Nei- ther the treatment, nor the depth x treatment interaction term was statistically significant. Varying quantities of dead but undecayed root matter and other organic debris were in- cluded with sample material and could not be separated from live roots. Inclusion of ex- traneous matter probably accounted, in part, for the low statistical sensitivity of root mea- surements. Most of the weight of roots was located in surface soil (Fig. 4). Material from the surface 15 cm of soil ranged from 36 percent of total root weight in imdisturbed sagebRish vegeta- tion to 54 percent of total root weight in sprayed vegetation. Conversely, only 1 to 2 percent of root weight for each treatment came from the deepest sampling depth. Herbaceous Production Herbaceous production of undisturbed sagebrvish vegetation was about a third great- er in 1973 than in 1969, but composition of vegetation was similar both years. Treatment differences within sagebrush, grass, and total production herbage classes were highly sig- nificant (Table 2). The response by sprayed vegetation the third year after treatment was typical to that reported from other locations. Grass production was 2.6 times greater than production in imdisturbed sagebmsh vegeta- tion, but forb production was still depressed below pretreatment levels. Total herbaceous production by sprayed vegetation was only 57 percent as large as production by undis- turbed sagebnish vegetation, the increase in grass production not compensating for loss of sagebrush. Grass production increased 27 percent where sagebrush was grubbed the previous fall, but the increase was not statistically sig- nificant (Table 2). Total production was 31 percent as high as that by undisturbed vege- tation because of the loss of sagebrush. Discussion and Conclusions This study indicates the soil water regime in the surface 91 cm of soil is unaffected by sagebrush control once herbaceous vegeta- tion responds to release from sagebrush com- petition. However, below 91 cm, substantial reductions in seasonal withdrawal can occur as reported by Tabler (1968) and Sturges (1977a). The overall reduction in soil water depletion caused by grubbing sagebrush com- pares closely with that detected on the same 0.4-ha experimental units in 1970 when sage- bnish was sprayed. Grubbing decreased sea- sonal water withdrawal 33 percent in this study, and spraying reduced withdrawal from the surface 137 cm of soil 37 percent (from the spray date on 22 June through 30 Sep- tember). The year after spraying, a 17 per- cent difference in seasonal withdrawal was observed with grass production doubling in response to sagebrush removal. Reductions in moisture withdrawal are re- lated to decreased aboveground herbaceous productivity of treated vegetation. Produc- tivity in grubbed and sprayed vegetation was 31 and 57 percent as large, respectively, as that of undisturbed vegetation. Development of vegetation in years immediately following sagebrush control also influenced water with- drawal patterns. Seasonal depletion under gnibbed vegetation was less than that of un- disturbed sagebnish vegetation at all depths, but appreciable differences existed only be- Table 2. Aboveground herbaceous production (kg/ha) by undisturbed, sprayed, and grubbed vegeta- tion in 1973. Treatment Sagebnish Grass Forb Total Undisturbed °120P Spraved 1^ C; rubbed (1*^ 347* 918^ 442'' 86* Ifif' 67* 1634* 93^ 509^" "Treatment means having different letters within a column are signifi- cantly different at the 0.05 level o^probability. June 1980 Sturges: Big Sagebrush 163 o o cr 0-15 15 ,;yyw^.i...»,.i„.iiiiii„„„„„„„„y,^ . . . ^;M. npot weigh t •mi Depletion Undisturbed _| Sprayed t i Grubbed 15-30 30 30-46 46 E 46-61 61 o IP Q. Q 61-76 76 76-91 91 91-107 107 107-122 122 I 2 3 Root weight (g) Seasonal water depletion/15 cm soiKcnn] Fig. 4. Seasonal water depletion and weight of roots in soil cores 7.6 cm in diameter and 122 cm long unde undisturbed, sprayed, and grubbed sagebrush vegetation. 164 Great Basin Naturalist Vol. 40. No. 2 low 61 cm. Sprayed vegetation, unlike grubbed vegetation, had fnlh- responded to release from sagebrush competition and depletion did not become appreciabK less tlian that of imdistiu-bed vegetation mitil a 91-cm depth was reached. Reductions in treatment effect through time within soil 60-90 cm deep were described by Hyder and Sneva (1955), Cook and Lewis (1963), and Shown et al. (1972). The reduction in seasonal water use and in root weight caused b\' treatments are similar when e.vpressed as a percentage of values for undisturbed vegetation. Seasonal depletion was 33 and 12 percent less for grubbed and spraved vegetation, respectively, and root weights were 29 and 16 percent smaller on these same treatments. Similar agreement be- tween depletion and root weight did not exist for individual measurement depths (Fig. 4). Thus, root weight measurements do not veri- f\' or refute the hypothesis that root devel- opment by herbaceous species in the surface 90 cm of soil subsequent to sagebrush remov- al accoimts for increases in moisture use from this zone. Measurement of root length, rather than root weight, probably would have pro- vided a better measure of potential moisture draft because of the differences in morpho- logy of grass and sagebrush roots. Comparisons of seasonal moisture change and root weight with depth does indicate that deep roots are extremely important in extracting soil water, even though they com- prised a small part of root weight in soil cores (Fig. 4). Summer precipitation is usual- ly ineffective in replenishing soil water levels in the sagebrush zone, so that deeper soil be- comes an important water reservoir when surface soil dries. A progressive, downward shift of major water use zones in August was especially evident for undisturbed sagebrush vegetation (Fig. 3). Results of this and other soil water deple- tion studies indicate that control of big sage- bru.sh with methods that do not destroy all vegetation on lands with an adequate popu- lation of herbaceous species has a relatively small effect upon the soil water regime. Changes in the soil water regime can, at most, result in small increases of streamflow. This response will onl\ t)ccur on lauds where soils are deeper than 90 cm and soil water recharge exceeds that retiuired to fulK wet the soil mantle. The maxinuuu reduction in depletion will usually occur in the treatment \ear because of productivity increases bv herbaceous species in xears immediatelv after treatment. Consecjuentlv. justification for big sagebiiish control must rest on the benefits derived from shifting site resources to species more desirable than sagebrush from a given land management perspective. Literature Cited Brown, G. R., and J. F. Thilenius. 1976. A low-cost machine for separation of roots from soil mate- rial. J. R;mge Manage. 29;506-507. 1977. .\ tool and method for extracting plant- root-soil cores on remote sites. J. Range Manage. 30:72-74. Cook, C. W., .\nd C. E. Lewis. 1963. Competition be- tween big sagebrush and seeded grasses on foot- hill ranges in Utah. J. Range Manage. 16:245-250. Goodwin. D. L. 1956. .\iitecological studies oi Artemisia tridentata Nutt. Unpublished dissertation. Wash- ington State Univ., Pullman. 79 pp. Hi LL, A. C, Jr., .and G. J. Klomp. 1974. Yield of crested wheatgrass under four densities of big sagebnish in southern Idaho. U.S. Dept. Agric. Tech. Bull. No. 1483. 38 pp. Hyder, D. N., a.nd F. A. Sneva. 1956. Herbage response to sagebrush spraying. J. Range Manage. 9:34-.38. Pierpoi.nt, G. 1966. Measuring surface soil moisture with the neutron depth probe and a surface shield. Soil Sci. 101:189-192. Shown. L. M., G. C. Lusby, and F. A. Branso.n. 1972. Soil-moisture effects of conversion of sagebrush cover to bunchgrass cover. Water Resour. Bull. 8:1265-1272. Sti rc;es, D. L. 1977a. Soil moisture response to spraying big sagebnish: A seven-year study and literature interpretation. USD.\ For. Serv. Res. Pap. R.\i- 188. 12 pp. Rocky Mt. For. and Range Expt. Stn.. Fort Collins, Colo. 19771). Soil water withdrawal and root character- istics of big sagebrush. Am. Midi. Nat. 98:257-274. SriRCES, D. L., AND .M. J. Trlica. 1978. Root weights and carbohydrate reserves of big sagebrush. Ecol- ogy 59:1282-1285. Tabi.er, R. D. 1964. The root system oi Arteniisia triden- tata at 9,500 feet in Wyoming. Ecology' 45:633-636. . 1968. .Soil moisture response to spraying big sage- brush with 2,4-D. J. Range Manage. 21:12-15. SWARMING OF THE WESTERN HARVESTER ANT. POGOXOMYRMEX OCCIDEXTALIS Dorald M. Allred' ,\bstract.— The swarming and mating of harvester ants was observed in Utah in July 1979. Workers groom the alate forms outside the mound before swarming occurs and are highly aggressive in protecting them. Mating pairs apparently are not disturbed by other ants. Fertile females likely use moving vehicles that extend their dispersal. Few detailed accounts of the swarming ac- tivities of harvester ants are existent in the literature. Wheeler (1910) reported the swarming of harvester ants in the desert along the Colorado River. Michener (1942, 1948) noted swarming and mating of Pogono- mymiex califomicus and P. barbatus. Strandt- mann (1942) recorded the mating activities of P. Comanche, and Chapman (1957) reported elevational swarming of P. occidentalis on mountain tops in five states. During the latter part of June and all of July in 1979, I traveled extensively over Utah collecting ants. In most cases when harvester ants were taken, I partially excavated each mound from which I collected to determine the presence of immature and winged forms. Although winged males and females were present in the majority of the moimds during this period, swarming was not seen until the latter part of July. On July 23 I stopped at about 11:30 a.m. (Mountain Daylight Time) to collect from an area of abimdant. large mounds one mile west of Elberta. Utah County, Utah, along- side highway US6 at an elevation of 5400 ft in a sagebrush-rabbitbmsh habitat [Artemisia tridentata-Chrysothamnus nauseosus). -\s I approached a large moiuid on that wanii, sunny day, an area of one-half square foot around two enlarged, south-facing openings was literally red with a mixture of \\orkers, winged males and females. I would have had difficulty finding an open space within tlie mass of ants where I could ha\'e touched the ground with a pencil. The winged forms were relatively inactive, none in flight, and the workers seemed to be grooming and at- tending them. As I approached the moumd to aspirate a sample of ants, the majority of the winged females and some males quicklv en- tered the nest openings. Some of the males, however, remained immobile outside the mound as though mesmerized bv the sToom- ing activities of the workers. The workers on the fringe of the mass immediatelv began ag- gressive tactics toward me in much more of a frenzied movement than I had heretofore en- countered with workers when winged forms were not outside the nest. After I had taken my sample. I returned to the car to record the data. Five minutes later I again visited the mound to see if the winged forms had left the burrow. The air was filled with fl\ing ants. At the mound the workers were no longer congregated around the openings, but were scurrvin^ around, near, and over the mound. .\ few winded forms were crawling around on the mound. I was quickly deluged with flNing ants, and a sting on my leg. presimiablv from a winged female, stimulated m\" hastv retreat to the car. where I quickly closed all windows. The outside of the car was soon covered with winged ants that were mating. Females seemed to be much more abundant than males. This was consistent \\ith m\ findings wherever I had excavated mounds through- out the state during Jime and July. In mating, the male moimted the female 'Department of Zcxilog), Bricham Vount; I'niversit) . Privo, I'tah S46Q2. 165 166 Great Basin Naturalist Vol. 40, No. 2 dorsally, clasped her around the thorax with his legs, and bent his abdomen strongly downward to contact her gentitalia. She si- multaneously bent her abdomen slightly up- wards to facilitate contact. Once joined, the partners sometimes assumed different posi- tions than described above, frequently both establishing leg contact with the substrate on which they were resting, although maintain- ing abdominal junction. Other crawling ants frequently came in contact with a mating pair, but the contact was brief. At no time did I observe other males remaining with mating pairs as described by Strandtmann (1942) for P. Comanche. Copulation lasted for perhaps 20 to 40 seconds, whereupon the two sexes immediately separated, with the male the first to fly away. No case was observed where either the male or female used its mandibles to grasp or chew on the other as described by Strandtmann (1942) for P. Com- anche, and by Michener (1948) for P. har- batus. About 12:.30 p.m., after I had observed the mating activities of numerous pairs and the numbers of ants crawling on the car and fly- ing in the area had considerably diminished, I returned to the mound. Few flying ants re- mained in the air in its vicinity, but a few winged forms were crawling around the mound. Most of the workers had reentered the entrances, and those that remained out- side had apparently resumed their normal, slower speed of routine activity. I moved to another area of several mounds situated about 25 yards north of the site of my previous observations. The ants of several of these mounds were in various stages of swarming activities. By periodical rotation between these, and with some excavations, I was able to summarize the overall activities associated with swarming of this species. When the males and females prepare to swarm, the workers enlarge the openings leading from the mounds, the winged ants and many workers leave the nest and congre- gate around its openings. The winged males and females are attended by myriads of workers who groom them as their bodies warm in the sun, and at the same time act in a protective capacity in relation to any would-be predators. Once the mating flight begins, the workers disperse and crawl around the mound for a few minutes, then re- enter the burrow. Normal worker activity outside the mound is resumed within a few minutes after the majority of the winged forms have departed. A few winged forms seem to linger around the mound, apparently hesitant to leave. Some also delay leaving the burrow to assume their flight of destiny. Swarming males and females seek some high point to mate, but many pairs mate on the ground. After mating, females migrate in all direc- tions to locate sites for establishing new colo- nies. Undoubtedly some of them hitch rides on moving conveyances such as cars, trucks, and trains. When I arrived at my home in Provo, 30 miles from and several hours after observing the swarming activity, a live winged female dropped from my car onto the driveway and crawled into the vegetation. Such methods of conveyance likely extend the dispersal of females over relatively great distances. Literature Cited Chapman, J. A. 1957. A further consideration of summit ant swarms. Canadian Ent. 89:389-95. Michener, C. D. 1942. The history and behavior of a colony of harvester ants. Scientific Monthly 55:248-58. 1948. Observations on the mating behavior of harvester ants. J. New York Ent. Soc. 56:239-42. Str.\ndtmann, R. W. 1942. On the marriage flight of Po- go(io»H/n)U'.v comanclic Wheeler. Ann. Ent. Soc. America .35:140. Wheeler, W. M. 1910. Ants: Their structure, devel- opment, and behavior. Columbia Univ. Press, New York, 663 pp. RELATIONSHIP BETWEEN ENVIRONMENTAL AND VEGETATIONAL PARAMETERS FOR UNDERSTORY AND OPEN-AREA COMMUNITIES William E. Evensoii', Jack D. Brothersoir, and Richard B. Wilcox' .XbstR'VCT.— Ten individnals from each of four tree species were selected, and their associated understory and ad- jacent open-area communities were sampled for both environmental and vegetational parameters, including light intensity, pH, litter depth, soil depth, and percentages of exposed rock, litter cover, living cover, shrubs, forbs, grasses, and annuals. The four tree species were ponderosa pine. Rocky Mountain juniper, Gambel oak, and snow- linish ceanothus. The studv site was in the lower Uinta Mountains about 10 miles east of Kamas, Utah. Correlations among the various biotic and abiotic parameters were examined. The interplay of these factors in differentiating the understory and open-area communities is discussed. Understanding the relationship of vegeta- tional patterns to environment is a primary goal of community ecology. One aspect of such relationships is the effect of overstory trees and shRibs on their associated under- story communities. In a previous report (Wil- cox, Brotherson, and Evenson 1981), we ex- amined the influence of four canopy species on their associated imderstory plant commu- nities in comparison to neighboring commu- nities in open areas outside the canopy in- fluence. The four canopy species were ponderosa pine {Piniis ponderosa Dougl.), Rocky Mountain juniper {Junipenis scopulo- nim Sarg.), Gambel oak {Quercus gambelii Nutt.), and snowbrush ceanothus (Ceanothus vehitinus Dougl.). Many previous studies have reported envi- ronmental and vegetational differences be- tween understories and open areas. Light in- tensity (del Moral 1972, Cline 1966, Blackman 1956) and spectral distribution (Federer and Tanner 1966) are known to strongly differentiate understory and open- area plant communities. Soil moisture and thickness of litter layer are also important factors (Anderson 1969, McQueen 1973), as is soil improvement due to nitrogen fixation by such common understorv plants as bitter- bmsh {Pursliia tridentata (Pursh) DC.) and snowbrush ceanothus (Wollam and Young- berg 1964, Rusel and Evans 1966, Webster, Youngberg, and Wollam 1967). Because these and other environmental modifications are influenced by the canopy species, cover (Anderson 1969, McQueen 1973, McConnell and Smith 1970) and diver- sity (Auclair and Goff 1971) of understory communities are strongly dependent on the canopy tree or large hebaceous species with which they are associated (Gordon 1962, Smith and Cottam 1967). The present study examines detailed rela- tionships between the various environmental and vegetational parameters measured under the canopies of fovir tree species and in nearby open areas. Study Area The study site is about 10 miles east of Kamas, Utah, along the Yellow Pine branch of Beaver Creek (Fig. 1). This area was cho- sen because of the homogeneity of the under- lying parent material (an alluvial outwash gravel bed) throughout the site, its constant slope and exposure, and its easy accessibility. The study site is an area of "zone jumbling" (Cottam 1930) and contains plant representa- tives from all life zones except lower sonoran. It is an area of highlv mixed vegetation, varying from Douglas fir {Pseudotsuga men- 'Department of Physics. Brigham Young I'niversity, Provo, Utah 84602. 'Department of Botany and Range Science. Brigham Young University, Provo, Utah 84602. 'Land Specialist, State of Utah, Room 440, Empire Buildnig, 231 East 400 South, Salt Lake City, Utah 84111. 167 168 Great Basin Naturalist Vol. 40, No. 2 ziesii (Mirb.) Franco.), white fir (Abies con- color Lindl.), and ponderosa pine to clumps of Gambel oak and snowbnish ceanothus. Also interspersed throughout the area are in- dividuals of lodgepole pine (Pinus contorta Dougl. ex Loud.), Rocky Mountain juniper, quaking aspen (Populus tretnuloides Michx.), and various other plant species. All can be found at the same elevation and in fairly close proximity. Because of its apparent uniformity, this site is especially well suited to measure the rela- tionship of environmental and vegetational parameters associated with understories of different tree species and nearby open areas. In such an area, the likelihood of factors oth- er than tree overstory affecting such relation- ships in a major way is small. "^76 00 y7(.SS Sfe iSrW Yellow Pine Campground Fig. 1. Map showing location of stiidv site. June 1980 EVENSON ET .\L.: PlANT EcOLOCY 169 Methods Ten individuals of each of four tree or shrub species (i.e., ponderosa pine, Rocky Mountain juniper, Gambel oak, and snow- bnish ceanothus) were chosen at random in the study area. Eight quadrats (0.25 m^) were placed aroimd each individual tree or .shrub. Four of these quadrats were placed inside the cylinder of the canopy, and four were placed outside the influence of the canopv. To elimi- nate bias, quadrats were consistently placed one at each direction of the compass. Quad- rats were subdivided into four equal units for species frequency measurements. Sample trees and quadrat sites were marked for relo- cation. Presence or absence of individual plant species in the understory was determined for all four subunits of each quadrat. All species rooted in the quadrat were recorded. Fre- quency of each plant species was determined by dividing the number of quadrat sub- divisions in which a species occurred by the number of subdivisions sampled. Total living plant cover and composition of plant cover by life form were measured at each quadrat using an ocular estimate method (Daviben- mire 1959). Light intensity was measured in foot-can- dles at each quadrat, and averages were com- puted for the understory and open-area quad- rats associated with each tree species. All readings were taken between 1200 and 1400 hours on cloud-free days, the last two days of the study. Soil pH was measured by the colormetric method in the field to avoid pH changes which can occur when soil is stored moist. Litter depth was measured at the center of all quadrats taken, and soil depth was deter- mined by the average of five penetrometer readings in each quadrat (one at each corner and one in the center). Correlations of all variables with each other were nm. Results .4nd Discussion Characteristics of the environment and the vegetation types associated with understory and open-area communities are summarized in Table 1 for the four canopy species. As ex- pected, light intensity values are consistently lower for the imderstory communities. Un- derstory communities have consistently high- er pH (more basic), except for ponderosa pine which shows an understory tending to be slightly more acidic than the open area. Both litter depth and soil depth are far great- er in tlie imderstory communities than out- side the canopies. Percent cover of litter is Table 1. Average measured values of environmental and vegetation parameters for understorv and open area comnumities ± their standard deviations. Pintis Ceanothus Juniper us Quercus ponderosa vchttintts scopulortim gamhelii Under- Open Under- Open Under- Open Under- Open story areas story areas story areas story areas Light intensity (foot candles) 117 ±52 225 ±65 123 ±29 .3.39 ±50 168 ±77 295 ±67 76 ±.32 242 ±67 Soil pH 6.3 ±0.2 6.4 ±0.1 6.6 ±0.1 6.4 ± 0. 1 7.4 ±0.6 6.4 ±0.2 6.6 ± 0.3 6.4 ±0.2 Litter depth (cm) 6.0±2.7 0.6 ±0.3 2.1 ±0.7 0.1±0.1 2.5 ±0.3 0.3 ±0.2 2.3 ± 0.6 0.2 ±0.2 Soil depth (cm) 10.5 ±2.6 5.3±1.1 6.7±1.7 4.4 ±1.2 6.1 ±.3.2 4.0 ±1.6 8.5 ±2.3 .5.7 ±0.8 % Living cover 20.2 ±6.0 25.8 ±6.8 20.0 ±4.7 31.5 ±7.6 28.0 ±19.1 .32.0 ±8.4 27.5 ±5.2 23.4 ±5.3 % Litter cover 83.6 ±6.0 39.0 ±11.6 45.6± 10.0 9.1 ±7.5 49.3 ±20.3 13.1 ±9.0 72.6 ±5.9 19.3 ±10.4 % Exposed rock 10.8 ±5.7 26.3 ±6.4 24.9 ±6.8 43.4 ±11.1 14.8±11.1 .32.4±8.1 14.6 ±9.9 28.5 ±4.7 % Shrubs 26.8 ± 1.3.0 27.1 ± 17.0 .37.0 ± 16.8 2.3.9 ± 1.3.2 48.0±21.9 22.9±17.8 27.7±11.9 18.4± 1,3.6 % Perennial forbs 5.2 ±7.4 11.4 ±11.3 18.9±9.7 .36.5± 12.9 8.0± 15.3 31.2± 14.9 15.6 ±9.2 27.4 ±12.8 % Perennial grasses 11.5 ±3.4 4.0 ±.3.1 13.9 ±8.5 7.1 ±5.9 16.9 ±1.3.9 10.4 ±9.0 22.8 ±10.2 14.0 ±9.3 % .\nnuals 56.6 ±11.9 57.5 ± 14.5 .30.2 ±12.5 .32.4 ±10.1 27.1 ±2.3.9 .3.5.6 ±2.5.5 .3.3.9 ±13.8 40.2 ±13.1 Average number of species/tree 10.5 11.9 14.2 14.9 10.9 14.3 12.7 14.0 Average number of species/ quadrat /tree 4.6 6.2 7.1 6.9 5.4 6.3 7.4 8.4 170 Great Basin Naturalist Vol. 40, No. 2 much greater in the understory, while per- cent of exposed rock is greater in the open- area communities. Percent living cover is higher in the open-area communities except for oak. Species preferences for understory and open areas were obtained by taking the dif- ference of total subquadrat occiu-rences for a species in canopy-covered and open stands and normalizing by dividing by the total oc- currences of that species in all stands. The re- sulting index nms from -1 to 1. Those species with the highest positive values are found most often under the canopy while those spe- cies with the greatest negative values are found most often in open areas. Table 2 shows species preference indices and total subquadrat occurrences for each species ob- served in the study (species are listed alpha- betically). Frequency is obtained by dividing the number of occurrences by 160, the total number of subquadrats sampled for each tree species within each type (canopy or open area). The species preference index was broken down into components relating to each of the four tree species as shown in Table 3. Each component was obtained by taking the differ- ence of the subquadrat occurrences for a spe- cies in canopy-covered and open-area stands Table 2. Number of ocxurrences of each species observed for each of the four canopy species; 160 subquadrats sampled in each category. Life form codes are f=forb, g = grass, c = cool-season or spring ephemeral but perennial (so cf = cool-season forb), sh = shrub, e = evergreen, a = annual, t = tree. Life Preference Understory stands Open-area stands Species form index Ceve Pipo Jusc Quga Ceve Pipo Jusc Quga Achillea millefolium Agoseris glauca Agropyron spicatum Agropyron suhsecundum Agropyron tnichycaulum Allium acuuiinatum Amelanchier alni folia f f g g g cf sh 0.4.3 -0.31 0.00 0.09 0.56 -0.30 -0.50 3 13 1 4 9 2 12 3 1 11 3 2 2 2 3 32 10 2 6 2 22 8 3 23 9 1 9 6 2 4 6 2 6 15 1 6 1 Antennario luzuloides f 1.00 1 Antennaria rosea f -1.00 1 Apocynum androsaemifolium Arctostapliylos uva-ursi f esh -0.78 1.00 2 4 9 Artemisia tridentata esh -1.00 1 Aster chilensis var. adscendens f -0.28 6 3 10 6 Bromus ciliatus g 0.32 11 11 4 9 2 5 6 5 Bromus tectorum g -1.00 1 Carex geyeri Carex rossii Ceanothus velutinus eg eg esh 0.36 0.31 0.09 6 23 9 35 1 6 43 18 1 14 15 22 5 6 1 Chenopodium amhrosioides Chenopodium fremontii Chrysopsis tillosa Cirsium undulatum a a f f 1.00 -0.,32 -0.79 -0..33 9 8 1 1 4 3 5 20 33 17 3 42 2 6 10 CoUinsia pnrviflora Collomia linearis Comandra umbellata a a cf -0.13 -0.19 0.07 7 20 11 .33 35 2 39 36 4 19 28 5 IS 8 41 66 1 36 39 8 .33 37 2 Cryptantha circttmscissa Eriogonum heracleoides a esh -0.42 0.07 6 1 1 14 16 1 9 1 9 Erysimum asperum Euphorbia rohusta Galium horeale f f f -1.00 1.00 -1.00 1 1 1 Gayophytum ramosissimum Uydrophyllum capitatum Ipompsis agoregata Juniperus communis Lomatium grayi Mahonia repens a cf f esh f esh -0.91 -0.02 -1.00 1.00 -0.05 0.08 1 4 7 70 2 1 9 40 1 5 4 2 60 1 10 48 11 2 15 47 5 5 1 2 43 30 4 3 44 59 10 50 June 1980 EvENSON ET AL.: PlANT EcOLOGY 171 associated with one of the four canopv spe- cies. This difference was then normahzed by dividing by the total occurrence of that spe- cies in all stands. The four components thus sum to the species preference index discussed above (within round-off error). The com- ponents of the species preference index clari- fy how a species preference for understories or open areas is associated with a particular canopy species. Some luiderstorv species are highly associated with a particular canopy tree species, and others are not. For example, Carex geycri's preference for the understorv is strongly associated with Gambel oak. Yet, Stellaria jamesiana prefers the understory much more evenly for three of the four cano- py species. Correlation analysis was performed to study the relationship of the environmental and vegetational parameters which were measured. Table 4 shows the significant posi- tive and negative correlations. Light intensity correlates significantly with all variables except percent annuals. The negative correlation of light intensity with pH, Itter depth, soil depth, and percent litter cover is to be expected because of the gener- ally higher values of these parameters under the canopies. Similarly, there is more exposed rock and living cover in the open areas. The correlations of light intensity with life forms reflect the preference of shrul>s and grasses for understory areas and forbs and annuals for open areas (Wilcox, Brotherson, and Evenson 1981). The significant correlations of pH with Table 2 continued. Understory Open area Life Preference stands stands Species form index Ceve Pipo Jusc Quga Ceve Pipo Jusc Quga Melica bulbosa g 1.00 2 Merten.mi hrcvistijle f -l.(X) 2 Osniorhizci obtttsa f -l.(K) 1 Pdcliystima nujisiiiites esh 0.19 3 2 10 4 2 3 6 2 Penstcmon sp. f -1.00 2 Piiuis coutorta at l.(X) 1 Pintis pondcnmi at 0.00 I 1 Poa curt a g -0.36 - 3 12 Poa fcndlcriana g -1.00 3 Poa pratensis g 0.11 5 4 Pohjgonuin cloui:,hi.sii a -0.51 65 17 29 28 129 92 87 119 Potcntilla olanduhmi f -0. 1 1 3 1 1 4 Pntniis viroiniana sh 0.54 1 1 6 2 2 1 Purshid tridcntatci sh l.(K) 1 1 {hicrru.s ^ambelii t l.(K) 1 Rosa ivoodsii sh 0.20 5 5 6 S 6 4 2 4 Seduiii stenopctaluni f -0.36 6 4 15 3 20 11 21 ( Scnccio u in tah ensis f -0.64 2 4 1 4 Silcnc menziesii ef 0.33 2 1 Solidaoo tnissouricnsis f -0.69 5 2 1 25 9 9 Solida^o multiradiata f 0.05 9 17 4 2 4 17 6 2 SoUdaop sparsiflora f -0.53 11 13 23 Stellaria janwsiana cf 0.51 55 94 27 105 11 36 16 27 y,tipa cohiiubiaiia g -l.(K) 5 Stipa k'ttcrmanii g 0.09 5 1 5 1 5 3 2 Sijmphoricarpos oreophilus sh 0.20 8 2 12 5 4 3 9 2 Taraxacum officinale f 0,(K) 4 6 2 2 2 4 3 5 Tlialictrum fendleri f l.(K) 6 Thlaspi montanutn f -0.20 .53 16 28 40 48 55 25 76 Tragopooon dubius a 0.25 5 1 2 Viguicra midtijlora f 0.00 9 2 2 5 6 2 Viola nuttallii cf 0.(X) 1 1 Viola purpurea f 0.52 11 5 1 2 2 Total number of species 39 34 36 37 38 32 41 .39 172 Great Basin Naturalist Vol. 40, No. 2 other variables again reflect the tendency to- nated communities with relatively low total ward higher pH under the canopies. Sim- living cover and significant amounts of litter ilarly, for litter depth, soil depth, percent ex- cover. posed rock, percent litter cover, and percent By contrast, shrubs and forbs are positively living cover the correlations generally reflect correlated with percent living cover, but the relationship of these parameters to the negatively correlated with each other. Shrubs canopy-covered or open-area condition. provided a significant proportion of living Correlations of these parameters with per- cover in any quadrat in which they occur just cent annuals are not so easily interpreted, because of their size. This fact accounts for however. Annuals tend shghtly to prefer the their positive correlation with living cover, open areas, hence the negative correlation However, shrubs tend to prefer understory with pH. Their positive correlation with per- habitats and forbs prefer the open areas. So cent litter cover, however, is better under- forbs are positively correlated with living stood by observing the negative correlation cover due to the greater cover outside the with all other life forms and percent living canopies, while being negatively correlated cover. Annuals tend to grow in annual-domi- with shrubs. Table 3. Preference index components by tree species. Life form codes are as in Table 2. Life form Preference _ index Preference index components Species Ceve Pipo Jusc Quga Antennaria luzuloides f LOO 0.00 0.00 0.00 1.00 Arctostaphylos uva-ursi ash LOO 0.00 0.00 1.00 0.00 Chenopodium ambrosioides a LOO 0.00 0.00 0.00 LOO Euphorbia robusta f LOO 1.00 0.00 0.00 0.00 Jttniperus conimiinis esh LOO 0.00 0.00 1.00 0.00 Melica bulbosa g LOO 0.00 1.00 0.00 0.00 Pinus contorta at LOO 0.00 0.00 0.00 1.00 Purshia tridentata sh 1.00 0.50 0.00 0.00 0.50 Querciis gambelii t 1.00 0.00 0.00 1.00 0.00 Tlialictrwn fendleri f 1.00 0.00 0.00 1.00 0.00 Agropyron trachycaitlum g 0.56 0.00 0.44 0.07 0.04 Pruntis virginiana sh 0.54 0.08 -0.08 0.38 0.15 Viola purpurea f 0.52 0.48 -0.10 0.00 0.14 Stellaria jamesiana cf 0.51 0.12 0.16 0.03 0.21 Adiillea millefolium f 0.43 0.01 0.01 0.03 0.37 Carex geyeri eg 0.36 0.04 -0.04 0.09 0.27 Silene menziesii af 0.33 -0.33 0.67 0.00 0.00 Bromus ciliatus g 0.32 0.17 0.11 -0.04 0.08 Carex rossii eg 0.31 0.14 0.00 -0.09 0.27 Tragopogon dubius a 0.25 0.50 0.00 -0.25 0.00 Rosa woodsii sh 0.20 -0.02 0.02 0.10 0.10 Symphoricarpos oreophilus sh 0.20 0.09 -0.02 0.07 0.07 Pachystima myrsinites esh 0.19 0.03 -0.03 0.12 0.06 Poa pratensis g 0.11 0.00 0.56 -0.44 0.00 Agropyron subsecundum g 0.09 0.00 0.18 -0.18 0.09 Ceanothus celutinm esh 0.09 0.00 0.00 0.09 0.00 Stipa lettennanii g 0.09 0.00 -0.09 0.14 0.05 Mahonia repens ash 0.08 0.06 -0.01 0.04 -0.00 Comandra umhellata af 0.07 0.07 0.02 -0.10 0.07 Eriogonum heracleoides ash 0.07 0.00 0.34 -0.05 -0.22 Solidago multiradiata f 0.05 0.08 0.00 -0.03 0.00 Agropyron spicatum g 0.00 0.00 0.00 0.00 0.00 Pinus ponderosa at 0.00 0.00 0.00 0.00 0.00 Taraxacum officinale f 0.00 0.07 0.07 -0.04 -0.11 Viguiera multiflora f 0.00 0.15 -0.23 0.08 0.00 June 1980 EVENSON ET AL.: PlANT EcOLOGY 173 Summary and Conclusions Canopy tree species clearly influence both vegetation and environment in their under- stories (Wilcox, Brotherson, and Evenson 1981). The effects of this influence on plant distributions are shown clearly for individual species in Tables 2 and 3. The correlations of the environmental and vegetational parameters among themselves can be imderstood on the basis of a few im- portant concepts. (1) The environmental parameters (light in- tensity, pH, litter depth, soil depth, percent- age of exposed rock, and percentage of litter cover) are directly influenced by the pres- ence or absence of canopy cover. All correla- tions among these parameters are as expected on that basis. (2) The vegetational correlations follow primarily from the facts that there is more living cover in open areas than in the under- stories; shnibs and grasses tend to prefer the understories; and forbs and annuals tend to prefer open areas. (3) Shrubs follow the pattern inferred from their tendency to prefer understory areas ex- cept for their positive correlation with the percentage of living cover. This is because shrubs themselves provide a large fraction of the living cover that is found in understory quadrats. (4) Annuals are different. They apparently tend to grow in annual-dominated patches with low total living cover and relatively high litter cover. Table 3 continued. Life form Preference index Preference index components Species Ceve Pipo Jusc Quga Viola ntittallii cf 0.00 0.00 0.00 0.00 0.00 Hydrophylltiiii capitatum cf -0.02 0.05 -0.10 0.02 0.(X) Lomatiwn gnnji f -0.05 -0.21 0.18 -0.03 0.(X) Potentilla glandulosa f -0.11 0..33 0.00 -0.11 -0.33 CoUinsia paniflora a -0.13 -0.05 -0.04 0.01 -0.06 Colloniki linearis a -0.19 -0.04 -0.11 -0.01 -0.03 TliUispi montanwn f -0.20 0.01 -0.11 0.01 -0.11 Aster chilensis tar. adscendens f -0.28 -0.16 0.00 0.00 -0.12 Allium acuminatutn cf -0.30 -0.19 -0.16 -0.08 0.14 Agoseris oUnica f -0.31 -0.09 -0.14 -0.03 -0.05 Chenopodiutn fremontii a -0.32 -0.25 0.02 -0.07 -0.02 CirsitiDi iindiilatuui f -0.33 0..33 ().(X) -0.67 0.00 Poo eurta g -0.36 0.00 0.00 -0.14 -0.23 Sedtim stenopetahnti f -0.36 -0.16 -0.08 -0.07 -0.05 Cnjptantha circuinscissa a -0.42 -0.42 0.04 0.00 -0.04 Amelanchier alnifolia sh -0.50 0.00 0.(X) -0.50 0.00 Polygonum dotiglasii a -0.51 -0.11 -0.13 -0.10 -0.16 Solidago sparsi flora f -0.53 0.00 -0.28 0.00 -0.26 Senecio uintahensis f -0.64 -0.18 0.00 -0.09 -()..36 Solidago m isso u riensis f -0.69 -0.39 -0.14 -0.16 0.(X) Apocynum androsaemifolium f -0.78 -0.39 -0..39 0.00 0.00 Chrysopsis villosa f -0.79 -0.22 -0.15 -0..33 -0.09 Cayophytum ramosissimum a -0.91 -0.09 -0.03 -0.26 -0..53 Antennaria rosea f -1.00 0.00 ().(X) -l.(X) 0.(X) Artemisia tridentiita esh -l.(X) 0.00 0.(X) -l.(X) O.tX) Bromus tectorum g -l.(X) 0.00 0.(X) ().(X) -1.00 Erysimum aspcrum f -1.00 -1.00 0.00 0.(X) 0.(X) Galium horeale f -1.00 0.(X) 0.(X) 0.(X) -l.(X) Ipompsis aggregata f -l.(K) 0.00 -1.00 0.(X) ().(X) Me rten s ia b re f istyla f -l.(K) -1.00 0.(X) 0.(X) 0.00 Osmorhiza obtusa f -l.(X) ().(X) ().(X) 0.00 -l.(X) Penstemon sp. f -l.(X) 0.(X) ().(X) -1.00 0.(X) Poa fendleriana g -1.00 -1.00 0.00 0.00 ().(X) Stipa Columbiana g -1.00 0.00 0.00 0.00 -l.(X) 174 Great Basin Naturalist Vol. 40, No. 2 Table 4. Significance levels for correlations of life form types, cover, and measured environmental parameters for all stands studied. Variables 7 8 10 11 1. Light intensity 2. pH 3. Litter depth 4. Soil depth 5. % exposed rock 6. % litter cover 7. % living cover 8. % shrubs 9. %forbs 10. % grasses 11. % annuals .001 -.001 +.001 -.001 +.1 -.05 +.001 -.001 NS NS NS -.05 NS NS + .05 -.01 NS -.05 + .(K)1 -.001 +.001 NS +.1 -.001 NS NS -.001 +.fK)l -.05 NS -.01 +.1 NS -.(K)l +.1 -.1 +.001 -.01 NS -.001 NS -.001 +.001 +.05 + .01 +.1 NS -.001 -.001 NS -.001 NS -.05 NS -.05 -.01 Literature Cited Anderson, R. C, O. L. Loucks, and A. M. Swain. 1969. Herbaceous response to canopy cover, light in- tensity and throughfall precipitation in co- niferous forests. Ecology .50:25.5-263. AucLAiR, .-v. N., and R. G. Goff. 1971. Diversity rela- tions of upland forests in the western Great Lakes area. American Naturalist 105:499-528. Blackman, G. E. 1956. Influence of light and temper- ature on leaf growth. Pages 151-169 in F. L. Mil- thorpe, ed. Growth of leaves. Butterworths Scien- tific Publications, London. 223 pp. Cline, M. G. 1966. Effect of temperature and light in- tensity on Scropltularia marihindica. Ecologv 47:782-795. CoTTA.M, W. P. 1930. Some unusual floristic features of the Uinta Mountains, Utah. Proceedings of the Utah Academy of Sciences 7:48-49. Daubenmire, R. 1959. A canopy-coverage method of vegetational analysis. Northwest Science .33:4.3-66. del Moral, R. 1972. Diversity patterns in forest vegeta- tion of the VVenatchee Mountains, Washington. Torrey Botany Club Bulletin 99:57-64. Federer, C. a., and C. B. Tanner. 1966. Spectral distri- bution of light in the forest. Ecologv 47:55.5-.561. Gordon, D. T. 1962. Growth response of east side pine poles to removal of low vegetation. U.S. Depart- ment of .'\gricvdture. Forest Service, Pacific Southwest Forest and Range Experiment Station Research Note 209. 3 pp. McCoNNELL, B. R., A.ND J. G. Smith. 1970. Response of understory vegetation to ponderosa pine thinning in eastern Washington. Journal of Range Man- agement 23:208-212. McQueen, D. R. 1973. Changes in understory vegeta- tion and fine root quality following thinning of 30-year-old Finns radiata in central North Island, New Zealand. Journal of .\pplied Ecology 10:13-21. RusEL, \. R., AND H. J. Evans. 1966. The nitrogen fixing capacity of Ceanothus vehitinns. Forest Science 12:164-169. Smith, B. E., and C. Cottam. 1967. Spatial relationships of mesic forest herbs in southern Wisconsin. Ecology 48:.546-548. Webster, S. C, C. T. Youngberg, .\nd A. G. Wollum II. 1967. Fixation of nitrogen by bitterbrush {Pur- sliia tridentata (Pursh) DC). Nature 216(51 1.3):392-93. Wilcox, R. B., J. D. Brotherson, a.nd W. E. Eve.nson. 1981. Canopy influence on understory commu- nity composition. (In press: Northwest Science). Wollum, A. G. II, and C. T. Youngberg. 1964. The in- fluence of nitrogen fixation by non-leguminous woody plants on the growth of pine seedlings. Journal of Forestry 62:216-321. SEASONAL ACTIVITY PATTERN OF COLUMBIAN GROUND SQUIRRELS IN THE IDAHO PRIMITIVE AREA Charles L. Elliott' and jerran T. Flinders' .\bstr.\ct.— Data were gathered concerning the seasonal activity pattern of a population of Columbian ground scjuirrels (Spermophihis colttmbianus) in the Idaho Primitive Area. Adult females were significantly more active in |une of all vears than were adult males. \ relationship between ground squirrel activity and temperature is postulat- ed in which the squirrels alter their activity so as to avoid high temperatures and possible heat stress. Members of the genus Spennophilus are characterized bv a seasonally short period of aboveground activity and a prolonged period of hibernation. During this time of surface activity ground squirrels must establish terri- tories, breed, reproduce, and gain .sufficient weight to survive the inactive season. The an- nual cycle of activity for various species of ground squirrels has been reported (Skryja and Clark 1970, Michener 1974, Loehr and Ris.ser 1977), but these data as they apply to populations of Columbian ground squirrels {Spennophilus cohimbianiis) are limited. Methods The study was conducted at Cold Mead- ows, an 87 ha meadow (elev. 2010 m) located in the northeastern portion of the Big Creek Ranger District, Idaho Primitive Area. A de- scription of the Big Creek area has appeared elsewhere (Wing 1969, Hornocker 1970, Sei- densticker et al. 1973). Ground squirrels were trapped from 12-19 June, 17-24 July, and 14-21 August, 1976-1978. Field work prior to 12 June was impractical because of bad weather and the inaccessibility of the study area. A 90 x 90 m grid with 36 trapping sta- tions 15 m apart was establi.shed on the cen- tral portion of the meadow. One live trap (15 X 15 X 48 cm) was placed at each trapping station. Traps were baited with carrot and checked every hour. Captured squirrels were marked using the toe clipping sequence of Melchior and Iwen (1965), sexed, measured. weighed, time-of-capture recorded, and re- leased back onto the grid. Vegetation was collected using the procedure outlined by Tadmor et al. (1975). All plant samples were weighed to the nearest gram in the field and then brought back to the laboratory, where they were oven dried at 64 C for three days. The dried specimens were then weighed to the nearest gram and percent moisture con- tent calculated. Daily ambient temperatures were obtained using a Taylor Maximum-Min- imum thermometer. Ground squirrel activity in this study was equated with the animals presence in the traps. Bias due to 'trap-shy' or 'trap-happy' squirrels may have occurred, but attempts to conduct hourly visual censuses proved unre- liable during the latter months due to the in- crease in vegetation height. Results and Discussion The number of Columbian ground squirrels captured, including recaptures, is depicted in Table 1. Adult female squirrels were signifi- cantly more active in June of each year than were adult males (Kolmogorov-Smirnov Two Sample Test, P <0.05). Activity for July and August was not significantly different be- tween sexes of adult or young squirrels. Males are territorial during the breeding season (Steiner 1970a), exliibiting extreme ag- gression toward other males and occasionallv raiding the nesting ground and colonies of adjacent males (Steiner 1970a, 1970b, 1971, 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 175 176 Great Basin Naturalist Vol. 40, No. 2 1972). This aggressive behavior and Lam- beth's (1977) findings that Columbian ground squirrels utilize a core area or arena of activi- ty between 21 and 40 m in size may account for the greater number of females captured in June. Dominant males would have been excluding other male squirrels from the trapping grid. The hibernation entry sequence described by Manville (1959) and Michener (1974) for S. columbianus was not apparent for the Cold Meadows colony. This disparity may be artificial because adult groimd squirrels dur- ing August become very lethargic and in- active and were extremely difficult to trap. Shaw (1925) has suggested that early spring activities of Columbian ground squir- rels are largely controlled by temperature alone, whereas estivation is induced by the drying of the vegetation. Howell (1938) felt that "the date of beginning estivation was de- termined chiefly by the ripening of the vege- tation and consequent reduction of the mois- ture content in their (ground squirrels) food, and in part also by the accumulation of fat in the body." Nansel and Knoche (1972) also ob- served Columbian ground squirrels and pos- tulated that hibernation was a response to drought prior to a decrease in temperature. Cold Meadows plant moisture content de- clined as the season progressed (Table 2), as did the squirrels' aboveground activity (Table Peak daily activity of ground squirrels at Cold Meadows appears to be determined by temperature. Table 3 shows the monthly time interval exhibiting the greatest percentage of adult ground squirrel activity. The mean maximum temperature for June 1976 (Table 4) was significantly greater than June 1977 and June 1978. The maximum percent of ac- tivity for June 1976 occurs later in the day than for June 1977 or June 1978. This same type of activity shift in relation to significant- ly greater maximum temperatures is evident for July; July 1978 is significantly greater Table 1. Live trapping results by age and sex for Columbian ground squirrels at Cold Meadows, Idaho Primitive Area, 1976-1978. June July August Adult Adult Juvenile Adult Juvenile Male : Female Male Femal e Male Female Male Female Male Female 1976 12 25 15 32 15 13 5 5 13 5 1977 15 39 7 12 11 8 3 2 2 4 1978 24 32 18 13 6 7 2 6 / 6 Table 2. Mean percent moisture content ( ± standard deviation) of plant species exhibiting the highest frequency of occurrence at Cold Meadows, Idaho Primitive Area, 1976-1978. 1976 1977 1978 Species' July August June July August June July August Achillea millefolium 66 ±1 72 ±4 72 ±1 72 ±5 58 ±1 82 ±3 77 ±2 60 ±4 Carex aquatilis 65 ±5 63 ±6 61 ± 7 51 ± 4 30 ± 1 67 ±2 58 ±3 45 ±5 Fragaria virginiana 67 ± 8 66 ± 2 65 ± 8 65 ± 7 52 ± 1 76 ± 1 67 ± 2 55 ± 1 Penstemon procerus 68 ±6 65 ±2 72 ±3 66 ±5 47 ±1 73 ± 1 69 ±2 63 ±2 Phleum alpimim 63 ±5 57 ±7 57 ±1 49 ±8 27 ±1 72 ±2 59 ±4 30 ±1 l:n = 10 samples/species/month. Table 3. Peak activity time intervals for adult Columbian ground squirrels at Cold Meadows, Idaho Primitive Area, 1976-1978. 1976 1977 1978 June 12-19 July 17-24 August 14-21 1700-1800 hours 13(K)-14(X) 1600-17(X) 1400-1500 1200-1300 1200-1300 1400-1500 1000-1100 13(K)-1400 June 1980 Elliott, Flinders: Columbian Ground Squirrels 177 Table 4. Monthly mean maximum temperatures (± standard deviation) at Cold Meadows, Idaho Primi- tive Area, 1976-1978. 1976 1977 1978 June 12-19 22±2C' 20±2l 20*2^ July 17-24 22±l2 22±li-2 24±l''2 August 14-21 17 ±2''^ 25±1^ 17±2'-2 1; Means tested for significance using unpaired t-test at the 0.05 level. 2: Means tested for significance using Cochran and Cox test at the 0.05 level (Ferguson 1959:143). than July 1976 and July 1977. August 1977 mean daily maximum temperature is signifi- cantly greater than August 1976 and August 1978; yet the expected shift in peak activity does not occur. This may be due to the small activity sample size for August 1977. The relationship of temperature to activity indicates that the higher the average daily maximum temperature, the earlier or later in the day peak activity will occur. If this hy- pothesis is valid, then, for those sampling pe- riods where the average maximum temper- atures were not significantly different, the time interval of peak activity should be sim- ilar. This relationship is observed for June 1977 and June 1978, and July 1976 and July 1977 (Table 3). Betts (1976) observed a lower elevation (1360 m) colony of S. columhianus in western Montana and reported that, with the increase in temperatures during lactation and post- lactation periods, there was an increase in morning and late afternoon activity and a de- crease in midday activity. Betts postulated that temperature or solar radiation may limit the amount of consecutive time Columbian ground squirrels can spent aboveground. The observations of Betts (1976) and data reported here indicate that the scheduling of surface activity for these squirrels is an ap- parent behavioral response designed to es- cape heat stress. We thank the University of Idaho for per- mission to use the facilities at the Taylor Ranch Field Station, Idaho Primitive Area. Literature Cited Betts, B. J. 1976. Behaviour in a population of Colum- bian ground scjuirrels, Spernu)philus coliuiibianus columhianus. .^nini. Behav. 24:652-680. Ferguson, C. \. 1959. Statistical analysis in psychology and education. McGraw-Hill Book Co., Inc., New York. .347 pp. HoRNocKER, M. G. 1970. An analysis of mountain lion predation upon mule deer and elk in the Idaho Primitive Area. Wildlife Monogr. 21:l-.39. Howell, \. H. 1938. Revision of the North American ground squirrels. North American Fauna 56:85-90. Lambeth, R. E. 1977. The Coliunhian groimd squirrel in subalpine forest openings in the Idaho Batholith. Unpublished thesis, Univ. Idaho, Moscow. 1 1.3 pp. Loehr, K. a., and a. C. Risser, Jr. 1977. Daily and sea- sonal activity patterns of the Belding ground squirrel in the Sierra Nevada. J. Mammal. 58:445-448. Manville, R. H. 1959. The Columbian ground squirrel in northwestern Montana. J. Mammal. 40:26-45. Melchior, H. R., and F. A. Iwen. 1965. Trapping, re- straining and marking .\rctic ground squirrels for behavioral observations. J. Wildlife Manage. 29:671-679. Michener, D. R. 1974. .\nnual cycle of activity and weight changes in Richardson's ground squirrel, Spermopliilus richardsonii. Can. Field-Nat. 88:409-413. Nansel, D., and L. Knoche. 1972. Blood changes in tor- pid and non-torpid Columbian ground squirrels, Spermophilus cohimbianus. Comp. Biochem. Physiol. 41A: 175-179. Seidensticker, J. C IV, M. G. Hor.nocker, W. V. Wiles, a.nd J. P. Messick. 1973. .Moimtain lion .social organization in the Idaho Primitive Area. Wildlife Monogr. .35:1-60. Shaw, W. T. 1925. Duration of the aestivation and hi- bernation of the Columbian ground squirrel. Ecology 6:75-81. Skryja, D. D., and T. W. Clark. 1970. Reproduction, seasonal changes in body weight, fat deposition, spleen and adrenal gland weight of the golden- mantled ground squirrel Spennophihis lateralis lateralis, (Sciuridae) in the Laramie Mountains, Wyoming. Southwestern Nat. 15:201-208. Stelner, a. L. 1970a. Etude descriptive de quelques ac- tivities et comportements de base de Spermo- philus columhianus columhianus (Ord). I. Loco- motion, soins du corps curiosite et alarme, reproduction. Rev. Comp. Animal 4:3-21. 1970b. Etude descriptive de quelques activities et comportements de base de Spennophilus colum- hianus coltimhianus (Ord). II. Vie de groups. Rev. Comp. .Animal 4:23-42. 1971. Plav activitv of Columbian ground squir- rels. Z. Tierpsychoi. 28:247-261. 1972. Mortality resulting from intraspecific fight- ing in some ground squirrel populations. J. .Mam- mal. 53:601-603. Tadmor, N. H., a. Briegher, I. Noy-Meir, R. W. Benjamin, and E. Eyal. 1975. .^n evaluation of the calibrated weight-estimate method for mea- suring production in annual vegetation. J. Range Manage. 28:65-69. Wlng, L. D. 1969. Ecology and herbivore use of five mountain meadows in the Idaho Primitive ,\rea. Unpublished thesis, Univ. Idaho, Moscow. 215 pp. HABITAT AND PLANT DISTRIBUTIONS IN HANGING GARDENS OF THE NARROWS, ZION NATIONAL PARK, UTAH George P. Malanson' ,\bstract.— Hanging gardens are insular plant comnuinities of the Colorado Plateau. This study examines hanging gardens in the Narrows, Zion National Park, Utah. The floristic similarity of gardens and the presence of species in classes of variables which characterize the habitat are disclosed. Although the gardens are isolated and dissimilar, the individual species are not restricted in the range of habitat found. Floristic dissimilarity cannot be attributed to differences in habitat. These results are compared to studies of hanging gardens in eastern Utah. Hanging gardens are plant communities growing at seeps on the canyon walls of the Colorado Plateau. The hanging garden envi- ronment is characterized by shallow soils at a seep from bedrock. Seeps occur where water has percolated through a porous formation until meeting a less permeable layer of rock. Then the water flows laterally until a canyon intersects this plane. The narrow canyons of- ten shade the hanging gardens. Compared to other environments of the Colorado Plateau, the hanging gardens are cool and moist. The Narrows of the North Fork of the Vir- gin River in Zion National Park is an arch- etypal hanging garden locale. In an 8 km sec- tion there are about 60 gardens, varying in size from a few square centimeters to over 100 m-. Most of these are at permanent seeps with small discharges of water. The hanging gardens of the Narrows assume a variety of shapes, but in general they occupy a place where erosion has modified the steepness of the canyon wall. Often these places are hori- zontal bands. Other gardens occur where ver- tical jointing has concentrated the seepage. Some gardens occupy remnants of potholes, and others are on bulges of travertine. A few are in alcoves. Most hanging gardens in the Narrows are close to the level of the river, where they may be vulnerable to flash floods. This study examines the relationship be- tween the plant species and the habitat in the hanging gardens of the Narrows. The con- cepts of Ramensky (1924) and Gleason (1926) are the basis for hypothesizing that the pres- ence and importance of species at sites are determined by their individual tolerances and requirements in relation to the habitat. This idea can be evaluated by examining the flo- ristic similarity between sites and the in- cidence of species across a range of variables that characterize the biotopes. The hypoth- esis leads one to expect a positive relation- ship between floristic similarity and sim- ilarity of biotope (i.e., if sites represent a single habitat). A few authors have investigated the vege- tation of hanging gardens. In a general eco- logical stvidy of Zion National Park, Wood- bury (1933) outlined the stages of primary succession that occur at seeps. Welsh and Toft (1976) described a variety of garden types in Glen Canyon, Utah, based on the form created by the erosion of the rock, and traced the geographical affinities of the spe- cies they found. They called hanging gardens "relictual refugia" because the gardens pro- vided sites for species from other southwest locations, boreal forests, and earlier epochs. Welsh and Wood (1976) concluded that hanging gardens have a stable structure, attri- buting change in species importance mea- -sured over a one-year interval to measure- ment error. Wood and Welsh (1976) found productivity to be relatively high for this 'Department of Geography, University of Utah, Salt Lake City. Utah 84112. Pre.scnt address: Department of Geography, University of California, Los Angeles, CaWfomia 90024. 178 June 1980 Malanson: Hanging Gardens 179 type of vegetation, and presumed it to be steady. Nebeker et al. (1977) believed that the flo- ristic dissimihiritv found in hanging gardens of eastern Utah indicated ''random assort- ments of individuals capable of exploiting the environments of individual sites." They con- clude that over 25 percent of the species were ecological specialists of hanging garden habitats, within the Colorado Plateau region. A few studies briefly mention hanging gar- dens. Clover and Jotter (1944), Flowers (1959), and Loope (1976) describe hanging gardens in general ecological studies of the Colorado River area. Raines (1976) noted that hanging gardens can be critical habitats for small mammals in drought years. All these studies emphasize the importance of habitat and support the hypothesis of this study. Methods I sampled 29 hanging gardens in an 8 km section of the Narrows and in 0.75 km of a tributary (Orderville Canyon) between June and September 1977. Sites were chosen infor- mally on the basis of access, but a representa- tive range of garden sizes was sought. To es- timate percent foliar cover of species, I recorded the number of decimeters inter- cepted by a species along line transects spaced at 2 m intervals and perpendicular to the long axis of each garden. In gardens nimi- bered 1, 2, and 3, I placed a 2500 cm- quad- rat every 2 m on transects spaced at 2 m in- tervals and recorded the number of 100 cm- grids occupied bv each species. This method proved to be impractical, although in com- parison tests the results were not significantly different. From the values of relative cover for each garden, I calculated the similarity of each pair of hanging gardens according to Ru- zicka's (1958) index: SI = (2A/2B)(100) where A is the smaller and B the larger value of a species in two sites, considering all spe- cies present in either site. I measured 11 variables to characterize the biotope of each hanging garden. Soil and wa- ter pH at each site was measured color- imetrically (Microessential Laboratory 0.2 unit paper). I gathered grab samples of soil from the surface where there was any depth, not taking any litter. I took 125 ml samples of water from the seeps. In a few cases sam- ples could not be taken because of a paucity of soil or water. The USDA Soils Testing Laboratory at Logan, Utah, measured soil and water conductivity as a surrogate of sali- nity, and the total phosphorous content of the soil. I measured the slope of each garden by taking the height:depth ratio and calcu- lating a percent slope. I sampled soil depth every 4 m across the center of each garden with a wire rod to obtain a single average soil depth measure; in smaller gardens, one or two evenly spaced measures were taken. Direct solar radiation was derived by a computer program of Williams et al. (1972), which accounts for slope, azimuth, and lati- tude of the site, and intervening topography that shades the site. The program computes the calories per square centimeter for any one day, and I summed the 120th, 180th, and 240th days of the year to bracket the grow- ing season. The scale of resolution does not account for all possible variations in micro- relief. In locations with particular sunblock- ing features, such as alcoves, I reduced the computed value 10 percent. At one garden where the three-day sum was zero, although I observed direct radiation at the site, I arbi- trarily assigned a value of 50 cal/cm-. I measured three spatial variables: area, isolation, and distance to the Gateway to the Narrows Trail. I derived the area of each gar- den from the grid formed by the species in- tercept transects. Isolation is defined here as the sum of the distances from each garden to its nearest three neighbors. These distances were measured on a topographic map (ZNHA 1977). The Gateway to the Narrows Trail is the scene of much pedestrian tourist traffic, but above the terminus human use de- clines rapidly. The distance to the trail, also measured on the topographic map (ZNHA 1977), may affect invasion by nonnative spe- cies. I analyzed the relationships between spe- cies presence and absence and the 11 envi- ronmental variables. These variables were considered separately because there was low correlation between any two. I followed 180 Great Basin Naturalist Vol. 40, No. 2 Strahler's (1978) use of the Gh statistic to dis- close significant differences between species presence and absence in ordinal categories of the environmental variables. Although the data are ratio scaled and the categories are ordinal, the presence of many zeros in the species matrix and the number of tie scores among the environmental variables make more powerful tests less dependable. Only the 13 most common species occur frequent- ly enough to be tested. I divided each of the 11 environmental variables into four or five ordinal categories (Table 1). The number of categories and the ranges are arbitrary, ex- cept that each category represents at least one garden. I attempted to categorize the variables so that the results of the Gh tests would represent the relationships between species presence and a continuous change in the variable. I also calculated a species-area curve of the gardens by a regression of log number of species on log area. Following the Whitehead and Jones (1969) treatment of island floras, I deleted the smallest garden on successive cal- culations to find the point below which the area detracted from the fit of the curve. Results Forty-eight species were coimted in the 29 hanging gardens. Nine taxa were identified only to genus or family, and four rare plants could not be identified (Table 2). Richness ranged from 2 to 20. The frequency of occur- rence ranged from 1 to 17. Although no spe- cies occurred in all gardens, the 13 species occupying five or more gardens are used here as diagnostic species, because species of mod- erate constancy are good indicators of varia- tion in the environment (Mueller-Dombois and Ellenberg 1974). The many rare species (35 occupying less than five gardens) are sim- ply not common enough to give useable cor- relations with garden environments. For complete tables of data on species cover, flo- ristic similarity, and the environmental varia- bles, see Malanson (1978). The floristic similarity of the hanging gar- dens is low, ranging from to 77, and aver- aging 10.23 for all gardens. This value of sim- ilarity is close to the results of Nebeker et al. (1977). All hanging gardens provide a mesic to hygric biotope. For water and soil, values of pH ranged from 5.9 to 7.2 and 6.3 to 7.4, and values of salinity from 0.3 to 2.5 jumhos/cm and 230 to 1420 mhos/cm, respectively. In this study, average soil phosphorous content ranges from 2.1 to 84 ppm. Direct solar radi- ation varies from to 683 cal/cm* for the three-day sum. Average soil depth ranges from 0.1 to 48.4 cm, but exceeds 10 cm in only three gardens. Slopes average between 40 percent and vertical. Area ranges from 2 to 100 m- among samples. Most values of iso- lation are low, 24 gardens are less than 300 m from the nearest three neighbors. All but four distances from the Gateway to the Narrows Table 1. Classes of the environmental variables. Classes Variables 1 2 3 4 5 Area (m-) 0-10 10-25 25-50 50 Distance from trail (m) 50O-ia30 1631-2760 2761-3890 3891-5020 5021-6150 Isolation (m) 0-80 81-160 161-240 241-320 ,320 Phosphorous (ppm) 0-5.0 5.1-15 16-36 37-87 Slope (percent) 1-50 51-125 126-275 276-525 526 Soil depth (cm) 0-1 1-2 2-4 4-8 8 Soil pH 6.3-6.4 6..5-6.6 6.7-6.8 6.9-7.4 Soil salinity (mhos/cm) 0.1-0.4 0.5-0.8 0.9-1.2 1.3-1.8 1.9-2.5 Solar radiation (cal/cm-) 0-150 151-300 .301-4,50 451-600 601-750 Water pH 5.9-6.2 6.3-6.5 6.6-6.8 6.9-7.2 Water salinity (mhos/cm) 0-300 301-600 601-900 900 June 1980 Malanson: Hanging Gardens 181 Trail are clustered bimodally between 500 and 2000 m and between 3000 and 5000 ni. The results of the species presence tests in- dicate that there are few relationships be- tween gradients of the environmental varia- bles and the presence and absence of important species in the sampled gardens (Table 3). Of the 143 tests, only 15 show a significant difference at p = 0.05, and one Table 2. Hanging garden plant species. Species Frequency Abies concolor Acer negiindo A did utti in cap i7/«.s- veneris Adiantuni pcdatum Amaranth tts c^raecizans Amiphalis margaritaceae Apocyniim cannahinum Aquiligia spp. Aralia racemosa A rteniisia ludovicia na Aster eatonii Berheris rcpens Brickelia grandiflora Bromus ciliattts Calaniagrostis scoptilorum Cirsiinn arizonicuin Cystoptcris fragiUs Dodecatheon pulchellum Dryopteris filix-nias Eleocharis sp. Epipactis gigantea Equisetttm hyemale Fraxintis vehitina Galium aparine Hepaticae Heuchera versicolor Junctts sp. Lobelia cardinalis Mimiilus cardinalis Mimtdtis guttatus Muhlenbergia andina Middenbergia mexicana Nasturtium officinale Pea nevadensis Rhus radicans Rubus leucodennis Riimex sp. Salix sp. Smilacina stellata Sphagnum sp. Sphagnaceae Taraxacum officinalis Thalictrum fendleri Viola spp. unidentified # 1 unidentified #2 unidentified #3 unidentified #4 1 3 15 6 1 4 1 13 12 1 8 3 1 3 5 1 17 9 2 3 3 1 3 4 10 2 2 3 13 1 1 2 2 2 2 3 1 1 7 14 9 4 2 3 1 1 2 1 would expect 7 of the tests to prove signifi- cant by chance alone. Only solar radiation consistently returns significant results. The hanging gardens are smaller isolates than those described by Whitehead and Jones (1969) (1.25 to 8.65 ha); yet the changing spe- cies-area curves are similar. The exclusion of the four smallest gardens (6 m-) improves the regression coefficient from 0.55 to 0.67, and changes the slope from 0.28 to 0.48. Discussion The biotopes of the hanging gardens were delimited in this study by the environmental variables of soil and water pH and salinity, soil phosphorous, soil depth, slope, direct so- lar radiation, area, isolation, and distance to the Gateway to the Narrows Trail. In gener- al, the biotopes are within the habitat of the species, and a particular species composition is not maintained by the environment. The distribution of species among the hanging gardens in the Narrows is not strongly affect- ed by their tolerances and requirements. High values of solar radiation seem to mete against the mosses and ferns, but these values occur in only 13 percent of the gardens sam- pled. The great dissimilarity between hang- ing gardens cannot be attributed to dissimilar habitats, although they are insular commu- nities. In this regard, the hanging gardens of the Narrows may be very different from those of the Arches and Canyonlands area. There the habitat differences found over a wide geo- graphic area are more likely to be significant in affecting species presence and garden sim- ilarity. In the Narrows, the proximity of many gardens and their probable suscepti- bility to flash floods prevents a strict com- parison with the research in eastern Utah. The Narrows presents a case in which we must look beyond the structure of the habitat to find an explanation of plant distributions. Malanson and Kay (in preparation) consider disturbance frequencies a likely alternative. Acknowledgments I am thankful for the encouragement and advice of Jeanne Kay, Kimball T. Harper, and Paul A. Kay on all aspects of this re- 182 Great Basin Naturalist Vol. 40. No. 2 search. Kezia M. Nielsen, Garry F. Rogers, Lois A. Amow, and Anne M. Travis helped in the gathering and preparation of the data. This research was funded in part by grants from the University of Utah Student Re- search Grants in Geography, the Zion Natu- ral Historv Association, and Sigma Xi, the Scientific Research Society, and an equip- ment loan bv Robert W. Austin. Literature Cited Clover, E. U., and L. Jotter. 1944. Floristic studies in the canvon of the Colorado and tributaries. .\merican Midland Naturalist 32:591-642. Flow'ers, S. 1959. \egetation of Glen Canyon. Pages 21-61 in A. M. \Voodbur\-. ed. Ecological studies of flora and fauna in Glen Canyon. University of Utah .\nthropological Papers 40. GuE.\so.\, H. .\. 1926. The individualistic concept of plant association. Bulletin of the Torrey Bot- anical Club 5.3:7-26. LooPE, W. E. 1977. Relations of vegetation to environ- ment in Canyonlands National Park. Unpub- lished dissertation. Utah State University. Logan. M.\L.\NSON, G. P. 1978. Distribution of plant species in hanging gardens of the Narrows, Zion National Park. Utah. Unpublished thesis. University of Utah, Salt Lake City. McELLER-DoMBOis. D.. AND H. Ellenberg. 1974. Aims and methods of vegetation ecolog) . John Wiley 6c Sons, New York. Nebeker, G. T., K. T. 1L\rper. J. D. Brotherson, .\nd S. L. Welsh. 1977. Characteristics of plants of com- mon occurrence in hanging gardens of the Colo- rado Plateau. Utah. Unpublished manuscript. Brigham Yoimg University. Provo. R\LNES, J. .\. 1976. Modeling studies of small mammal trapping, phenolog)', and plant succession in the Kaiparowits region. Kane County, Utah. Unpub- lished dissertation. Brigham Young University, Provo. R\MENsia-, L. G. 1924. The basic lawfulness in the struc- ture of the vegetation cover. [In Russian.] \'estnik opy tnogo dela Sredne-Chemoz. Obi.. Voronezh. Pages 37-73 excerpted in E. J. Kormondv, ed. Readings in ecologw Prentice-Hall. Englewood Cliffs, N.J. RuzicKA, M. 1958. Anwendung mathematisch— statistis- cher methoden in der geobotanik, (s\nthetische bearbeitung von aufnahmen). Biologia. Bratislava 13:&47-661. Str.\hler, a. H. 1978. Binary discriminant analvsis: a new method for investigating species-environ- ment relationships. Ecolog\- 59:108-116. Welsh, S. L., and C. .\. Toft. 1976. Biotic conununities of hanging gardens in southeastern Utah in J. R. Murdock,^S^ L. Welsh, and B. W. Wood, eds. Navaho-Kaiparowits environmental baseline studies 1971-1974. Center for Health and Envi- ronmental Studies. Brigham Young Universitv. Provo. Welsh, S. L., .a.nd B. W. Wood. 1976. Structure of a se- lected hanging garden in Glen Canvon of the Colorado River drainage in J. R. Murdock. S. L. Welsh, and B. W. Wood. eds. Navaho-Kaiparo- wits environmental baseline studies 1971-1974. Center for Health and Environmental Studies. Brigham Young University, Provo. \\ hitehead. D. R.. and C. E. Jones. 1969. Small islands and the equilibrium theory of insular biogeo- graphy. Evolution 23:171-179. Williams, L. D., R. G. B.arry, and J. T. .\ndrews. 1972. -Application of computed global radiation for areas of high relief. Journal of Applied Mete- orology 11:526-5.33. Wood. B. W., and S. L. Welsh. 1976. Productivit\ of hanging gardens in J. R. Murdock, S. L. Welsh, and B. W. Wood, eds. Navaho-Kaiparowits envi- ronmental baseline studies 1971-1974. Center for Health and Environmental Studies, Brigham Young Universitv. Provo. WooDBiRY. .\. M. 19.33. Biotic relationships of Zion Canyon. Utah, with special reference to succes- sion. Ecological .Monographs 3:147-245. ZNH.\. 1977. Topographic map of Zion National Park and vicinity. Zion Natural Historv .\ssociation. Springdale, Utah. Table 3. Class of environmental variable in which a species was most significanth limited (classes from Table 1). Variables Species Soil Soil Solar Water Distance Isolation Slope Depth Salinitv Radiation Salinity Adiantum capillus-veneris Adiantum pedatum Aster eatonii CaUimagrostis seoptilorum Cijstupteris fragilis Dodeea th eon p tt Ich ellu m Mimtdus cardinalis Smilacina stellata Sphagnimi sp. Sphagnaceae sp. 4-5 1-2 I 4-5 3-5 3-5 5 4-5 5 3 1&5 3-4 1-2 SHORT-TERM EFFECTS OF LOGGING ON RED-BACKED VOLES AND DEER MICE Thomas M. Cainpbcll HI and Tim VV. Clark- Abstract.— Clearcutting and selective logging effects on red-backed voles (Clethrionomtjs gapperi) and deer mice (Peromysciis maniculatus) were studied (September-November, 1975; June-October, 1976) in Bridger-Teton Nation- al Forest, Wvoming. Five selective cuts (total 137 ha) removed 57 percent (range 34-74 percent) of the trees. One clearcut (9.6 ha) eliminate 84 percent of the trees. Soils remained mesic in selective cuts, but became xeric in the clearcut. Snap-trapping indicated that voles were most abundant on the imlogged and selectively cut mesic sites (76 percent of 408 captures), whereas deer mice were more common on the xeric clearcut (80 percent of 60 captures). Species composition remained unchanged on selective cuts following logging (77 percent voles of 256 captures), but changed from predominantlv voles to mostly deer mice (80 percent of 60 captures) in the clearcut. Intraspecific age and sex ratios, litter sizes, and morphological measurements were compared between logged and unlogged areas. The short-term logging effects on the structure and dynamics (i.e., habitat, num- bers, and morphological and reproductive characteristics) of red-backed voles {Cleth- rionomys gapperi) and deer mice {Peromysciis maniculatus) were examined on a 646 ha study area in the Bridger-Teton National Forest, about 40 km north and 8 km east of Jackson, Wyoming. It is on the backslope of an escarpment that nms west from Mt. Leidy to a point southwest of Toppings Lakes. This backslope is a series of benches with L5 to 40 percent slopes (Brady 1974) at elevations of 2300 to 2700 m. Soils include loams at the surface, with siltv clay to clav loam subsoils (Knight 1973). Climate is characterized by long, cold win- ters with deep snow, a short growing season (average 60 days), and a low mean annual temperature of 1 C. Snowfall averages 345 cm annually and can occur any month. Mean annual precipitation is 69 cm, predominantly snow (Department of Commerce 1975). Jackson Hole vegetation has been de- scribed by Read (1952), and Beetle (1961). A climax spruce-fir {Picea engelmannii-Abies lasiocarpa) forest covers the study area ex- cept on recent bums (last 80 years) and at lower elevations, where lodgepole pine {Pinus contorta) is dominant, and in areas of intermixed coniferous forest and meadows where Douglas fir (Pseudotsuga menziesii) also occurs. Limber pine {Pinus flexilis) is sparsely scattered throughout. Understory is dominated by highbrush huckleberry {Vaccinium globuhre), grouse whortleberry (V. scoparium), mountain ash {Sorbus scopu- lina), and sedges {Carex spp.). Methods The density of all trees (dbh > 15 cm), saplings (dbh 7.5-15 cm), and seedlings (dbh < 7.5 cm regardless of height) by species was measured on 10 randomly placed quadrats (0.004 ha each) in each of the six harvest blocks before and after logging. Soil beneath the ground litter was classified as mesic if it felt damp or xeric if it felt dry. Small mammals were snap-trapped on each harvest block just prior to and immedi- ately after logging and on each harvested block and adjacent undisturbed sites for up to one vear thereafter at monthly intervals. Each sample consisted of 60 traps in three lines with 10 trap stations 16 m apart per line and 2 traps 3 m apart per station. Traps were baited with peanut butter and checked daily for 3 consecutive days for a total of 180 trap davs (TD) per sample (one trap set for 24 hours equals one trap day). The species; sex; age (juvenile or adult); length of body, tail. 'Department of Fishery- and Wildlife Biology, Colorado State University. Fort Collins, Colorado 80522. Reprints: Box 2705, Jackson. Wyoming 83001. ■Department of Biology, Idaho State I'niversity. Pocatello. Idaho 83209. 183 184 Great Basin Naturalist Vol. 40, No. 2 right hind foot, and right ear; and reproduc- tive status were recorded for all individuals trapped. Testes length for males and numbers of embryos or placental scars for females were also recorded. Age was based on pelage, size, and reproductive status. Statistical tests were based on the Chi square method of analysis imless otherwise indicated. Results and Discussion Effects of Logging on Vegetation Collectively, logging altered approx- imately 24 percent of the study area. Table 1 shows the harvest schedule. Selective cutting removed a mean of 52, 52, and 77 percent of the trees, saplings, and seedlings, respective- ly, and mesic ground conditions persisted for one year on these sites (Table 2). Clear- cutting removed 88, 70, and 79 percent of the trees, saplings, and seedlings, respective- ly, but ground conditions changed from mes- ic to xeric within 9 months after harvest. Al- though not quantitatively measured, logging and skidding drastically disturbed understory vegetation and litter in both clear and selec- tive cuts. Effects of Logging on Small Mammals Five species were captured: red-backed voles, deer mice, western jumping mice {Zapiis princeps), yellow pine chipmunk (£«- tamius amoenus), and masked shrews {Sorex cinereus). Ninety-eight percent of 478 cap- tures were voles and deer mice; only these two species are discussed here. Abundance: Voles were significantly (P<0.01) more abundant on mesic soils (76 percent of 408 captures. Fig. 1) throughout the study, but deer mice were significantly (P<0.01) more numerous on the one xeric site sampled 10 months after clearcutting (80 Table 1. Logging schedule for the six timber harvest blocks. Toppings Lakes Study Area, Bridger-Teton National Forest, Wyoming. Size Harvest Harvest schedule Block Interval No. (ha) method Cut Skidded (days) 1 19.4 Selective cut 8/19/76 8/21/76 2 2 9.6 Clearcut 9/08/75 9/10/75 2 3 75.0 Selective cut 9/15/76 10/31/76 46 ^ 8.1 Selective cut 9/10/76 9/10/76 5 9.6 Selective cut 9/09/75 8/20/76 315 6 36.4 Selective cut 9/13/76 9/23/76 10 ^Logging was suspended after 4 ha were logged aiid it became apparent that the expected timber voKimes were not there. Table 2. The changes in tree overstory densities for clear and selective cuts on the Toppings Lakes Study Area, Bridger-Teton National Forest, Wyoming. Mean size of harvest block (\mf Mean number per hectare* before logging after logging Selective cuts Trees Saplings Seedlings Clearcut Trees Saplings Seedlings 29.7 ± 27.7 SD 9.6 1064 ± 233 866 ± 84 11027 ± 4124 1213 743 17203 507 ± 47 416 ± 196 2592 ± 545 150 230 3770 "Means and standard deviations for selective cuts (N = 5). June 1980 Campbell, Clark: Rodent Ecology 185 percent of 60 captures). Koehler et al. (1975) found similar results in Idaho on undisturbed niesic sites and on sites xerified by forest fire. Red-backed voles apparently require a heavy cover of vegetation or logs (Gashwiler 1959, LaBue and Darnell 1959, Hooven 1969, Krefting and Ahlgren 1974) and reside pri- marily in cool, damp forests (Townsend 1935, Bailey 1936). The greater diversity of under- story plants of mesic sites (Daubenmire and Daubenmire 1968) apparently provide food and cover for red-backed voles, but deer mice prefer xeric habitats. Species composition on logged and un- logged mesic sites (selective cut #5) did not differ significantly (P>0.10) one year after logging (Fig. 2). Conversely, the disturbed xeric site (clearcut 2) had a highly significant (P<0.01) change in species composition fol- lowing harvest (Fig. 2). Preharvest and imme- diate postharvest data (September and Octo- ber 1975, respectively) showed a community composed primarily of red-backed voles (73 percent of 113 captures). Nine to 12 months after harvest (June to September, following winter inaccessibility) composition had changed to 80 percent deer mice of 60 ro- dent captures. These deer mouse capture rates indicate a larger population than that on the original, undisturbed forest. This is at- tributed to the xerification of this site as a re- sult of logging. Similar increases of deer mice were observed by Tevis (1956), Gashwiler (1959, 1970), Koehler et al. (1975), and Hoov- en and Black (1976) in forests altered by tim- ber harvest and forest fires. Clearcuts may be more attractive to deer mice (Gashwiler 1959) because they tend to move into dis- turbed, depopulated areas (Stickel 1946). Age ratios: On newly logged selective cuts, juveniles outnumbered adults 4.1 : 1.0 for red-backed voles and 5.0 : 1.0 for deer mice, but 9 to 12 months later juveniles had decreased significantly (P<0.01) to 0.8 : 1.0 for voles and 0.6 : 1.0 for deer mice. Juveniles also outnumbered adults on the newly clear- cut site 1.6 : 1.0 for voles and 1.8 : 1.0 for deer mice. 90 -■ 80 70 60 50 -■ 40 30 20 10 RED-BACKED VOLES DEER MICE LOGGED (mesic ) UN LOGGED (mesic) LOGGED (mesic) UNLOGGED (xeric) SELECTIVE CUTS CLEAR CUTS Fig. 1. Percent captures of red-backed voles and deer mice on logged and unlogged areas. Bridger-Teton National Forest, Wyoming. Figures at the top of the bars equal the number of animals trapped. 186 Great Basin Naturalist Vol. 40, No. 2 Juveniles also comprised 75 percent of all deer mice captured in a recent burn (Sims and Buckner 1972). Powell (1972) found three times as many red-backed vole juve- niles in a recently blown down forest than in an undisturbed forest. He concluded that standing forests were preferred red-backed vole habitat and adults drove juveniles into the less preferred, disturbed habitat (presum- ably via aggressive behavior). Sex ratios: The overall sex ratio for red- backed voles was 1.2M:1.0F, comparable to 1.3M:1.0F found in an Oregon study (Gash- wiler 1959). Adults had an even sex ratio, but juvenile males significantly (P<0.05) out- numbered juvenile females (1.5M:1.0F). Vole sex ratios (adults, juveniles, and total) did not differ significantly (P>0.05) in unlogged mesic, logged mesic, or logged xeric sites. Sex ratios for all deer mice captured (1.0 M:1.0 F); for adults (1.6 M:1.0 F); for juve- niles (0.6 M:1.0 F); and for logged mesic, un- 70 50 ■- 40 30 -- 20 10 y 70 40 30 20 -- 10 Block 2 - Clearcut Sept Jun Jul Aug Sept 1975 1976 Block 5 - Selective Cut ^ # # Total Captures #— — —• Red-backed Voles # # Deer Mice -\ 1 1 H Block 2 - Unlogged y —I 1 1 1 H Sept Jun Jul Aug Sept 1975 1976 Block 5 - Unlogged Fig. 2. Numbers of red-backed voles and deer mice trapped on logged and unlogged sites, Bridger-Teton National Forest, Wyoming (September 1975, Jvme-September 1976). June 1980 Campbell, Clark: Rodent Ecology 187 logged mesic, or logged xeric sites did not differ significantly (P>0.05) from 1.0 M : 1.0 F. Reproduction: Reproductive data for June-September 1976 showed that most re- production ceased after August (Tables 3 and 4). Males of both species with scrotal testes were captured through August but not in September. Packard (1968) and Clark (1973) used male rodents exhibiting scrotal testes as an indication of population breeding. Fe- males of both species carried embryos and were lactating from June to September, but September embryos were near term (Table 4). Mean litter size and SD based on embryo and placental scar counts was 6.0 ± 0.9 (range 4-9) for 44 red-backed voles and 6.1 ± 0.3 (range 6-7) for 10 deer mice. These were similar to other studies of red-backed voles in Crand Teton National Park (Clark 1973) and of deer mice in northwestern Wyoming (Clark 1975, Long 1964). Repro- ductive timing or litter sizes for either spe- cies did not differ between logged and im- logged areas. Body measurements: Linear measure- ments for red-backed voles and deer mice are shown in Tables 5 and 6, respectively. Adult females of both species were insignificantly larger than males. No significant (P>0.05, AOV) intraspecific age differences in mea- surement of either sex were observed be- tween logged and unlogged areas 9 to 12 months after harvest. Male red-backed voles from this study and nearby Grand Teton Na- tional Park (N' = 7, Clark 1973) did not differ significantly (P>0.10, t-test). Conclusions Both selective cutting that removed about 57 percent of the trees and clearcutting of spruce-fir forest resulted in an increase of juvenile red-backed voles and deer mice two weeks to a month after logging that was not apparent 9 to 12 months later. Clearcutting changed species composition 9 to 12 months later from predominantly red-backed voles on unlogged areas to predominantly deer mice. The change was attributed to soil xeri- Table 3. Male reproductive condition of red-liacked voles and deer mice, Bridger-Teton National Forest. Wyo- ming (June-September 1976). Number of Number of red-hacked voles deer mice Condition June J"iy Aug. Sept. Total June July Aug. Sept. Total Scrotal 9 7 12 28 4 9 1 14 Nonscrotal 11 9 20 1 13 6 20 Total 9 7 2.3 9 48 4 10 14 6 34 Table 4. Reproductive condition of red-backed vole and deer mouse adult females. Bridger-Teton National For- est, Wyoming (June-September 1976). N umber of Number of red-backed voles deer mice Condition June July Aug. Sept. Total June July Aug. Sept. Total Nimiber of females examined 8 5 13 18 44 1 6 10 1 18 Percent pregnant 50 80 23 72 54 100 67 20 100 44 Niunber of embrvos Mean 6.3 6.8 6 5.9 6.1 6 6.5 6 6 6.3 Range 6-7 5-9 6 4-9 4-9 6 6-7 6 6 6-7 Percent with placental scars 50 40 -"- 28 48 .33 80 .56 Number of placental scars Mean 5.5 6 6 .5.6 5.8 6 6 6 Range 5-6 6 6 4-6 4-6 6 6 0-6 Number of females lactating 4 6 13 17 40 1 10 1 17 188 Great Basin Naturalist Vol. 40, No. 2 fication on the clearcut. Voles continued to predominate on selectively cut sites, and soils remained mesic there. For both species adults consistently outnumbered juveniles in June, July, and August in both logged and unlogged areas, but juveniles outnumbered adults in areas newly logged in September. Sex ratios, timing of reproduction, litter size, and body measurements for both species did not differ significantly between unlogged, selectively cut, or clearcut areas 9 to 12 months after logging. Acknowledgments This study was supported by the Bridger- Teton National Forest, Jackson, Wyoming. Special thanks is extended to Mr. G. GreuU, Wildlife Biologist, U.S. Forest Service. Mr. W. Barmore offered valuable criticism of the manuscript. Table 5. Body measurements of 258 red-backed voles from unlogged, clearcut, and selectively cut areas, Bridger- Teton National Forest, Wyoming (June-September 1976). Sex and age* Mean length, SD, and (range) in mm Body Tail Ear Hind Foot Control M2+ (n=18) F2+ (n = 23) Ml-(n = 40) Fl-(n = 26) 139.4 ±8.8(1 19-152) 143.4 ±10.9(118-158) 115.2 ±12.1(88-137) 112.2 ±10.5(90-132) 39.1 ±2.6(34-45) 40.5 ±4.4(26-47) ,33.0 ±4.5(26-43) .33.2 ±3.7(25-42) 15.8 ±1.8(1,3-21) 16.5± 1.1(14-19) 14.9± 1.8(10-17) 14.9± 1.7(10-18) 18.4 ±0.9(17-19) 18.0 ±1.0(16-19) 16.9 ±1.2(15-18) 17.0± 0.7(15-18) Clearcut M2+ (n = 3) F2+ (n = 4) Ml-(n = 4) Fl-(n = l) 147.7 ±6.4(144-155) 41.0 ±3.5(39-45) 14.3 ±0.6(14-15) 18.7 ±0.6(18-19) 139.6 ±10.4(125-147) 40.3 ±2.9(37-44) 17.0± 0.8(16-18) 17.8 ± 1.0(17-19) 113.3 ±13.8(100-130) 33.5 ±6.2(27-40) 13.0± 2.0(12-16) 16.8 ±0.5(16-17) 117.0- - .35.0- - 17.0- - 18.0- - Selective cut M2+ (n = 27) F2+ (n = 17) Ml-(n = 59) Fl-(n = 36) 138.9± 10.1 (115-160) 40.8 ±3.5(26-48) 15.9± 1.7(12-18) 18.4 ± 1.0(16-19) 142.7 ±11.2(122-159) 40.6 ±4.7(37-47) 15.9± 1.8(11-18) 17.9± 0.8(16-19) 121.9 ±8.5(96-1.37) ,35.6 ±.3.5(26-42) 15.6± 1.6(11-19) 17.3 ±0.7(16-19) 115.0± 10.7(93-138) .3,3.5 ±,3.6(25-40) 15.4 ± 1.9(10-18) 17.2 ±0.7(16-19) *2+ = adult; 1- = juvenile. Table 6. Body measurements of 118 deer mice from unlogged, clearcut, and selectively cut areas, Bridger-Teton National Forest, Wyoming (June-September 1976). Mean length, SD, and (range) in mm Sex and age° Body Tail Ear Hind foot Control M2-I- (n=ll) F2+ (n = 5) Ml-(n = 8) Fl-(n = 9) Clearcut M2+ (n = 12) F2+ (n = 10) Ml-(n=9) Fl-(n=17) Selective cut M2+ (n = 12) F2+ (n = 4) Ml-(n = 6) Fl-(n=15) '2+ = adult; 1- = juvenile. 157.5± 11.1(140-17,5) 156.6± 10.1(146-170) 139.4 ±5.0(133-146) 146.0±6.,3(137-155) 68.4 ±4.8(62-77) 69.2±4..3(63-7.3) 6.3.9 ±2.0(61-66) 67.0 ±3.4(62-72) 158.7± 11.1(148-175) 69.8 ±5.0(6,3-77) 166.3 ± 14..3(144-188) 7.3.4 ± 7.1(62-85) 146.4 ±7.2(133-156) 67.4 ±3.6(61-74) 142.5 ± 7.6(130-155) 63.4 ± 5.2(55-72) 154.1 ± 10.1(141-174) 67.4 ±5.2(61-76) 160.1 ± 17.8(145-183) 69.8 ± 8.3(62-80) 146.8 ±6.6(139-157) 66.5 ±3.7(64-74) 143.2 ±9.6(128-160) 65.0 ±5.0(57-74) 18.5 ±0.8(1.3-20) 18.8 ±0.5(1.3-19) 18.3 ±1.0(17-20) 18.8 ±0.8(17-20) 18.0 ±1.7(15-20) 18.3 ±1.3(17-20) 18.7 ±1.0(18-20) 17.6 ±1.8(14-20) 18.3 ±1.4(15-20) 18.5± 1.0(17-19) 18.2 ±0.8(17-19) 17.9 ±1.5(16-20) 18.4 ±0.9(19-20) 19.6 ±0.9(19-21) 19.1 ±0.6(18-20) 18.6± 1.0(17-20) 19.4 ±0.7(19-21) 19.3 ±0.7(18-20) 19.1 ±0.9(18-20) 18.5 ±1.0(17-20) 19.1 ±0.,5( 18-20) 19.3 ±1.0(18-20) 19.2 ±1.0(18-20) 18.8 ±0.8(17-20) June 1980 Campbell, Clark: Rodent Ecology 189 Literature Cited Bailey, V. 1936. The mammals and life zones of Oregon. North American Fauna 55. 416 pp. Beetle, A. A. 1961. Range survey of Teton County, Wyoming. Part 1: Ecology of range resources. Univ. Wyoming Agr. Exp. Stn. Bull. No. 376R:l-42. Brady, S. 1974. Environmental impact study of Top- pings Lake timber sale. Bridger-Teton National Forest, Jackson, Wyoming. Mimeo. 20 pp. Clark, T. W. 1973. Local distributions and interspecies interactions in microtines. Grand Teton National Park, Wyoming. Great Ba.sin Nat. 33:205-217. 1975. Ecological notes on deer mice in Grand Te- ton National Park, Wvoming. Northwest Sci. 49(1): 14-16. Clark, T. W., and T. M. Campbell. 1976. Population organization and regulatory mechanisms of pine martens in Grand Teton National Park, Wyo- ming. Paper presented at Conference on Re- search in National Parks, Nov., 1976, New Or- leans, Louisiana. Daube.nmire, R., and J. B. Daubenmire. 1968. Forest vegetation of eastern Washington and northern Oregon. Washington .\gr. Exp. Stn. Tech. Bull. No. 22, 60 pp. Department of Commerce. 1975. Climatological data of Wvoming. Natl. Ocean, and Atmos. Admin., .\sheville, N.C. Nos. 1-12. Gashwiler, J. S. 1959. Small mammal study in west- central Oregon. J. Mammal. 40(1): 128-1.39. 1970. Plant and mammal changes on a clearcut in west-central Oregon. Ecology 51(6): 1018-1026. Hoffman, G. R. 1960. The small mammal components of six climax plant associations in eastern Wash- ington and northern Idaho. Ecology 41(.3):571-572. HoovEN, E. F. 1969. The influence of forest succession on populations of small animals in western Ore- gon in H. E. Black, ed. Wildlife and reforestation in the Pacific Northwest. Oregon State Univ., Corvallis. HoovEN, E. F., and H. C. Blac:k. 1976. Effects of some clear-cutting practices on small-mammal popu- lations in western Oregon. Northwest Sci. 50(4): 189-208. Knight, C. A. 1973. Soil resource inventory manual. Bridger-Teton National Forest, Jackson, Wyo- ming. KoEHLER, G. M., W. R. Moore, and A. R. Taylor. 1975. Preserving the pine marten: management guidelines for western forests. Western Wildlands 2(3) :3 1-36. Kreftinc, L. W., and C. E. .\hlgren. 1974. Small mam- mals and revegetation changes after fire in a mixed conifer-hardwood forest. Ecology 55(6): 1391-1398. LaBue, J., and R. M. Darnell. 1959. Effect of habitat disturbance on a small mammal population. J. Mammal. 40(3):425-437. Long, C. A. 1964. Comments on reproduction in the deer mouse of Wyoming. Trans. Kansas Acad. Sci. 67:149-153. Love, J. D., and J. C. Reed, Jr. 1968. Creation of the Teton landscape. Grant Teton Nat Hist, .\ssoc. Publ., Moose, Wyo. 120 pp. Packard, R. L. 1968. An ecological study of the fulvous harvest mouse in eastern Texas. Amer. Midi. Nat. 79:68-88. Powell, R. A. 1972. A comparison of populations of bo- real red-backed voles {Cletlirionomys gapperi) in tornado blowndown and standing forest. Can. Field Nat. 86(4):377-379. Reed, J. F. 1952. The vegetation of the Jackson Hole Wildlife Park, Wyoming. Amer. Midi. Nat. 48(3): 700-729. Sims, P., and C. H. Buckner. 1972. The effects of clear- cutting and burning of Pinus barksiana forests on the populations of small mammals in south-east- ern Manitoba. Amer. Midi. Nat. 90(1):228-231. Stickel, L. F. 1946. The source of animals moving into a depopulated area. J. Mammal. 27(4):302-307. Tevis, L. 1956. Response of small mammal populations to logging of Douglas fir. J. Mammal. 37(2): 189-196. Townsend, N. T. 1935. Studies on small mammals of central New York. Roosevelt Wildl. .\nnals. 4:6-120. TERMINAL BUD FORMATION IN LIMBER PINE Ronald M. Lanner' and James A. Bryan' ,\bstr.'\ct.— The progress of bud development was studied in limber pines growing in the mountains of north- eastern Utah. Initiation of new bud scales began in mid-June, several weeks after elongation of the current shoot had begim. Needle primordia first appeared in September and continued to form through the winter, until all were pres- ent in May. This winter activity is believed to be fostered by surface temperatures on the terminal buds considerably higher than ambient air temperatures. The annual shoot of limber pine (Pinus flexilis James), a five-needled species com- mon in the Rocky Mountains, consists of a monocyclic spring shoot formed by the elon- gation of a winter bud. This is the most fa- miliar shoot development pattern in northern pines and has been classified as the Resinosa pattern (Lanner 1976). But even among spe- cies of this habit, there is diversity in the de- velopmental schedule of bud morphogenesis. For example, in some species formation of the new winter bud begins while the old one is still elongating, but in others bud formation is delayed until the cessation of current-sea- son shoot growth. Further, the timing of .short-shoot and needle morphogenesis is also subject to variation. In this report we de- scribe the annual cycle of development of limber pine terminal buds to resolve the questions of when the short shoots and nee- dles are formed, both in terms of calendar date and in regard to the growth stage of the spring shoot. The trees studied grow at an elevation of 2130 m on a steep southeast slope in Logan Canyon, northeastern Utah. They vary from 12 to 27 cm DBH and 4 to 7 m in height. At each of 18 collection dates during 1978, 2 terminal buds of vigorous first-order branches in the upper crown were harvested from each of at least 2 of the 18 study trees. Buds were stored in formaldehyde-acetic acid-eth- yl alcohol (FAA), dissected with standard mi- cro-dissecting tools, and examined at 12-lOOx with a Wild M-5 stereomicroscope. Two shoots had steel pins inserted at the base of tlie bud as a datum for observations of shoot elongation. On each of the harvested buds we dis- sected at least two short-shoot budlets from the proximal (basal) end of the bud, and two from the distal (apical) end. We counted the budlet scales (future fascicle sheath scales) and needle primordia, if any. The study took place during a single calendar year, so we ac- tually observed the late development and elongation of the 1978 spring shoot and the early development of the 1979 spring shoot. Ideally, a study of this kind should begin with early morphogenesis of a bud and end with the maturation of the resulting shoot. Results One of the marked shoots started to elon- gate during the interval 20-30 May, and the other during the interval 30 May-5 June. These shoots completed their elongation growth prior to 8 July and 13 July, respec- tively. Final lengths of these shoots were 30.5 and 21.5 mm. Pollen was shed during the pe- riod 9-13 July. Initiation of the primary bud scales (cata- phvlls) of the newlv forming terminal bud be- gan between 11 and 18 June. After cataphylls formed, meristems appeared in the axils of most of them. These axillary meristems be- came the apical meristems of the short-shoot 'Department of Forestry and Outdoor Recreation, Utah State University, Logan, Utah 84322. 190 June 1980 Lanner, Bryan: Limber Pine 191 budlets that would later become needle fas- cicles. Scales formed on these budlets are fu- ture fascicle sheath scales. Sheath scale ini- tiation began in proximal budlets in early July, and in distal budlets in early August. Scale production continued in both types of budlets luitil about mid-Januarv. The earliest needle primordia started to appear in proximal budlets in late August and continued to appear over a period of three months. Needle primordia in distal budlets did not begin to form until about mid-No- vember, but continued appearing up to early May, about 5.5 months later. When bud scale initiation began, about two-thirds of the elongation growth of the marked shoots had been completed. Bud scale initiation probably continued through- out the remainder of the elongation period. Sheath scale initiation began in proximal budlets about the time shoot elongation was ceasing, and it began in distal budlets after the cessation of shoot growth. When needle primordia started to form, shoot growth had been inactive for several weeks. The needle primordia formed in the spring developed prior to the onset of shoot elongation in late May. When the first buds were harvested 22 Jan- uary 1978, short-shoot budlets from proximal parts of the terminal bud contained 10-12 scales (x = 11.2) and all had their full com- plement of 5 needle primordia. But budlets located at the distal end of those terminal buds tended to have fewer scales (8-11; x = 10.0) and averaged only 1.7 needle primordia (Table 1). The difference in scale number was maintained even beyond 8 May, when, for the first time, all the distal budlets contained their full complement of 5 needle primordia (Table 1). Discussion and Conclusions The initiation of bud scales was first noted on 18 June, when the 1978 shoot had attained 68 percent of its final length, showing that bud morphogenesis began during the period Table 1. Progress of sheath scale and needle initiation in proximal and distal short shoot budlets of first-order terminal buds of Limber Pine, and of shoot elongation. Proximal budlets Needle Distal budlets Needle Length of 1978 number rimordium primordium bud/shoot as Scale ninnber Scale number number percent of final length 1978 Date X range X X range X 1978 annual shoot 22 Jannary 11.2 10-12 5.0 10.0 8-11 1.7 31.5 21 March' 10.0 9-12 5.0 9.6 9-10 2.5 31.5 12 April 10.3 9-12 5.0 9.5 9-10 3.3 31.5 8 May 10.0 9-11 5.0 10.6 10-11 5.0 31.5 20 May 9.5 8-11 5.0 9.5 9-10 5.0 31.5 4 Jnne 10.0 10 5.0 9.5 9-10 5.0 45.0 1 1 Jnne 13.0 13 5.0 1979 annual 11.0 shoot 11 5.0 55.5 1 1 Jnne 55.5 IS June 68.0 2Jnly 95.0 13 July 2.0 2 100 23 July 4.5 3-6 1(X) 2 August 5.0 5 1(K) 16 August 6.0 6 0.75 0-2 100 26 August 6.0 .5-7 2.3 2-3 1(X) 15 September 8.0 8 5.0 7.3 6-9 100 5 October 8.3 8-9 3.0 7.3 6-9 100 15 November 8.8 8-10 2.2 7.0 6-8 100 9 December 8.0 8 5.0 8.0 8 1.8 100 192 Great Basin Naturalist Vol. 40, No. 2 of shoot elongation. In this regard limber pine resembles P. strobus L. and P. lamber- tiana Dougl. (Lanner 1976). It provides fur- ther evidence that shoot elongation does not inhibit the initiation of lateral structures on the shoot apical meristem. Bud morphoge- nesis in limber pine is delayed, however, in comparison to that of lodgepole pine (P. con- torta Dougl.), one of its associates in this area (Van Den Berg and Lanner 1971). In lodge- pole, initiation of the new bud and elonga- tion of the shoot began almost simultaneously early in May. A given stage of development— i.e., attain- ing a certain number of sheath scales or a certain number of needle primordia— is reached earlier in proximal than in distal short shoots. Thus, in the 1978 winter buds, the full complement of 5 needle primordia was present in proximal short shoots in Janu- ary collections, but was not found in distal short shoots until May. This is another case of the developmental gradient in short-shoot maturation described in the more complex buds of lodgepole pine (Van Den Berg and Lanner 1971), the much larger buds of slash pine, P. elliottii Engelm. (Lanner 1978), and in eastern white pine (Owston 1969). Perhaps the most unusual finding reported here is that morphogenetic activity contin- ued in wintering limber pine buds. Buds col- lected early in 1978 showed periodic increas- es in needle primordium number in distal budlets. Buds collected late in 1978 showed consistent increases in needle primordium number in both proximal and distal budlets, and in sheath scale number in distal budlets (Table 1). Yet temperatures at a nearby tem- porary weather station at a comparable ele- vation fell as low as -21 C in January and -12 C in November and December (pers. comm., S. A. Loomis). In lodgepole pine studied just a few miles from this site, budlets overwintered with less than their full com- plements of sheath scales and needle pri- mordia, but no changes were noted during the winter (Van Den Berg and Lanner 1971). Budlets of slash pine actively initiated scales and needle primordia during the winter months, but this was in the much milder cli- mate of Florida (Lanner 1978). Seed cone primordia have also been re- ported as morphogenetically active during the winter. Duff and Nolan (1958) observed changes in ovulate strobili of red pine (P. resi- nosa Ait.) between October and January in the cold climate of Chalk River, Ontario. Gif- ford and Mirov (1960) also reported female strobilus development in ponderosa pine (P. ponderosa Laws.), but in the milder climate of the Sierra Nevada west slope. Such meristematic activity may be permit- ted by surface temperatures considerably higher than those of the ambient atmosphere. For example, Tranquillini and Turner (1961) have reported maximum monthly temper- atures of Swiss stone pine (P. cernbra L.) nee- dles 2.7 and 7.4 C higher than air temper- atures during November and March, respectively, in the Austrian Alps. In March, needles reached a maximum of 18.4 C, though the mean air temperature for that month was only 0.3 C. At our study site, even in January, the coldest month of the year, air temperature on three occasions attained al- most 6 C. Bud surface temperatures may have reached as high as 12 C or more, well above the apparent threshold for meristemat- ic activity. Literature Cited Duff, C. H., and N. J. Nolan. 1958. Growth and mor- phogenesis in the Canadian forest species. III. The time scale of morphogenesis at the stem apex of Pintis resinosa Ait. Canadian J. Bot. 36:687-706, 4 pis. GiFFORD, E. M., Jr., and N. T. Mirov. 1960. Initiation and ontogeny of the ovulate strobilus in ponde- rosa pine. Forest Sci. 6:19-25. Lanner, R. M. 1976. Patterns of shoot development in Pinus and their relationship to growth potential. Pages 223-243 in M. G. R. Cannell and F. T. Last, eds. Tree physiology and yield improve- ment. Academic Press, New York and London. 1978. Development of the terminal bud shoot of slash pine saplings. Forest Sci. 24:167-179. Owston, P. W. 1968. The shoot apex in eastern white pine: its stnicture, seasonal development, and variation within the crown. Canadian J. Bot. 47:1181-1188. Tranquillini, W., and H. Turner. 1961. Untersuchu- ngen iiber die Pflanzentemperaturen in der sub- alpinen Stufe mit besonderer Beriicksichtung der Nadeltemperaturen der Zirbe. Mitteil. Forstlich. Bundes-Versuchs. Mariabrunn 59(1): 127-151. Van Den Berg, D. A., and R. M. Lanner. 1971. Bud de- velopment in lodgepole pine. Forest Sci. 17:479-486. STINGER UTILIZATION AND PREDATION IN THE SCORPION PARUROCTONUS BOREUS Bnice S. Gushing' and Anne Matherne' .\bstr.\ct.— The iitihzution of the stinger and the predatory technique of the scorpion, Paruroctanus boretis, was studied under laboratory conditions. During the study, 83 feedings were observed. Age of the scorpions and the per- centage of prey stung by them were used to classify the scorpions into groups. The scorpions aged 1.3-61 days always stung prey, .\fter 62 days the scorpions began to selectively utilize the stinger. Utilization declined until it reached .30 percent in the adult stage. The stinger is apparently necessary for prey capture only in the earlv life stages. The role of the stinger in scorpion behav- ior has never really been studied or estab- lished. The available information is vague and inconsistent. Alexander (1959) reported that different groups of scorpions utilize the stinger differentially. Pocock (1893) and Stahnke (1966) stated that scorpions paralyze prey only when it does not submit to passive consumption. Finally Hadley and Williams (1968) reported that Vepvis confusus and Paruroctonus mesaensis and P. baergi stung prey (Williams 1972). These reports leave the adaptive significance of this potent device in doubt. In this study, we attempted to estab- lish the use of the stinger through controlled experimentation and observation. Methods and Materials Scorpions were collected from south- western Oregon on the Malheur National Wildlife Refuge and adjacent region. They were kept in 10-gallon terrariums containing three inches of soil and flat rocks from the natural habitat. In most ca.ses, two scorpions were placed in each tank to induce in- traspecific responses. A large population of grasshoppers and a small number of beetles, crickets, and termites were found in the area, and the.se were selected in relative propor- tions as the prey. To maintain natural conditions, the terra- riums remained outdoors except during peri- ods of observation. Observations were con- ducted at night using red light. Red light provided visibility for us, but apparently did not affect the scorpions, which possess vision in the blue and ultraviolet wavelengths (Machan 1968). Results Over a two-year period several different groups of Paruroctonus boreus were ob- served. Predation techniques were the same for scorpions of all ages and sizes. Emergence from cover occurred between 2130 and 2300 hours. If emergence did not occur by 2.300 the scorpions did not forage that night. After emergence, contact with prey was estab- lished through random encounter or active stalking. When actively stalking, the scor- pions traveled with the pedipalps extended forward and held apart at a distance approx- imately equal to the maximum width of the abdomen. The telson was arched over the ab- domen with the caudal vesicle above the midabdomen. When a potential prey was de- tected the scorpions rushed it. Upon contact, the scorpions used their pedipalps to grasp the prey by one or more appendages. If stinging occurred at all, it oc- curred at this time. The telson was arched over the abdomen and at the same time the abdomen was quickly raised. This imparted a downward stabbing motion which allowed 'Department of Wildlife Biology, University of Montana. Missoula, Montana 59812. "Department of Zoology, Louisiana State University. Baton Rouge, Louisiana 7080.3. 193 194 Great Basin Naturalist Vol. 40, No. 2 the stinger to penetrate the prey's abdomen. Resistance by the prey subsided within one minute after it was stung. Whether or not the prey was paralyzed, it was held motionless in the pedipalps for 10 to 30 minutes and then transferred to the chelicerae. Upon leaving the site of capture the prey was slung, ven- tral side up, onto the cephalothorax.The prey was carried about the terrarium in this posi- tion for up to several hours. When walking, the scorpions waved their pedipalps in front of their path in a "blindman" fashion, a slow exploratory touching. Except when scorpions were with young, prey was taken beneath cover for con- sumption. Consumption time varied between 2 and 48 hours. Feeding began at the head of the prey and continued until the prey was consumed. The hard exoskeleton of beetles was left as an empty husk. Several of these husks were found with scorpions in the field. All scorpions used the same feeding tech- niques. However, the scorpions also under- went regular periods of nonfeeding which lasted up to five months. The stinger was removed from two adult scorpions. These scorpions fed six times and utilized the same techniques as unimpaired individuals. However, they never attempted to sting any of the prey. During intraspecific aggression or canni- balism, the method of capture was as de- scribed above with minor modifications. If there was a significant difference in size, the smaller scorpion attempted to avoid conflict, but the larger one often pursued. When ag- gression occurred, the scorpions grasped each other by the pedipalps and repeatedly at- tempted to sting. A size difference always re- sulted in the death of the smaller scorpion. Consumption proceeded normally after im- mobilization. In two instances of the scor- pions being the same size, both animals were killed. Immature scorpions did not capture prey until they were 13 days old. Prior to this time they consumed their casting left at birth and their first exuvium. After the juveniles dis- persed at 9 to 11 days, the female began feeding with an alteration in feeding tech- nique. The adult female consumed prey in the open and its young congregated about the adult's cephlathorax. On day 14 the young began to capture prey. Table 1 sum- marizes the percentages of prey stung by the early instars and all other age groups. Discussion and Analysis No age group beyond 84 days was ob- served due to our inability to keep juveniles alive. This resulted mainly from a high de- gree of cannibalism and mishandling of a few remaining scorpions. Data on cannibalism were excluded from Table 1 because of the bias it would introduce inasmuch as in- traspecific aggression always elicited stinger utilization. Two scorpions per tank and the limited dispersal range of the young led to an unnatural increase in incidents of cannibal- ism. Data on the six feedings by the scorpions with stinger removed were also excluded due to their inability to sting. During active stalking, the scorpions rushed prey. Some stimulus must have been present which alerted the scorpions. Paruroc- tonus boreus, like other desert scorpions, may be able to detect and utilize Rayleigh waves for prey location (Brownell 1977). A Rayleigh wave is a slow-moving secondary vibration created by movement and propagated through sand. The stinger was not essential for feeding by the adult scorpions. A low percentage of prey was paralyzed, and stinger-impaired individ- uals were able to feed without difficulty. Scorpions are also capable of surviving pro- longed periods without food (Stahnke 1966). Considering these factors, we suggest that the amount of food which would be lost to an adult incapable of stinging would not have a significant or deleterious effect upon its sur- vival. If stinging occurs, it is triggered by two stimuli. One, as stated by Pocock (1893) and Table 1. Relationship between age and stinger utili- zation. Age group Number of Number Percent in days feedings stung stung 01-12° 00.0 13-61 40 40 100.0 62-84 13 9 69.2 Adult 30 10 30.0 "Fed on exuviae and adult pellets. June 1980 CusHiNx;, Matherne: Scorpion Predation 195 Stahnke (1966), is an attempt by the prev to resist capture. This is not the onlv stiuiuhis. Struggling hard-bodied or powerful prey, such as grasshoppers, were stung. Termites and other soft-bodied prey were held in the pedipalps until resistance subsided. The fac- tors that elicited a sting were resistance in combination with the characteristics of the prey species. This strongly suggests a form of selective stinger utilization. Tlie selection process develops over time. The first instars feed on the exuviae and pos- sibly on small pellets dropped by the adult during feeding (Stahnke 1966). Utilization of pellets for food suggests the reason for the change in feeding technique by the adult fe- male, with the young gathered about her cephalothorax. The next age group, 13-61 days, paralyzed all prey (Table 1). The cause of this may be that the pedipalps were not sufficientlv developed at this point to hold prey against struggle. Therefore, in order to insure the maximimi number of feedings and promote growth and development, the juve- niles must sting prey at first contact. As de- velopment occurs the pedipalps strengthen, and certain prey types, such as small ter- mites, no longer must be paralyzed. This would reduce the use of toxin and be energy efficient by reducing the manufacture of more toxin. Stinger utilization drops from 100 to 30 percent in the adults (Table 1). This decrease began about the second month and continued until the adult stage. The actual percentage utilization in the group aged 62-84 days may have been biased in that this group was not fed a representative class of prey, but instead was fed whatever small insects and arachnids happened to be available. Nevertheless, this group still demonstrates the beginning of the process of differential selection for stinger utilization in that not all prey was stung. In conclusion, the stinger fimctions as a necessary device for prey capture by the early instars. As physical development oc- curs, the pedipalps can hold certain types of prey and there is a reduction in the use of the stinger. This decline continues until the adult stage, where only a small percentage of prey is stung and these are not essential for survi- val. However, the stinger is still utilized for intraspecific aggression and possibly for de- fense. Acknowledgment We thank Dr. John Mates, University of California, Davis, for his early help in this project, and Dr. Andrew Sheldon, Depart- ment of Zoology, University of Montana, for his critical review of this manuscript. Literature Cited Alex.\nder, a. J. 1959. A survey of the l)iologv of scor- pions of South Africa. Afr. Wildl. 13(2):99-106. Brownell, p. H. 1977. Conipressional and surface waves in sand: used by desert scorpions to locate prey. Science 197:479-481. RIDLEY, N. F., AND S. C. WiLLiAMs. 1968. North .Ameri- can scorpions in relation to feeding. Ecology 49(4):726-734. Macha.n, L. 1968. Spectral sensitivity of scorpions' eyes and possible role of shielding pigment effect. J. Exp. Biol. 49(1):95-105. PococK, R. I. 189.3. Notes upon the habits of some living scorpions. Nature. 48:1()4-I()V. Stahnke, H. L. 1966. Some aspects of scorpion behavior. Southern Calif. Acad. Sci. Bull. 62(2)65-80. W'illia.ms, S. C. 1972. Four new scorpion species be- longing to the genus Paruroctonus. Occasional Papers^Calif. Acad. Sci., No. 94. NOTICE TO CONTRIBUTORS Original manuscripts in English pertaining to the biological natural history of western North America and intended for publication in the Great Basin Naturalist should be directed to Brigham Young University, Stephen L. Wood, Editor, Great Basin Naturalist, Provo, Utah 84602. Those intended for the Great Basin Naturalist Memoirs should be similarly directed, but these manuscripts are not encumbered by a geographical restriction. Manuscripts. Two copies of manuscripts are required. They should be typewritten, double spaced throughout on one side of the paper, with margins of at least one inch on all sides. 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Reprint Schedule of the Great Basin Naturalist 2 pp. 4 pp. 6 pp. 8 pp. 10 pp. 12 pp. Each 100 copies $20 $24 $28 $32 $36 $40 additional 200 copies 28 32 36 40 44 48 2 pp. 300 copies 36 40 44 48 52 56 $4 Great Basin Naturalist Memoirs No. 1 The birds of Utah. By C. L. Hayward, C. Cottam, A. M. Woodbury, H. H. Frost. $10. No. 2 Intermountain biogeography: a symposium. By K. T. Harper, J. L. Reveal et al. $15. No. 3 The endangered species: a symposium. $6. TABLE OF CONTENTS Feeding ecology of Gila boraxobius (Osteichthyes: Cyprinidae) endemic to a thermal lake in southeastern Oregon. Jack E. Williams and Cynthia D. Williams 101 First record of the pallid hat (Antrozous palliclus) from Montana. Jeff Shryer and Dennis L. Flath '. 115 A Cliiivcantliiuiti spider bite. Dorald M. ,\llred 116 Identity of narrow-leaved Chitisotliamnits visciclifloni.s (Asteraceae). Loran C. .\nder- ll'i Ribulose diphosphate carboxylase activities in cold-resistant common mallow, Malva neglecta Wallr. and a cold-sensitive tomato, Lycopersicon esculentum L., Ace 55 var. William R. Andersen and Jack D. Brotherson 121 Recovery of Gambel oak after fire in central Utah. L. M. Kunzler and K. T. Harper .. 127 Relationships among total dissolved solids, conductivity, and osmosity for five A;- temia habitats (Anostraca: Artemiidae). Nicholas C. Collins and Gray Stirling 1.31 Spawning of the least chub (loticJtthijs phlegt'thuntis). Thomas M. Baiigh 1.39 Transferrin polymorphism in bighorn sheep, Otis canaclcn.^is, in Colorado. Patrick W. Roberts, Donald J. Nash, and Robert E. Keiss 141 The genus Eriogoniim Michx. (Polygonaceae) and Michel Gandoger. James L. Reveal 143 Parasites from two species of suckers (Catostomidae) from southern Utah. J. Craig Brienholt and Richard A. Heckmann 149 Soil water withdrawal and root distribution under grubbed, sprayed, and undis- turbed big sagebni.sh vegetation. David L. Sturges 157 Swarming of the western harvester ant, Pogononnjnnex occiclentalis. Dorald M. Allred 165 Relationship between environmental and vegetational parameters for understory and open-area communities. William E. Evenson, Jack D. Brotherson, and Rich- ard B. Wilcox 167 Seasonal activity pattern of Columbian ground sqviirrels in the Idaho primitive area. Charles L. Elliott and Jerran T. Flinders 175 Habitat and plant distributions in hanging gardens of the narrows, Zion National Park, Utah. George P. Malanson 178 Short-term effects of logging on red-backed voles and deer mice. Thomas M. Camp- bell III and Tim W. Clark 183 Terminal bud formation in limber pine. Ronald M. Lanner and James A. Bryan 190 Stinger utilization and predation in the scorpion Paiuwctoniis boreits. Bruce S. Gush- ing and -•Vnne Matherne 193 HE GREAT BASIN NATURALIST lume 40 No. 3 September 30, 1980 Brigham Young University MUS. COMP. ZOOl_. MAY 14'^ ' HARVAl-vO UMfVERSI-TY GREAT BASIN NATURALIST Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Life Science Museum; Stanley L. Welsh, Botany; Clayton M. White, Zoology. Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agricul- tural Sciences; Ernest L. Olson, Director, Brigham Young University Press, University Editor. The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been published from one to four times a year since then by Brigham Young University, Provo, Utah. In gener- al, only previously unpublished manuscripts of less than 100 printed pages in length and per- taining to the biological and natural history of western North America are accepted. The Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natu- ral history longer than can be accommodated in the parent publication. The Memoirs appears irregularly and bears no geographical restriction in subject matter. Manuscripts are subject to the approval of the editor. Subscriptions. The annual subscription to the Great Basin Naturalist is $12 (outside the United States $13). The price for single numbers is $4 each. All back numbers are in print and are available for sale. All matters pertaining to the purchase of subscriptions and back num- bers should be directed to Brigham Young University, Life Science Museum, Provo, Utah 84602. The Great Basin Naturalist Memoirs may be purchased from the same office at the rate indicated on the inside of the back cover of either journal. Scholarly Exchanges. Libraries or other organizations interested in obtaining either journal through a continuing exchange of scholarly publications should contact the Brigham Young University Exchange Librarian, Harold B. Lee Library, Provo, Utah 84602. Manuscripts. All manuscripts and other copy for either the Great Basin Naturalist or the Great Basin Naturalist Memoirs should be addressed to the editor as instructed on the back cover. U-80 650 48558 ISSN 0017-3614 The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 Volume 40 September 30, 1980 No. 3 SPATIOTEMPORAL VARIATION IN PHENOLOGY AND ABUNDANCE OF FLORAL RESOURCES ON SHORTGRASS PRAIRIE V. J. Tepedino''^ and N. L. Stanton^ Abstr.\c:t.— Phenolog\- and abinidance of the floral resources used by bees and other flower-visiting insects were recorded weekly from permanent quadrats for two years on two shortgrass prairie sites in the Laramie Basin. Wyom- ing. Each site was composed of three distinct plant communities. Residts suggest considerable spatiotemporal varia- bilit\ in floral resources. Specifically, most species at both sites showed the following temporal variation: (1) bloom times between one and two weeks earlier in the second year; (2) differences of approximately 1-2 weeks in bloom span between years; (3) substantial differences in abundance of flowers between years. Species blooming during the last half of the census period were significantly more variable in flower abundance between years than those bloom- ing earlier. Spatial variation was shown both by differences between sites and between plant communities within sites in the direction and magnitude of between-year changes in floral abundance. For example, floral production at Boulder Ridge in 1976 was much lower than in 1975, but at The Dirt Farm between-year differences were less pro- nounced and depended upon season. Similarity measures and cluster analysis .suggest differences in the structure of the bloom season at both sites between years, and a relatively rapid turnover of floral composition within years such that bees face a very different flora over the latter part of their flight seasons relative to that encountered initially. Evidence from other reports support the hypothesis of spatiotemporal unpredictability of floral resources. Spatial and temporal heterogeneity in re- source supply is receiving increasing atten- tion in ecological hypotheses concerning both species diversity and consumer foraging strategies (see reviews in Wiens 1976, Levin 1976). An important component of such re- source heterogeneity is its predictability (Wiens 1976). When resources are unpredic- table in time or space, generalists are hypoth- esized to dominate the consumer faima be- cause they are less vulnerable than specialists to fluctuations in the supply of particular re- sources (Pianka 1966, 1970, Levin 1968, Cody 1974, Moldenke 1975). Though this hy- pothesis is intuitively attractive, existent evi- dence is equivocal (Futuvma 1976, Rabenold 1978). In addition, the resource predictability hy- pothesis has been extended to explain latitu- dinal diversity gradients by proposing that high tropical diversity is the result of clo.se species packing of specialists in a predictable environment (Pianka 1966, 1970). However, the assertion of large differences in predict- ability between temperate and tropical re- gions has recentlv been challenged bv Wolda (1978), who has shown that precipitation pat- terns appear no less variable in the tropics and that insect populations in the seasonal tropics fluctuate as much as their temperate counterparts. The impression that high .spatiotemporal variability in resources is more common in certain regions is primarily intuitive or rests on anecdotal evidence; convincing documen- tation is lacking (Ricklefs 1973). Clearly, if resource predictability and .spatiotemporal heterogeneity are to play more than a hypo- thetical role in ecology, more quantitative field studies are needed that measure changes in resource levels and their use across space and time. Department of Zoology and Physiolog)'. University Station Box 3166, University of Wyoming, Laramie, Wyoming 82071. Present address: USD.VSE.\-Ali-\VR, Bee Biology- and Systematics Laboratory, Utah Stale University, UMC 53, Logan, Utah 84322. 197 198 Great Basin Naturalist Vol. 40, No. 3 For many consumer guilds, it is difficult to distinguish between what is and what is not a resource (Haigh and Maynard Smith 1972). In contrast, flowers, the trophic resources of bees and other animals, are easy to delimit and quantify. In this report we present two years' data on variation in phenology and abimdance of entomophilous flowers on two shortgrass prairie sites in SE Wyoming. Fu- ture papers will relate these data to the struc- ture of the bee community (Tepedino and Stanton, in preparation). The Great Plains are subject to the wide and unpredictable fluctuations in temper- ature and moisture availability which typify interior, temperate climates. Shortgrass prairie experiences the most unpredictable fluctuations in precipitation of all North American grassland biomes (Wiens 1974). For example, two of every five years may be expected to deviate by a minimum of 25 per- cent from mean yearly precipitation and one of 12 deviates by at least 50 percent; this var- iation is temporally impredictable (Wiens 1974). In addition, precipitation frequently occurs in localized patches and is spatially unpredictable (Coupland 1958). Low climatic predictability prompted the following hypotheses for flower production: 1. Floral phenology and abundance exhibit wide year-to-year variations at a given site, both at the level of the species and, more generally, for the whole commu- nity. 2. If climatic variation is spatially local- ized, then within-year differences be- tween sites in floral phenology and abundance should be evident. 3. Between-year variability is modified by seasonal effects. In particular, the spring flora exhibits greater year-to- year variability than the summer flora (Leopold and Jones 1947). 4. Within-year predictability, i.e., the probability of encountering similar flor- al composition for several consecutive weeks, is greatest during the summer blooming season. Methods and Study Sites Study Sites The study was conducted in the southern part of the Laramie Basin, a semiarid inter- mountain valley in Albany County, Wyom- ing. In general, soils are shallow, rocky, and poorly developed. Precipitation in nearby Laramie averages 25.6 cm per year, with 70 percent falling from April through Septem- ber. The growing season is short, varying be- tween 80 to 100 days, with killing frosts com- mon in June and early September. With few exceptions, the flora is composed of perennial species. The Dirt Farm Located approximately 16 km southeast of Laramie, The Dirt Farm site is 1.6 ha in area at an altitude of 2250 m. The vegetation is divided into three contiguous communities. A cushion plant commimity covers 0.77 ha and is located on windswept shallow soils with frequent bedrock exposure. Abundant species include Phlox bryoides. Astragalus serico- leucus, A. spatidatiis, Arenaria Jiookeri, and Paronychia sessiliflora, all of the caespitose, herbaceous growth form. Adjoining the cush- ion plant community is a level area of 0.33 ha dominated by the shrub, Cercocarpus montanus, mountain mahogany. Associated species include Allium textile, A. cernuum, and the half-shrub Chrysothamnus vis- cidiflorus. Soils are very shallow or nonex- istent here also, with the roots of the shrubs penetrating cracks in the bedrock. Extending south from the mountain mahogany commu- nity is a 0.5 ha section of typical shortgrass prairie. Soils are deeper here than in the other two communities. Boulder Ridge The Boulder Ridge site covers 1 ha and is located approximately 38 km southwest of Laramie (22 km southwest of The Dirt Farm) at an altitude of 2425 m. The vegetation is foothill scrub (Porter 1962) and is divided into three communities. A central section of 0.35 ha is dominated by the shrub Cerco- carpus montanus with a few individuals of the shrubs Primus virginiana, Amelanchier al- nifolia, and Ribes cereum. Abundant associ- ated herbs are Allium textile, A. geyeri, A. cernuum, Ranunctdus ranuncuUnus, and Ce- rastium arvense. The western part of the communitv is level, but the eastern end September 1980 Tepedino, Stanton: Bee Ecology 199 slopes at an angle of approximately 30 de- grees. At slope bottom movmtain mahogany gives Way to a community of 0.28 ha domi- nated by sagebrush {Arteinisia tridentato). The most abimdant associates are Castilleja flava, Astmoalus flexuosiis, CoUinsia parii- folia, Orthoccirpus luteus, and Chrysopsis vil- losa. Bordering the study site at its eastern and western ends is a heterogeneous commu- nity of 0.37 ha that includes representatives of the cushion plant, shortgrass prairie, and sagebrush communities. Nomenclature is that of Harrington (1954), Porter (1965), and We- ber (1967). Methods Censusing.— Floral phenology and abun- dance of species with entomophilous flowers were estimated for two years at each site by weekly censuses of the number of flowers by species in permanent m^ quadrats. Censusing extended from the last week in May through the last week of August, except for the Boul- der Ridge site in 1975, which began one week later because of a spring snow storm. Quadrats were chosen by using a stratified random sampling technique to insure that each community was sampled in proportion to its percentage of the entire study area. Ap- proximately 1.6 percent of the total area of each site was sampled. Individual flowers were counted in all cases except for most Compositae, where heads were counted, and the Umbelliferae and Polygonaceae, where umbels were counted. Analysis.— To avoid the assumptions of normality and homoscedasticity, non- parametric statistics were used. The Sign Test (Conover 1971) was used to test for be- tween-year differences in total floral abun- dance at each site by comparing the total number of flowers in each permanent quad- rat for each set of paired sampling dates (Table 1). Comparisons were made for each site as a whole and by vegetation tvpe. To provide a measure of the similarity be- tween sampling dates both within and be- tween years for each site we used the Czeka- nowski measure (also known as the Brav- Curtis Index: Goodall 1973) to generate sim- ilarity matrices, which were then subjected to cluster analysis. The Czekanowski Index is written PS = 22 min (x.y.)/ 2 (x. + y.), 1=1 i= 1 where Xi and y^ are the number of flowers of species i on dates x and y. The matrices were analyzed by single, complete, and average linkage clustering al- gorithms using the BMDP computer package (Dixon 1975) and the best grouping method, decided by calculating the cophenetic corre- lation coefficient of Sokal and Rohlf (Sneath and Sokal 1973). The average linkage method provided the best results, and only these are reported. Matrices for each site for all census date comparisons as well as separate within- site, within-year matrices were clustered. Only the four within-site, within-year clusters are reported here because we consider them most informative. Table 1. Actual floral census dates and their corresponding census code numbers for each study site. Census Dirt Farm Boulder Ridge Code 1975 1976 197,5 1976 1 26 May 24 Mav No census 27 May 2 2 June 31 May 5 June 3 June 3 9 June 7 June 13 June 10 June 4 16 June 16 June 20 Jiuie 18 June 5 23 June 21 June 27 June 24 June 6 30 June 28 June 4 July 1 July 7 7 July 6 Julv 11 July 7 Julv 8 14 Julv 12 Julv 18 July 18 Julv 9 21 Julv 21 Julv 25 July 25 Julv 10 28 July 28 July 31 July 31 July 11 6 August 5 August 7 August 8 .\ugiist 12 13 August 11 August 14 August 14 August 13 19 August 18 August 23 August 19 August 14 26 August 28 August 29 August 28 August 200 Great Basin Naturalist Vol. 40, No. 3 Results Precipitation and temperature.— Precipi- tation patterns over the two years of study were quite different (Data from weather sta- tion at Laramie Airport). Rainfall in spring and early summer 1975 was much heavier than normal (Fig. la), and total precipitation for the year was 6.1 percent above normal. Conversely, 1976 was a dry year with below normal rainfall for every month from March through June. By the end of June precipi- tation was 32.0 percent below normal. July and August received greater than average rainfall, and by the end of the study precipi- tation was only 17.0 percent below normal for the January through August 1976 period. In general, temperatures were warmer in 1976 (Fig. lb). In particular 1976 was warm- er from April through July, a period which was also (July excluded) much drier than nor- mal (Fig. la). The frost-free period in 1975 extended from 16 June to 5 September (79 days) and in 1976 from 25 June to termi- nation of the study (29 August) (63 days). JFMAMJJASOND 6- I a E,. u c « o"- , /\ — ,\ 1 x^\ .«- 0] / ^' , J /"'/ ''\ **"* 3' \ Q. 1/ K^ U y 1 ^'v f / a;2 A. / / ' \ / I /\ / T ■ -i 1 "7 /i \j ~^v'V> y^ / 1975 MAM 1976 Fig. 1. Monthly precipitation (a) and temperature (b) records from the Laramie .\irport (Brees Field); a) solid line = 1975, 1976, dashed line = normal; b) solid line = 1975, dashed line = 1976. Species Composition.— We recorded 63 and 73 entomophilous plant species during the two years at The Dirt Farm and Boulder Ridge, respectively (Appendix A). Total flow- ers by species and year are also shown in Ap- pendix A. The family Compositae was repre- sented by the largest number of species at both sites, followed by the Cruciferae at The Dirt Farm and the Scrophulariaceae at Boul- der Ridge. Dirt Farm Phenology.— 'The flowering phenology of selected species is shown in Figure 2. Phe- nological variation between years took two forms: differences in first bloom and in bloom span. Evidence for variability in first bloom comes from several sources. First, when total floral abundance is graphed by date for each year, it is clear that both June and August peaks were advanced in 1976 (Fig. 3). It is worthwhile to note, however, that the ad- vancement in each peak is not due to similar responses in each vegetation type. The early peak is advanced due to responses of the cushion plant and mountain mahogany com- munities, and the shortgrass and mountain mahogany communities account for advances in the late peak (Fig. 3). Earlier flowering in 1976 was due to a warmer spring and sum- mer relative to 1975. Phenological advancement in 1976 is also seen when first bloom dates are compared by species. Analysis shows that 27 of 38 species differed by at least one week in anthesis. Eleven species were excluded because they were in bloom when censusing began, and 14 others were eliminated because they flower- ed in only one year. Of the 27 species that differed in phenology, 24 were earlier by an average of 10 days in 1976 (X2 = 16.33, P<0.001). Are there seasonal differences between early- and late-blooming species in phenolo- gical predictability? lo test this the census period was halved and species grouped ac- cording to the half in which they began blooming (Fig. 3). Twelve of 18 first-half spe- cies showed a mean difference of one week in beginning bloom, and 15 of 20 second-half species showed mean advance of 12.1 days. Species blooming during the last half of the September 1980 Tepedino, Stanton: Bee Ecology 201 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 CO 20 021 U22 d) 23 CL24 00 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 Census week ^456789 10 11 12 13 14 -J I Fig. 2. Bloom spans of selected species at The Dirt Farm for 1975 (solid) and 1976 (dashed); stars = did not flow- . Clensiis date numbers refer to Table 1. species numbers to Appendix A. 202 Great Basin Naturalist Vol. 40, No. 3 MOUNTAIN MAHOGANY SHORTGRASS 200r 120 - en UJ Q tr lOOr UJ CD :^ 3 I 2 3 4 5 6 7 8 9 10 II 12 13 14 JUNE h- JULY -\ AUG 2 3 4 5 6 7 8 9 10 II 12 1314 JUNE — +- JULY H — AUG CENSUS WEEK Fig. 3. Total floral abundance per ni^, irrespective of species, at The Dirt Farm for each census date in 1975 and 1976. Census date numbers refer to Table 1. Asterisks mark significant differences (P<.05) between years for paired dates. September 1980 Tepedino, Stanton: Bee Ecology 203 season showed significantly greater phenolo- gical differences than those blooming during the first half (Mann-Whitney U-Test, P<0.05). Between-year comparisons of bloom spans are a second indicator of phenological varia- bility. Differences of at least one week in bloom span were shown by 22 of 38 species. Average difference in bloom span for the 22 species was two weeks with a range of one to seven weeks. Ten species had longer spans in 1975 and 12 had longer spans in 1976. A comparison by seasonal grouping of bloom span data into early and later blooming spe- cies shows that longer bloom spans during the last half of the season occurred mostly in 1976, and 1975 had more longer blooming species during the first half (X^ = 2.76, P = 0.097). Floral Abundance.— Differences in floral abundance between years is first shown by DIRT FARM comparing total number of flowers by date (Fig. 3). Total abundance comparisons show the early peak to be higher but the late peak lower in 1975. The late peak difference is due largely to profuse flowering of Eriogo- nitm effusum. Exclusion of this species re- sults in much closer agreement of abundances from late July to mid-August. When total floral abundance is partitioned into component communities it is again evi- dent that there is no typical, overall site re- sponse (Fig. 3). For example, the mountain mahogany community shows nine significant between-year differences in abundance, with 1976 having more flowers on eight dates. In contrast, between-year differences in the cushion plant community show 1975 with more flowers for seven or nine significant comparisons and the shortgrass community with more flowers in 1975 for five of six dates. A cold period during the week begin- BOULDER RIDGE 60r 50 - 40 - iij o 30h UJ Q. 20 ■ 10 - z < X a: < X o o llJ r 1.00- 2.00- >3.00 1.99 2.99 < X o o UJ (A) F < X CO T 1.00- 2.00- >l3.00 1.99 2.99 RHL Fig. 4. Frequency distril)iition of the ratio of the number of flowers in the most abundant year to the number of flowers in the least abundant year for each species (RHL) at both sites. 204 Great Basin Naturalist Vol. 40, No. 3 ning 16 Jiine 1976 reduced floral production appreciably on the mountain mahogany com- munity and probably affected the shortgrass community as well. During this period tem- peratures were below freezing for three nights and snow and sleet fell twice. The ef- fect of such periods upon organisms in the Rocky Mountains has been described by Ehrlichetal. (1972). Floral abundance was also compared by species between years. An expression of dif- ferences in abundance is the ratio of total number of flowers observed in the year of highest production divided by total number of flowers in the year of lowest production (RHL). We eliminated from this analysis those species that either flowered in only one year or failed to produce at least 50 flowers in either vear. For the remaining 44 species mean RHL = 5.39 (SD = 8.56, range = 1.04-48.60), suggesting that the abundance of flowers of most species may show significant bet ween-y ear- variation. The data are shown as histograms in Figure 4. Of the 44 species, 26 were more abundant in 1975 and 18 were more abundant in 1976. Thus, no year effect was observed. Do RHL values display a seasonal pattern? Again, species were grouped according to the half of the season in which they began flow- ering and the Mann- Whitney U-Test used to test for differences in RHL between groups. The comparisons show that plants blooming during the latter half of the season were sig- nificantly more variable (P = 0.05), i.e., had larger RHL values than those blooming ear- lier. No year effect was evident since both years had an almost equal proportion of spe- cies with highest RHL values in each half of the year. Similarity and Cluster Analysis.— We used cluster analysis to elucidate differences be- tween census dates within and between years and to illustrate seasonal groupings. High similarity values for paired between-year sampling dates were expected; however, the data do not support this hypothesis. Mean similarity for paired census dates was only 0.519 (SD = 0.196, range 0.204-0.854). Mean similarity was highest (0.593 [SD = DIRT FARM BOULDER RIDGE 0.8 r 23456789 10 II I JUNE 1— JULY H 0.8 r 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 / / /|976 \ J I I L J I I I I 23456789 10 JUNE \— JULY — I CENSUS WEEK Fig. 5. Czekanowski similarity measures between floral ahiindance for each census date and the third subsequent week at both sites. Census date numbers refer to Table I. September 1980 Tepedino, Stanton: Bee Ecology 205 0.149, range = 0.265-0.772]) when each 1975 date was compared with the week prior to that date in 1976. Thus, the similarity data provide additional evidence for phenological advancement in 1976. Within-year similarity comparisons were also quite variable. We reasoned that, since bees are the predominant pollinators on shortgrass prairie and because females of most species of solitary bees fly for a mini- mum of four weeks (Linsley 1958), a conserv- ative estimate of within-year resource varia- bility for a bee would be the similarity between the week of emergence and three weeks later. This measure is conservative be- cause we used similarity measures between dates that are one week less than the mini- mum flight span. In Figure 5 we graph the results for each year. Except for the last month of the census period, values are very low. Any species emerging during the first two-thirds of the blooming season would face a very different flora during the latter part of its flight season relative to that encountered initially. Cluster analysis aids in depicting seasonal groupings and transitional periods within the blooming season (Fig. 6). The number of clus- ters formed at a value of .50 differs between years as does the number of unclustered dates, suggesting that the "structure" of the blooming season may differ from year to year. For example, five clusters plus one un- clustered date form in 1975, but four clusters and four unclustered dates form in 1976. Within both years late season dates cluster strongly, again indicating higher within-year predictability for late summer bees. Almost all other clusters are composed of only two consecutive census dates, indicating a high rate of turnover in floral composition for the first two-thirds of the blooming sason. Boulder Ridge Phenohgij.— Flowering also began earlier in 1976 at Boulder Ridge (Fig. 7).^ Of the 73 species censused, 33 were either in bloom when censusing began of flowered in only one year and were eliminated from this anal- ysis. Of the remaining 40 species, 32 showed phenological differences of at least one week. Twenty-six of the 32 species were earlier an average of 10 days in 1976 (X^ = 12.50, P< 0.001). The seasonal differences in ad- vancement between first- and second-half DIRT FARM 0.0 r 0. 0.2 0.3 > 0.4 q: < 0.5 1 '^ 0.6 0.7 0.8 0.9 1.0 1975 1976 a I 2 3 4 5 6 7 8 9 10 II 12 13 14 I JUNE 1— JULY —I AUG I 2 3 4 5 6 7 8 9 10 II 12 13 14 I JUNE (— JULY — I AUG CENSUS WEEK Fig. 6. Dendrograms of floral similarity between census dates for each year at The Dirt Farm. Census date niim- hers refer to Table 1 . 206 Great Basin Naturalist Vol. 40, No. 3 CO a; u (D Cl 00 Census week 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 3 I 4 5 6 7 I 8 9 I 10 11 _J 1_ 12 13 14 _i I I Fig. 7. Bloom spans of selected species at Boulder Ridge for 1975 (solid) and 1976 (dashed); stars = did not flower. Census date numbers refer to Table 1, species numbers to Appendix A. species observed at The Dirt Farm were not detected here. Most species differed in bloom span be- tween years. After eliminating species that began blooming prior to censusing, 29 of the remaining 40 differed by at least one week. Average difference in bloom span for the 29 species was 17 days with a range of one to five weeks. Chi-square analyses for sea.sonal patterns were insignificant. Year effects, however, were present; longer bloom spans were concentrated in 1975 (20 of 29, X2 = 4.17,F<0.05). Abundance.— Differences in total floral abundance between years were remarkable (Fig. 8). Twelve of the 13 comparisons September 1980 Tepedino, Stanton: Bee Ecology 207 showeci significantly greater abundance in 1975. There was a marked midyear peak in 1975 that is only suggested in 1976. In contrast to The Dirt Farm, floral abun- dance for all vegetation types at Boulder Ridge was higher in 1975 (Fig. 8). Between- year differences on the heterogeneous area were primarily responsible for the total flow- er differences. The last 10 dates showed sig- nificantly more flowers on this section in 1975 (Fig. 8). The second peak in the total abundance curve in 1975 (Fig. 8), absent in 1976, was due mainly to the heterogeneous area. Differences observed on the other sec- tions showed floral abundances in 1976 to be consistently below those of 1975, although many of these comparisons were not signifi- cant. The RHL ratio was used to compare abun- dance by species between years. We elimi- nated all species with less than 50 flowers in either years and those that flowered in only one year. For the remaining 38 species, mean RHL = 5.47 (SD = 6.56, range = BOULDER RIDGE c/) q: UJ o UJ CD 60 50 40 30 20 10 I 2 3 4 5 6 7 8 9 10 II 12 13 14 28i- TOTAL 24 h ' HETERO- GENEOUS r- ,1975 " A i 1 v^ - >y V V * y \ __^ \ \*y / ^ ^ jy ^ \ y ^ \ 1976 \ 1111 -L — L 1 1 1 -L. 8 - 4 - \^\ .1976 -- \ MOUNTAIN MAHOGANY J I L J. J_J l_l L-J I 20 28 24 20 16 12 8 4 2 3 4 5 6 7 8 9 10 II 12 13 14 SAGEBRUSH 2 3 4 5 6 7 8 9 10 II 12 13 14 I 2 3 4 5 6 7 8 9 10 II 12 13 14 |— JUNE I JULY— I AUG |— JUNE — | JULY 1 — AUG CENSUS WEEK Fig. 8. Total abundance per m^, irrespective of species, at Boulder Ridge for each census date in 1975 and 1976. Census date numbers refer to Table 1. Asterisks mark significant differences (P<.05 between years for paired dates. 208 Great Basin Naturalist Vol. 40, No. 3 1.07-27.50; Fig. 4). Again, there were large between-year differences in abundance for most species on the site. We tested for seasonal differences in RHL values using the same methods as for The Dirt Farm. Again, RHL values for the latter part of the blooming season are significantly higher (P<0.05). However, in contrast to The Dirt Farm, each year is not equally rep- resented by species with high RHL values. Twenty-eight of the 38 species were more abundant in 1975 (X^ = 8.53, P< 0.005). This phenomenon is particularly striking for the latter part of the blooming season, when only one of 16 species was more abundant in 1976. Similarity and Cluster Analysis.— As with The Dirt Farm data, the expected high sim- ilarity for paired between-year comparisons was not evident. Mean similarity for paired census dates was only 0.424 (SD = 0.200, range = 0.184-0.896). Again, highest mean similarity was between 1975 dates and the week prior to the identical date in 1976 (x = 0.518, SD = 0.161, range = 0.309-0.869). Within-year similarities [between each census date and the census taken three weeks later] were as low as at The Dirt Farm (Fig. 5). The graph for 1975 is uniformly low with- out the appreciable rise la^te in the season present in The Dirt Farm and Boulder Ridge 1976 data. Cluster analysis again suggests "structural" differences between the blooming seasons (Fig. 9). Four clusters plus three unclustered dates form in 1975, and five clusters and three unaffiliated dates are found in 1976. As at The Dirt Farm, late season dates cluster most densely and all but one other cluster is composed of only two consecutive dates at the .50 level. Discussion Plant species varied substantially between years in the onset and length of the blooming period and in the number of flowers pro- duced. Comparison of phenology and abun- dance, both between sites and among vegeta- tion types within sites, shows other important differences. Though phenological advance- ment in 1976 was a uniform occurrence at both sites, changes in the direction and mag- nitude of floral abundance were not. A com- parison of total floral abundance between The Dirt Farm and Boulder Ridge shows that the two sites behaved quite differently. Floral production at Boulder Ridge in 1976 was consistently well below that of 1975. In con- 0.0 - 0.1 - 0.2 - 0.3 I- >- 0.4 o: 2 0.5 h ^ 0.6 0.7 0.8 0.9 1.0 1975 BOULDER RIDGE 1976 Q 2 3 4 5 6 7 8 9 10 II 12 13 14 I— JUNE —I JULY \ — AUG CENSUS WEEK I 2 3 4 5 6 7 8 9 10 II 12 13 14 I — JUNE —\ JULY 1 AUG Fij;. 9. Dciidrograiii.s of floral similarity between census dates for each year at Boulder Ridge. Census date num- bers refer to Table 1 . I September 1980 Tepedino, Stanton: Bee Ecology 209 trast, floral production was significantly high- er at The Dirt Farm in 1976 over the latter part of the blooming season. Evidently, the spring and early summer drought was either more severe at Boulder Ridge or the plant communities at Boulder Ridge were more susceptible than those at The Dirt Farm. Between-site differences appear due to varying responses to weather by each vegeta- tion type within each site. At Boulder Ridge all commmiities displayed consistently lower floral productivity in 1976, but at The Dirt Farm each community responded indepen- dently. Indeed, at The Dirt Farm each spe- cies seemed to display an independent re- sponse as shown by the lack of any year effects on the distribution of RHL values ei- ther in between-year comparisons or be- tween-season comparisons. These observa- tions suggest that variation in floral production is expressed as spatially localized patches of high or low abundance that change from year to year. Because we collected data for only two years, it is necessary to ask how representa- tive of routine variability these results are. Schemske et al. (1978), in a study of seven spring herbs, found the onset of flowering to range, by species, from 8 to 22 days over three years. More importantly, peak flower- ing did not usually coincide with optimal pollinator conditions. Long-term bloom re- cords for several regions in North America are available for analysis. In several cases data are available for period of up to 30 years in the same area (Lindsey and Newman 1956-Indiana; Smith 1915-Ohio; Hulbert 1963-Kansas; Hodson 1971-Minnesota). In examining these data we have used varia- bility in first flowering as an indication of re- source predictability since this phenophase is common to all studies. In brief, we find that almost all variability in the date of first bloom is accounted for with 10 years of ob- servations and that the range of first bloom is between four and five weeks for most species (Tepedino and Stanton, impublished manu- script). Other support for phenological varia- bility exists. Recently West and Gasto (1978) reported that the onset of bloom of two arid land shrubs in northwestern Utah varied over seven years by 44 and 39 days. Thus, the sub- stantial phenological variability recorded in our study over two years is low relative to what can be expected over a 10-year period. Long-term studies of floral abundance are few. Tamm (1948, 1956, 1972a, b) counted flowers of several species in permanent quad- rats in forest and meadow in mid-Sweden for 14 to 29 years. All species showed large ir- regularities in year-to-year floral abundances from no flowers in some years to profuse abundance in others. Short-term studies are more numerous. Ackerman and Bamberg (1974) reported large variation in floral abundance over a three-year period for Lijciinn andersonii in Nevada. Bykov (1974), in a general review of vegetation dynamics of the arid Turanian Plain, reported wide variation in floral abun- dance of both ephemerals and perennials. Sa- rukhan (1974) supplied floral abundance data for three species of Ranunculus for two years from permanent plots, with all species pro- ducing many more flowers in the first year. Holway and Ward (1965), studying the alpine plant community in northern Colorado over two years, noted that floral production in the second year was much lower. Davies (1976) used the same five trees of each of two spe- cies and recorded the number of individuals flowering over an eight-year period in west- ern Australia. Combining data for both spe- cies {Acacia pruinocarpa, Hakea lorea; Davies 1976; Table 6) showed that in three of eight years the number of individuals flowering was 40 percent or less. Data on fruit crops also were presented for 10 species of shrubs and trees for 10 years. If we can assume that fruit crop bears at least a partial relation to floral production (Grubb 1977), floral pro- duction was irregular in 9 of the 10 species. Schemske (1977, 1978) has shown that the number of flowers of two woodland herbs censused in 78 permanent m^ quadrats varied considerably between years. Moldenke (1976) noted that floral production varies widely be- tween years in California grasslands. Treshow (1979), in a six-year study of the pinyon- juniper community in Utah, has shown that forb cover in almost every year differed sig- nificantly from each other year. Year-to-year variation in floral abundance is not restricted to "unpredictable" temper- ate zone communities (Federov 1966). Mass flowering via synchronization of all members 210 Great Basin Naturalist Vol. 40, No. 3 of a particular species or many species in a community in the tropics at periodic inter- vals is well known (Whitmore 1975). Med- way (1972) and McClure (1966) provided data showing widespread irregularity in flow- ering for 46 species of tropical rain forest trees in Malaya. Although most of the obser- vations were recorded on very few individ- uals, it is enlightening to learn that the per- cent of species flowering each year ranged from 44 to 88 over the period from 1963 to 1968, with an average of 58 percent. Of 42 species observed for the entire six-year peri- od, only 11 (26.2 percent) flowered every year, and 24 (57.1 percent) flowered in three or fewer years. In a study of flowering phenology in Ceylon, Koelmeyer (1959) re- ported: "There was no regularity in the se- quence of years of flowering and years in which there was no flowering in the individ- ual trees. The result is the absence of a defi- nite cycle of flowering." The data seem clear. Where data on year- to-year floral abundances have been record- ed, large variations in floral production are the rule rather than the exception. Variability in floral resources may also be modified by seasonal influences. First, some parts of the blooming season may exhibit more year-to-year variability than others. Leopold and Jones (1947) hypothesized that early blooming species are more "turbulent" in first bloom than those which bloom later in the year. We reexamined the phenology data of Leopold and Jones (1947) for Wiscon- sin using multiple regression analysis and found that their Sauk County data do show a significant inverse correlation between range of flrst bloom and average first bloom date. Though the Dane County data show the same pattern, it is not significant (Tepedino and Stanton, unpublished manuscript). The Wyoming data do not support the hy- pothesis of greater year-to-year "turbulence" in the spring flora. At The Dirt Farm, be- tween-year differences in flrst bloom were signiflcantly greater for plants that bloomed over the second half of the census period. At Boulder Ridge no significant difference be- tween flrst- and second-half plant species was detectable. In addition, between-year differ- ences in abundance (as judged by RHL ratios) were significantly greater for the last half of the blooming season at both sites. Whether these differences were due to only two years' data from Wyoming or to conditions that are site specific is not clear. The second way in which floral resources may vary seasonally is in predictability of subsequent resource abundance and composi- tion. From this perspective the spring flora is, indeed, more turbulent; predictability, as judged by floral similarity values calculated at three-week intervals (Fig. 5), was low rela- tive to average flight time for bees until the latter third of the bloom season when com- posites became dominant. This result may be somewhat misleading however, because more species begin bloom in spring than in late summer and low spring similarity values are due in large part to species additions. Low year-to-year predictability in floral phenology and abundance must exert strong selection on flower-visiting insects. This is particularly true for bees because every stage in their life cycle is obligately dependent upon floral resources for food. When re- sources are unpredictable in time and/or space, selection should favor generalized con sumers. Alternatively, specialization would require precise synchronization between bee emergence and anthesis of the host plant, particularly when the host has a brief bloom span. It is unclear how such precise synchro- nization might be affected. In most plant spe- cies studied photoperiodic stimuli initiate flower formation, but subsequent devel- opment and anthesis is profoundly modified by diverse factors such as moisture and nutri- ent availability and temperature (Evans 1969, Schwabe 1972). Our knowledge of the stimuli used by bees to cue emergence in a natural setting is scanty (Linsley 1958), but in the laboratory temperature alone is a reliable stimulus for several species {Megochile rotiin- data (Fabricius), Osmia lignaria Say, Hylaetis bisintiatiis Forster, Nomia melanderi Cock- erell, and several others; G. E. Bohart, F. D. Parker, P. F. Torchio, pers. comm., pers. obs.). Thus, though anthesis is determined by a complex of factors, bee emergence may be primarily responsive to temperature. Because of these differences in potential stimuli used by the two groups, synchronization may be rare. In this regard, Linsley (1958) noted that September 1980 Tepedino, Stanton: Bee Ecology 211 studies of oligolectic bees frequently reveal consistent with documented fluctuations in poor synchronization. Even if specialized bees could achieve close synchronization with host plant an- thesis, the problem of year-to-year variation in resource quantity still remains. Attempting to track specific floral resources that vary widely could cause large fluctuations in the populations of bee specialists, thereby in- creasing the probability of local extinction (Tepedino 1979). The frequently expressed view that most temperate bees are special- ized (van der Pijl 1966, Faegri and van der Pijl 1971, Heinrich 1976, Raw 1976, Heithaus 1979) will probably require modifi- cation because such specialization seems in- floral resources. Acknowledgments We thank J. M. Loar and T. M. Root for providing meticulous assistance in the field, J. Meyer for drawing the figures, and Dr. L. L. McDonald for advice on cluster analysis. The manuscript was improved by comments by M. S. Boyce, K. T. Harper, D. H. Knight, P. Lincoln, M. D. Marcus, and A. R. Moldenke. The study could not have been conducted without support from NSF Grant BMS75- 14044. "Doc" and Peggy Wollbrinck of Lar- amie and the proprietors of The Dirt Farm, Inc., graciously allowed use of private land. Appendix A. The number of flowers recorded in permanent m^ quadrats at The Dirt Farm and Boulder Ridge in 197.5 and 1976. Nomenclature: Harrington (1954), Weber (1967), Porter (1965). Numbers in the far left column refer to Figures 2, 7; the first number to Figure 2 (Dirt Farm), the second to Figure 7 (Boulder Ridge). A zero signifies nonrepresentation. Species Dirt Farm 1975 1976 Boulder Ridge 1975 1976 Berberidaceae (0,8) Berberis repens Lindl. Boraginaceae (19, 0) Crijptantha flovoculata (A. Nels (23, 0) Cijnoglossitm officinalis L. (14, 0) Hackelia florihwida (Lehm.) Lappulo redoicskii (Horneni.) (10, 0) Lithospermtim incisum Lehm. (0,5) Mertetisia Iwinilis Rydb. (38, 0) (0, 23) (24, 19) (0,3) (22, 20) (40, 34) (35, 28) (39, 0) (42, 0) (0, 24) Cactaceae Opiintia pohjacantha Haw. Capparidaceae Cleorne serrtilata Pursh Caryophyllaceae Arenoria fendlcri A. Gray Arcnaria hookcri Nutt. Cerastitim arvense L. Paronychia sessiliflora Nutt. Stellaria media (L.) Compositae Acliillea miUefoliuin L. Antennaria microphyUa Rydb. Antennaria rosea Greene Artemisia frigida Willd. Aster rtibrotincttis Blake Chnjsopsis villosa (Pursh) Chrysothamntts nauseosus (Pallas) Chrysothamntis viscidiflorus (Hook.) Cirsitim tmdulatum (Nutt.) Erigeron canus A. Gray 101 52 68 575 60 - 32 50 47 — — 67 109 — — _ — 5 2 133 16 9 9 12 5 16 42 _ _ 122 58 1165 628 1577 474 _ — .332 50 3489 .3000 718 40 — — 9 — 382 19 _ .3.343 959 _ 83 - 311 114 .393 - ,393 73 87 18 262 116 2426 876 1882 73 - - 1906 4216 11 3 17 4 2 — - 2 817 314 212 Appendix A continued. Great Basin Naturalist Vol. 40, No. 3 (12, 2) (41, 35) (34, 29) (30, 14) (6,0) (13, 9) (0,30) (0, 18) (32, 0) Species Erigeron nematophyUus Rydb. Erigeron pumilus Nutt. GailUirdia aristaia Pursh Gutierrezia sarothrae (Pursh) Haplopappus nutiaUii T. & G. HelkintheUa uniflom (Nutt.) Hijmenoxijs acaiiUs (Pursh) Hymenoxijs torreijana (Nutt.) Senecio catum Hook. Senecio integerrimus Nutt. Solidago spathulata DC Taraxicttm sp. Hall Townsendid scricea Hook. Crassulaceae S«/i("i stenopetalum Pursh Cniciferae (4,0) Arabis fendleri (Wats.) Arcihis holbocUn Hornem. (21, 0) Dcsnirainia sophia L. Dmha nemorosa L. (15, 11) b'.rijsimum capitatum (Dougl.) Halimolohos virgata (Nutt.) (8, 10) LcsqucrcUa ludoviciana (Nutt (0,7) Phiisaria amtralis (Payson) (17, 0) Smjmbrium ultissimum L. Sunjmhrium linefolium Nutt. Euphorbiaceae Euphorbia sp. L. Gentianaceae Swertia radiata (Kellogg) Labiatae Hedcoma drtDtitiiondii Benth. Snitelhirid bhtionii Porter Legiuuinosae (25, 0) Astragcdm hisulaitus Hook. Astragalus crassicarpus Nutt. (0, 21) Astragalus flexuosus Dougl. (9, 0) Astragalus sericolettcus Gray Astragalus shortianus Nutt. (7, 0) Astragalus spatulatiis Sheld. (0, .32) Astragalus striatus Nutt. (0,12) TItcrmopsis rhoinhifolia Nutt. Liliaceae (37, 31) Allium cernuum Roth (0, 16) Ellium getjeri Wats. (18, 17) .A//ii/m textile Nels. & Macbr. Caloehorttis nuttallii Torrev (0, 1) Leticocrintim montanum Nutt. (11, 0) VAjgadenus venenosus Wats. Linaceae Lidi/Hi lewisii Pursh Dirt Farm Boulder Ridge 1975 1976 1975 1976 434 406 895 1014 59 — 12 - _ 7 - 2163 3170 1512 735 45 — — 45 35 50 3 62 81 82 24 342 248 — — 499 701 21 53 10 11 — 19 227 100 _ — 57 — 4 _ 2 — 279 971 125 9 325 75 9 88 1768 366 535 89 166 — — — 13 17 128 225 — - 35 581 207 — 122 - 160 12 — - - 1839 332 466 360 48 — 219 84 24 52 - - 27 — — — 187 _ — 4.38 31 21 57 - - _ — 18 - 386 329 - - _ 41 — - - 84 43 1771 2693 478 98 _ _ 853 607 622 1284 187 200 1 — — — — 73 122 211 314 - - 25 17 September 1980 Appendix A continued. Tepedino, Stanton: Bee Ecology 213 Species Malvaceae SpluicKiIcea coccinea (Pursh) Onagraceae (31, 0) Gaum coccinea Nutt. Oenothera cownopifolia T. & G. Poleinoniaceae Gilia apicata Nutt. Mirrostcris htivtilis (Dougl.) (1,0) Plilox hnjoides Nutt. (2,6) Phlox hoodii Rich. Polygonaceae Eriogonum ahitttm Torr. (36, 0) Eriogonum effiistini Nutt. (0, 33) Eriogonum janicsii Benth. (28, 22) Eriogonum umheUatum Torr. Portulacaceae Clatjtonia hinceohita Pursh Primulaceae Androsace septentrionalis L. Ranunculaceae (26, 0) Delphinium nehoni Greene (0. 4) Ranunctihts ranunetdinus (Nutt.) Rosaceae (0, 15) Amelanehier ahiifolia Nutt. (16. 13) Ccrcoearpus montonus Raf. Potcntilla coneinna Richards PotcntUla fi-ssa Nutt. (29, 0) PotentiUa liippiana Lehm. Santalaceae Commandra umbellata (L.) Saxifragaceae Ribes cereum Dougl. Scrophulariaceae (0, 25) CastUIeja flava Watson CoUinsia parviflora Dougl. Oddiocarpus hiteus Nutt. (20, 0) Penstemon angustifolius Nutt. (27, 0) Penstemon erianthcrus Pursh (33, 27) Penstemon laricifohus exHifolius (A. Neis. (0, 26) Pemtemon strictus Benth. Umbelliferae (5, 0) Harbouria trachijpleura (A. Grav) (3, 0) Violaceae Viola nuttallii Pursh Dirt Farm 1975 1976 28 317 41 67 Boulder Ridge 1975 1976 - - 4 6511 7526 — 91 42 12 192 27 85 28,869 57,115 - - - - 84 298 852 284 282 73 19 22 11 56 13 14 - - 4284 5509 _ 220 8 mi 3752 128 219 - - 9 1 - — 6 2 862 375 20 3 243 5 14 98 - - 3 7 912 .361 — — 565 402 — - 571 32.3 67 ^ - - 61 12 4 1 449 220 966 a3 - - 44 7 667 196 - 8 73 52 4 Species total Totals (both years) 56 5.5 63 57 60. 214 Great Basin Naturalist Vol. 40, No. 3 Literature Cited AcKERMAN, T. L., AND S. A. Bamberg. 1974. Phenologic- al studies in the Mojave Desert at Rock Valley (Nevada Test Site). Pages 215-226 in H. Lieth, ed.. Phenology and seasonality modeling. Springer- Verlag, New York. Bykov, B. \. 1974. Fluctuations in the semi desert and desert vegetation of the Turanian plain. Pages 243-251 in R. Knapp, ed., Vegetation dynamics. Part 8 of Handbook of vegetation science. Dr. VV. Junk, Tlie Hague. Cody, M. L. 1974. Competition and the structure of bird communities. 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Linsley, E. G. 1958. The ecology of solitary bees. Hil- gardia 27: 543-599. McClure, H. E. 1966. Flowering, fruiting and animals in the canopy of a tropical rain forest. Malay For- ester 29: 182-203. Medway, Lord. 1972. Phenology of a tropical rain forest in Malaya. Biol. J. Linn. Soc. 4: 117-146. Moldenke, a. R. 1975. Niche specialization and species diversity along a California transect. Oecologia 21: 219-242. 1976. California pollination ecology and vegeta- tion types. Phytologia 34: 305-.361. PiANKA, E. R. 1966. Latitudinal gradients in species di- versity: a review of concepts. Amer. Natur. 100: .3.3-46. 1970. On r- and k-selection. Amer. Natur. 104: .592-597. Pijl, L. van der. 1966. Pollination mechanisms in or- chids. Pages 61-75 in J. G. Hawkes, ed.. Repro- ductive biology and taxonomy of vascular plants. Pergamon, New York. Porter, C. L. 1962. A flora of Wyoming. Part 1. Univer- sity of Wyoming Agric. Exp. Sta. Bull. 402, Lar- amie, Wyoming. 1965. A flora of Wyoming. Part IV. University of Wyoming Agric. Exp. Sta. Bull. 4.34, Laramie, Wyoming. Rabenold, K. N. 1978. Foraging strategies, diversity, and seasonality in bird communities of .Appala- chian spruce-fir forc,^ts. Ecol. Monogr. 48: 397-424. RicKLEFS, R. E. 1973. Ecology. Chiron, Newton, Mas- sachusetts. Sarukiian, J. 1974. Studies on plant demography: Ra- nunctthis repens L., R. biilbosiis L. and R. acris L. II. Reproductive strategies and seed popu- lation dynamics. J. Ecol. 62: 151-178. ScHEMSKE, D. W. 1977. Flowering phenology and seed set in Clai/tonia virginica. Bull. Torrey Bot. Club 104: 254-263. September 1980 Tepedino, Stanton: Bee Ecology 215 1978. Sexual reproduction in an Illinois popu- lation of Sanguinarici ((iiuidcnsis L. Am. Midi. Nat. 100: 261-268. ScHEMSKE, D. W., M. F. WiLLsoN, M. N. Melampy, I.. J. Miller, L. Verner, K. M. Schemske, and L. B. Best. 1978. Flowering ecology of some spring woodland herbs. Ecology 59: .351-.366. Sc:nvvABE, \V. W. 1971. Physiology of vegetative repro- duction and flowering. Pages 2.33-411 in F. C. Steward, ed.. Plant physiology: a treatise. Vol- ume VI.\: Physiology of development: plants and their reproduction, .\cademic Press, New York. Smith, J. W. 1915. Phenological dates and meterological data recorded by Thomas Mikesell between 1873 and 1912 at Wauseon, Ohio. Monthly Weather Review Supplement 2: 23-93. Sne,\th, P. H. A., AND R. R. SoKAL. 1973. Numerical tax- onomy. Freeman, San Francisco, California. Tamm, C. O. 1948. Observations on reproduction and survival of some perennial herbs. Botaniser Noti- ser 1948: .305-321. 1956. Further observations on the survival and flowering of some perennial herbs. I. Oikos 7: 27.3-292^^ 1972a. Survival and flowering of some perennial herbs. II. The behavior of some orchids on per- manent plots. Oikos 23: 23-28. 1972b. Survival and flowering of some perennial herbs. III. The behavior of Primula veris on per- manent plots. Oikos 23; 159-166. Tepedino, V. J. 1979. The importance of bees and other insect pollinators in maintaining floral species composition. Great Basin Nat. Mem. 3: 139-150. Treshow, M., and J. Allan. 1979. Annual variation in the dynamics of a woodland plant communitv. Environ. Cons. 6: 231-2.36. Weber, W. A. 1967. Rocky Mountain Flora, liniversity of Colorado Press, Boulder, Colorado. West, N. E., and J. Casto. 1978. Phenology of the aer- ial portions of shadscale and winterfat in Curlew Valley, Utah. J. Range Mgmt. 31: 43-45. Whitmore, T. C. 1975. Tropical rain forests of the Far East. Clarendon Press, ().\ford. WiENS, J. \. 1974. Climatic instability and the "ecologi- cal saturation" of bird communities in North American grasslands. Condor 76: 385-400. 1976. Population responses to patchv environ- ments. .\nnu. Rev. Ecol. Svst. 7:81-120. Wolda, H. 1978. Fluctuations in abundance of tropical insects. Amer. Natur. 112: 1017-1045. DOG OWNERS AND HYDATID DISEASE IN SANPETE COUNTY, UTAH' Peter M. Schantz' and Ferron L. Andersen' .\bstract.— a questionnaire survey was conducted in Sanpete County, Utah, to determine the knowledge of dog owners concerning hydatid disease and an identification of some basic sheep management practices there. The households surveyed included 21 (Group I) that had one or more dogs infected with Echinococciis gmuiilostis tape- worms at more than one annual field clinic, and 19 others (Group II) that had one or more dogs infected when the studv first began in 1971-72, but had not had any infected dogs identified at field clinics during subsequent years. The results showed that 92.5 percent of households knew the cause of the disease and how it is transmitted, and that 9() percent knew of someone who had been operated on for surgical removal of hydatid cysts. There was no signifi- cant difference in the level of knowledge of the disease between the two groups of respondents, nor in their sheep management practices. Even though the level of infection of the parasite in dogs has decreased since the project started, certain sheep management practices persist among respondents in both groups that allow for continued transmission of the parasite in this region. Hydatid disease is an infection of people, sheep, and some other animals that produces fluid-filled (hydatid) cysts in the liver, lungs. or other organs (Fig. 1). The cysts are the lar- val (immature) forms of a tapeworm parasite, Echinococciis granulosus (Fig. 2), which lives Fig. 1. Fluid-filled hydatid cysts in the livers and lungs of infected sheep. 'Supported in part by U.S. Public Health Grant AI-10.588. 'Parasitic Diseases Division, Center for Disease Control, Atlanta. Ceorgia 303.33. 'Department of Zoology, Brigham Young University, Provo, Utah 84602. 216 September 1980 ScHANTz, Andersen: Hydatid Disease 217 ^ a. • ^ti ga ^-'Ti '^- Fisi;. 2. Tlie adult tapeworm. Echinococni.s oranitlosus (approximatelv 5-6 mm in leiiiith). removed from the small intestine ot an infeeted dog. as an adult in the .small intestine of dogs. People and sheep contract the hydatid cysts when they inadvertently ingest the tapeworm eggs passed in the stools of infected dogs. This may occur when people handle dogs that harbor the parasite, and when sheep graze on contaminated pa.stures. Dogs be- come infected with the tapeworm when they ingest hydatid cy.sts in the viscera of sheep. The parasite occurs throughout the world wherever dogs, sheep, and other suitable ani- mal hosts are kept together. The common practice among .sheep ranchers of allowing dogs to eat the uncooked viscera of home- killed sheep provides optimum conditions for continued transmission. In the United States, transmission of Ech- inococctis granulosus in the dog-sheep cycle is known to occur in several western states, including California (Araujo et al. 1975), Ari- zona and New Mexico (Schantz 1977), and Utah (Spniance et al. 1974). The most serious problem is in Utah, where nearly 50 human cases have been diagnosed since 1944. Sever- al of these cases were fatal, and most of the others have required surgical removal of the hvdatid cvsts. Many of the victims were resi- dents of Sanpete County, which is in the cen- tral part of the .state. Since 1971 hvdatid disease has been stud- ied and control measures initiated through the combined efforts of Brigham Young Uni- versity (Provo, Utah), the Utah State Depart- ments of Health and Agriculture (Salt Lake Citv, Utah), and the Center for Disease Con- trol (Atlanta, Georgia). These measures have included (1) the development and distribu- tion of educational displays and brochures on the life cycle of the hydatid tapeworm, (2) the development of adequate methods for disposal of sheep carcasses at community dumping grounds, (3) the periodic holding of public health clinics to detect new ca.ses of human infection, and (4) annual field clinics to detect new or persistent cases of infected dogs (Fig. 3). Following the implementation 218 Great Basin Naturalist Vol. 40, No. 3 Fig. 3. Sheep dogs from Sanpete County restrained at cocnts gr«nr//o.v(/.v tapeworms. of these control measures, the number of dogs found infected at the field clinics has decreased from 27 percent in 1971 (Loveless et al. 1978) to 14 percent in 1978 (unpub- lished ms.). Most sheep ranchers have shown a cooperative attitude with regard to proper disposal of sheep carcasses or viscera. Certain individuals, however, have not been success- ful in preventing reinfection of their dogs as evidenced by the fact that some of their dogs were found repeatedly infected on numerous occasions. We believed that if the reasons could be determined why some dog owners were imable or unwilling to comply with the recommended preventive measures, it might be possible to change or modify the recom- mendations to obtain more cooperation, and ultimately an improved control program. Materials and Methods A questionnaire survey was conducted of the owners of dogs that had been found to be field clinic during examination for detection of Echino- infected in Sanpete County. The survey in- cluded 40 households, 21 of which had one or more dogs found infected at more than one annual clinic (Group I) and 19 others that had one or more dogs infected only at either the first or second annual clinic (1971 or 1972), but did not have infected dogs at sub- sequent clinics (Group II). During the visits, questions were asked about dog-feeding prac- tices, dog control, sheep-killing procedures, and knowledge of the life cycle and control of hydatid disease. Results What emerged from our study may be con- sidered a general description of the habits and practices of dog owners that tend to maintain the cycle of hydatid disease in San- pete County. Each household selected had both sheep and dogs. The average number of dogs per household was 2.5 and the average flock size was approximately 1000. We found that nearly everyone was aware of the dis- September 1980 ScHANTZ, Andersen: Hydatid Disease 219 ease. Persons interviewed in 90 percent of the households knew of someone who had been operated on for the disease. This was usually someone from the same town, and in 10 households (25.0 percent) the victim known was a member of the nuclear or ex- tended family. Moreover, persons inter- viewed in 92.5 percent of households knew the cause of the disease and how it is trans- mitted. Specifically, they knew that people become infected with hydatid cysts by in- gesting eggs passed in the feces of infected dogs, and that dogs become infected with the hvdatid worms by ingesting the cysts in the lungs and livers of sheep. More than four-fifths of the households in- dicated they sometimes killed and butchered sheep on their premises or in the fields. De- spite their awareness and understanding of how hvdatid disease is transmitted, nearly two-thirds admitted their dogs had access to the sheep-killing area, and nearly half said the dogs sometimes ate part of the sheep car- cass. The main diet of dogs in more than 85 per- cent of households was commercial dog food, and in none was the main diet reported as sheep muscle or organ meat. Nevertheless, it was clear that most dogs could possibly eat sheep at least occasionally, since in two- thirds of households dogs were allowed to roam free, and, therefore could scavenge on sheep carcasses at the town dump or in the fields. Less than one-third of households regularly tied or locked up their dogs when the dogs were not working. Persons interviewed at more than 80 per- cent of households indicated they believed that the recommended control measures were adequate to break the chain of transmission and eliminate the infection. Persons at only 6 (15 percent) of households indicated they had taken no active measures to eliminate the in- fection. At the 34 households that indicated they had done something, the most frequent- ly mentioned steps taken were (1) periodic treating of dogs for tapeworms, and (2) dis- carding of viscera from home-killed sheep in such a way that dogs could not get to it. Four households indicated they no longer had dogs because of the potential of contracting hyda- tid disease. There was a general consensus (82.5 percent) that government authority should not make it illegal for dogs to eat parts of the sheep carcass. When the households were categorized ac- cording to whether their dogs had been found infected at only one of the first clinics or whether their dogs had been found repeat- edly infected, there were no obvious differ- ences that would allow us to conclude whv the first group of households was apparently successful in preventing reinfection. There were no statistically significant differences in the two groups regarding the number of dogs or sheep they owned, the frequency that sheep were butchered for home consumption, the apparent access of dogs to sheep viscera, the household members' knowledge and un- derstanding of hydatid disease, nor willing- ness to take measures to prevent the infection in the dogs. In fact, the responses to our ques- tions appeared to suggest that dog owners with repeatedly infected dogs were more likely to have tied their dogs up when not working and to have taken other deliberate measures to prevent their dogs from eating parts of the sheep carcass. This apparent anomaly is most likely explained by the fact that. owners of repeatedly infected dogs had more recently been made aware of what they should be doing to prevent infection than the other group of dog owners whose dogs had been given a "clean bill of health " at the most recent dog clinics. In summary, we did not learn from our study why some dog-owning households were successful in preventing reinfection of their dogs and why others were not. What was clear, however, was that numerous opportu- nities still existed at these households for dogs to become infected with hydatid tapeworms. As a result of health education and other con- trol activities, virtually all the Sanpete Coun- ty dog owners interviewed in our survey knew the basic facts ab^nit hydatid disease; however, few had actually taken all the nec- essary steps to insure its elimination. Evi- dence obtained from the survey suggests that manv dog owners apparently believe that pe- riodic treatment of dogs is sufficient to solve the problem; however, that may be an over- simplified solution. To effectively break the chain of transmission, all dogs must be pre- vented from eating the viscera of infected an- imals. This means not only that dog owners 220 Great Basin Naturalist Vol. 40, No. 3 must refrain from feeding such organs to their dogs, but, since dead sheep are fre- quently discarded in open pits and are acces- sible to roving dogs, dogs must be kept under control at all times. An additional feasible control measure would be the installation of large metal pit covers or sturdy fences at the animal pits in order to prevent ready access of roving dogs to animal carcasses discarded at those sites. From its inception in 1971, the Hydatid Disease Control Program has been an entire- ly voluntary campaign. Results of this survey suggest that some additional incentives may be necessar\' to insure that all dog owners take the necessar\' steps to stop the transmis- sion of hydatid disease. Literature Cited Aracjo. F. p.. C. W. Schwabe. J. C. Sawder, and ^\^ G. Davis. 1975. Hvdatid disease transmission in California. A study of the Basque connection. Am. J. Epidemiol. 102: 291-302. ScH.\NTZ. P. M. 1977. Echinococcosis in American In- dians living in Arizona and New Mexico. Review of recent studies. Am. J. Epidemiol. 106: .370-.379. Spru.ance, S. L., L. F. Klock. F. Cha.vg. T. Fikushima. F. L. .\ndersen, .and 1. G. K.\g.an. 1974. Endemic hvdatid disease in Utah. A review. Rocky Mtn. Med. J. 71: 17-23. Loveless, R. M., F. L. .\ndersen. M. J. Rwis.ay, and R. K. Hedelius. 1978. Echinococciis granulosus in dogs and sheep in central Utah. 1971-1976. Am. J. Vet. Res. 39: 499-502. NEW GRASS DISTRIBUTION RECORDS FOR ARIZONA, NEW MEXICO, AND TEXAS' Stephan L. Hatch- Abstr^ct.— New distribution records are given for seven grass species now found in Arizona, New Mexico, and Texas. Recent collections have revealed new dis- tribution records for seven southwestern grass species. These records are extensions to the known distributions of these species as re- corded in general for the United States bv Hitchcock (1951), in Arizona and Te.xas by Gould 1 1951. 1975^. and in the intermountain area by Cronquist et al. il977>. The checklist for New Mexico published by Martin and Castetter (1970) is the basis of distributions for that state. Voucher specimens for these new records have been distributed in the Tracy Herbarium (^TAES). Eremopyron triticeum (Gaertn.^ Nevski .\rizona: Coconino Co.: Locallv abundant as an adventive on disturbed sites. 0.5 km north of Fredonia, east side of Highway 89A. on a hard gra\ clav soil at 1600 m elevation, 17 May 1978. Brown 652 (TAES). This in- troduced grass is a new state record for Ari- zona. Previous collections have been report- ed from Oregon, Idaho, Nevada, and Utah by Cronquist et al. il977'. Montana (Hitchcock 1951k and New Mexico ^Hatch 1977). Afrna barbota Brot. New Mexico: Dona Ana Co.: Rare adven- tive along Interstate 10. 10 km south of Las Cnices, 3 April 1978. Mocliange 21 (TAES). This is a new state record for New Mexico. Previous reports show a distribution from Washington to Arizona ^Gould 1951). Bothriochloa ischaemum yh.) Keng. New Mexico: Colfax Co.: Locally abun- dant as an adventive. 3.5 km south of Raton. 18 August 1978, Hatch 4072 (TAES). Dona Ana Co.: Locally abundant. 11 km south of San Agustin Pass on the north slopes of the Organ Mountains. 18 September 1977, Dick- Peddie 55 i^TAES). Grant Co.: .\bundant, 32 km north of Silver City along Highwav 15, 9 October 1977. Hoefler'33 TAES). Sierra Co.: Five km south of Williamsburg along Inter- state 25, 26 August 1977, Hatch 2604 I TAES). This is a new state record for New Mexico. Gould (1975) reported this species as being introduced in Te.xas as a pasture grass. Dactyloctenium aegyptium (L.i Beauv. New Mexico: Dona Ana Co.: Rare as an adventive in the New Mexico State Univer- sitv Agronomy Field Laboratory land. Las Cruces, 10 September 1977, Hatch 2565 TAES). This is a new record for New Mexi- co. Gould ^195L reported this species from Arizona and later ^1975) from Te.xas. Gould (1975) stated that this species was well adapt- ed to sandy soils of the southern United States. Eragrostis superba Peyr. New Me.xicg: Dona Ana Co.: A relatively rare introduced grass along Interstate 10. 6 km south of Las Cruces in sandy soil. 15 Oc- tober 1977, Yehca 3S (T.\ES). This is a new state record for New Mexico. The plant has been collected in Texas 22 May 1957. Gould 7550 (TAES), but was not included in Gould's (1975) Grasses of Texas. This species has been collected in .\rizona. 3 November 1961. Mat- tox and White sn. (T.\ES). It is native to South Africa and was introduced to the United States as P.I. 185516 (No. 39) 12 De- cember 1949. Technical Bulletin T..\. 15803, Texas Agricultural Eipenment Stabon. Teias A & M L'niversit). College Station. Texas . iS*3. 'Department of Range Science. Tesas A & Vt Universitv-. College Station. Teias 77843. 221 222 Great Basin Naturalist Vol. 40, No. 3 Leptoloma arenicola Swallen New Mexico: Lea Co.: A rare-abundant perennial grass on sandy sites, 22 km east- northeast of Maljamar along Highway 82, 10 August 1976, Elhworth 23 (TAES). This col- lection is a new state record for New Mexico. Hitchcock (1951) and Gould (1975) report this taxon as occurring only in Kenedy Co., Texas. This is a rhizoinatous form of the widespread L. cognatum (Schult.) Chase, classified by Gould (1975) as L. cognatum var. arenicola (Swallen) Gould. Texas: Kent Co.: Locally abundant in stands of shinnery oak {Querciis havardii Rydb.), 3.0 km south-southeast of Girard, 9 June 1979, Slosser sn. (TAES). The Slosser collection is a distribution extension within Texas of several hundred miles. Neeragrostis reptans (Michx.) Nicora New Mexico: Sierra Co.: Locally abun- dant on the mudflats of Elephant Butte Res- ervoir, near Alamosa, 10 September 1978, So- pyn sn. (TAES). This is a new record for New Mexico. Gould (1975) reported this species as being abundant on exposed lake beds in the central United States from Kentucky and South Dakota to Louisiana, Texas, and Flor- ida. Literature Cited Cronquist, a., B. H. Holmgren, N. H. Holmgren, J. L. Reveal, and P. L. Holmgren. 1977. Inter- mountain flora. Vol. 6. Columbia Press. New York. 584 pp. Gould, F. W. 1951. Grasses of the southwestern United States. University of Arizona Press, Tucson. 352 pp. 1975. The grasses of Texas. Texas A & M Univer- sity Press, College Station. 65.3 pp. H.\TCH, S. L. 1977. New grass distribution records for New Mexico and the United States. Great Basin Nat. 37:530-531. Hitchcock, A. S. 1951. Manual of the grasses of the United States. USDA Miscellaneous Publication 200, rev. by Agnes Chase. 1051 pp. Martin, W. C, and E. F. Castetter. 1970. A checklist of gymnosperms and angiosperms of New Mexi- co. Published privately. A COMPARISON OF EPIPHYTIC DIATOM ASSEMBLAGES ON LIVING AND DEAD STEMS OF THE COMMON GRASS PHRAGMITES AUSTRALIS Judith A. Grimes'. Larry L. St. Clair', and Samuel R. Rushforth' Abstract.- Diatoms epiphytic on Phragmites austmUs (Cav.) Trin. ex Steaded stems were collected from a single clone at the .southern end of Provo Bay, Utah Lake, Utah. Diatom populations from both living and dead stem sec- tions were analyzed. Species diversity in each .sample was high, indicating that the stems provide a relatively stable habitat for diatom epiphytes. Of the 23 genera found, only Gomphonema and \avicula showed significant trends toward stem preference. The diatoms in this study support the current view that Utah Lake is a slightly saline, eutrophic system. The occurrence of diatom assemblages as epiphytes on httoral, emergent macrophytes is well documented (Godward 1934, 1937, Knud.son 1957, Prow.se 1959). Likewise, the impact of such epiphytes on primary produc- tivity and community trophic structure has been examined in several estuarine environ- ments (Mclntire et al. 1971, Stowe et al. 1971, and Main et al. 1974), but has been largely ignored in freshwater systems (Wetzel 1964). Tlie epiphytic diatom communities at- tached to emergents inevitably play a role in the overall productivity of lakes and es- tuaries. They also contribute to regulation of the overall metabolism of such waters by al- tering the amount and quality of alloch- thonous organics entering the lake by acting as physical and metabolic traps or filters. The attached diatom flora also serves as an au- tochthonous source of particulate organic and dissolved organic matter that is readily available to pelagic animals. The degree of influence of these epiphytic organisms on the productivity of standing waters has rarely been determined. However, Allen (1971) esti- mated that up to 31.3 percent of the total lit- toral production could be attributed to epiphtyic algae, with up to 21.4 percent of the total lake production being attributable to such attached communities. In addition, a comparison between phytoplankton and epi- phyte production demonstrated that the lat- ter was equivalent to 75 percent of the phytoplankton production over the annual period (Allen 1971). Even though epiphytic commimities have been demonstrated to be important, the dis- tribution patterns of such assemblages on the basis of variation in ho.st species and host sub- strate conditions have received little atten- tion. Likewise, the complex physiological relationship between the host macrophyte and the attached diatom species has received less attention than warranted (Wetzel 1964, 1965, 1969b, Allen 1971, Hough et al. 1975). The impact of this relationship is fimdamen- tal to understanding the basic distribution patterns of epiphytes not only on different macrophyte host species but also on members of the same species at different levels of se- nescence. The purpose of this study is to illuminate distribution patterns of diatom epiphytes on living and dead specimens of a single macro- phyte host {Phragmites australis (Cav.) Trin. ex Steaded) in Utah Lake, Utah. The data from this study will be used as a baseline for extended research in Utah Lake on epiphyte distribution patterns and epiphyte impact on lake productivity and trophic structure. Methods Samples were collected 20 September 1978 from a single clone of Phragmites ous- trcilis located at the southern end of the mouth of Provo Bay in Utah Lake. Five sam- ples of living and five of dead Phragmites australis stems were collected as cut 10 cm sections, measured from the water level 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 223 224 Great Basin Naturalist Vol. 40, No. 3 down. Samples were prepared according to standard acid-oxidation methods, and per- manent diatom slides were made using Naph- rax diatom momitant (St. Clair and Rushforth 1977). Slides were examined and diatom spe- cies were identified at lOOOX with a Ziess RA research microscope with bright field and Nomarski interference phase-contrast acces- sories. Quantitative data on the diatom assem- blages were recorded by counting 250-400 diatoms for each sample. Previous studies have shown that a statistically valid count can be obtained within this range (Squires 1977). Each slide was then thoroughly scanned to record the rare species. The re- sults were converted into percent relative density values for all species for each site. Shannon-Wiener diversity indices were cal- culated for individual samples (Shannon and Wiener 1963). The relative density figures for each sample were compared to all other samples and similarity indices were calculated (Ru- zicka 1958). These indices were clustered (Sneath and Sokal 1973) to identify unique community associations within and between the living and dead Phragmites aiistralis stem sections. The most prevalent diatoms encountered in the study and the diatoms that significant- ly differed between the living and the dead specimens of Phragmites australis were deter- mined. Results Twenty-three genera and 114 diatom spe- cies were found on the 10 Phragmites austral- is stem sections (Table 1). The most prevalent diatoms throughout the study were Navicula graciloides, Nitzschia inconspicua, and Nit- zschia filiformis (Table 2). Nitzschia dis- sipata, Stephanodiscus astrea var. minutula, and Nitzschia palea were also common. Among the prevalent species, Amphora ve- neta was found to occur only on the living stems, whereas Navicula schroeteri var. es- camhia was essentially restricted to the dead stems. Table 1. Alphabetical list of the diatom taxa found on living and dead Phragmites australis stem sections from Provo Bay, Utah Lake, and their average relative density. Species Living Dead Achnanthes hauckiana" .07 _ Achnanthes lanceolata var. dtibia" — P Achnanthes lewisiana .06 P Achnanthes minutissima 2.20 .57 Achnanthes sp." P — Amphora ovaUs .31 P Amphora ovalis var. pedicuhis' — .08 Amphora perpusilla .76 1.79 Amphora veneta 8.78 1.26 Amphora sp.°° .19 - Anomoeoneis sphaerophora" ° P — AsterioneUa formosa .09 .08 Colonels fenzlioides' ° .02 — Cocconeis placenttila var. eughjpta' — P Cocconeis placentula var. lineata .07 .07 Coscinodiscus sp. .06 .06 Cyclotella kiitzinghiana' — .13 Cijclotella meneghiniana 1.87 1.80 Cijmbella affinis' — P Cymbella miniita" — .06 Cijmbella muelleri" .14 - Cymbella prostrata .07 .05 Cymbella sp. ° ° P — Diatoma tentie var. elongatum .13 .23 Diploneis oblongella' ° .02 — Epithemia adnata var. porcellus P P Epithemia adnata var. proboscidea" - P Fragilaria brevistriata" — P Fragilaria brevistriata var. inflata .06 .15 Fragilaria construens' — P Fragilaria constriiens var. binodis .06 P Fragilaria constrtiens var. venter .65 .11 Fragilaria crotonensis' — .17 Fragilaria lapponica" .19 - Fragilaria pinnata" .37 — Fragilaria pinnata var. lancettula .07 .13 Fragilaria similis .19 .17 Fragilaria vaucheriae .90 .63 Fragilaria virescens" P — Gomphonema affine" .13 - Gomphonema gracile" .02 — Gomphonema intricatum' — P Gomphonema olivaceiim .30 .05 Gomphonema parvulum .44 .21 Gomphonema subclavatiim var. commutatum .08 P Gomphonema tenellum" .02 — Gomphonema ventricosum' ° .44 — Gomphonema sp. ° ° .06 — Melosira granulata .55 .65 Melosira granulata var. angustissima 1.58 1.05 Melosira italica .21 P Navicula arvensis' _ .15 Navicula arenaria' — P Navicula aurora" .02 - Navicula capitata var. hungarica' — .1 Navicula cincta' — .59 Navicula circumtexta' — .15 Navicula cryptocephala .29 .35 September 1980 Grimes et al.: Diatom Assemblages 225 Table 1 continued. Species Living Dead Navicula cryptocephala var. veneta 3.45 4.38 Navicuhi cxigua" — P Navicuhi gmciloides 12.35 9.17 Navicula Iwiifleri var. leptocephala .08 P Navicuhi lanccolata ° — .08 Navicula tnitiima .07 .84 Navicula oblonga P .08 Navicula peregrina' - .08 Navicula pupula P .08 Navicula radiosa var. tcnclla 1.13 2.42 Navicula rh yncocephala ° — P Navicula salinarum ° — .13 Navicula salinarum var. intc rmedia .08 .05 Navicula schroetcri var. cscambia 3.06 6.23 Navicula tcnclloidcs .15 1.27 Navictila tripunctata P .33 Navicula tripunctata var. sch lizonemoides .44 1.51 Navicula sp.° — .63 Nitzschid acicularis .51 .66 Nitzschia amphibia .37 .25 Nitz-schia apiculata .06 .18 Nitzschia di.ssipata 6.00 4.95 Nitz-'ichia filiformis 7.10 6.83 Nitzschia frustulum .80 .53 Nitzschia gracilis" .23 — Ni tzsch ia han tzsch ia na 4.94 4.40 Nitzschia holsatica 1.67 2.35 Nitzschia inconspicua 12.40 13.25 Nitzschia linearis .06 .08 Nitzschia longissima var. clostcrium .08 .08 Nitzschia ovalis .18 ..32 Nitzschia palea 6.90 13.50 Nitzschia paleaceae 4.63 3.92 Nitzschia punctata' — P Nitzschia sigmoidca ° — P Nitzschia stagnorum' " P — Nitzschia s-p. 1 P .42 Nitzschia sp. 2°° .21 — Ophephora martiji' - P Rhoicosphenia curvata 3.96 6.32 Rhopalodia gibba° - .05 Rhopalodia gibberuhi var. vanheurckii .10 P Stcphanudiscus astrea" .06 - Stcphanodiscus astrea var. minutula 5.15 4.34 Stcphanodisctis niagarae ° - .08 Surirella angustata P P Surirella ovalis var. brightwe llii° — .15 Surirella ovata°° .08 — Sijnedra actis .08 1.60 Sijnedra dclicatissima var. angiistissima .15 .02 Sijnedra fasciculata var. trunctata' - .08 Sijnedra mazamaensis .15 .08 Sijnedra socio P .08 Synedra ulna P .11 Synedra ulna var. contractu .81 .02 °° Species unique to living Phragmites australis stems. 'Species unique to dead Pbragmites australis stems. P Species not recorded on the transects taken for relative de ures but found on other sections of the diatom slide. iity fig- Species diversity according to number of species encountered was high, averaging 48 species per sample. However, there were generally 2 to 4 dominant species ranging be- tween 10-18 percent relative density in each sample, which allowed for only moderately high Shannon-Wiener diversity values (Table 3). Forty-nine percent of the diatom species was found in 30 percent of all samples, and 34 percent of the diatom species was found in 50 percent of all samples. The results of the cluster analysis (Fig. 1) demonstrate the high degree of similarity en- countered for all ten .samples. Even so, mar- ginal separation into .samples from living and dead stems was obtained. A similarity matrix comparing all 10 sam- ples was constructed. Means of similarity indices for living stems, dead stems, and between living and dead stems were computed. T-tests were per- formed and it was determined that there was no significant difference in similarity within or between these samples sets. Discussion Each of our 10 samples consistently con- tained approximately 50 identifiable species. In general, no one species represented more than 18 percent of the total population of any sample. These conditions are indicative of a diverse flora that is further supported by our Shannon-Wiener diversity values and the average number of species per substrate (Table 3). Such conditions indicate that the epiphyte flora in Utah Lake is more diverse than we previously believed. T-tests were computed comparing the means of the Shan- non-Wiener diversity indices of both .sub- strates as well as the average number of spe- cies from both living and dead stems. No significant differences between the values in either compari.son existed. A total of 23 diatom genera were encoun- tered during this study. The number of spe- cies included in these genera was nearly equally distributed between living and dead stems (Table 4). However, substratum prefer- ences were noted in the genera Navicula and Gomphonema and in individual .species with- in several other genera. Of the 114 species found in the study, 22 were unique to living and 29 were unique to dead stems. 226 Great Basin Naturalist Vol. 40, No. 3 T\BLE 2. Important species encountered on Phragmites australis stem sections from Provo Bay, Utah Lake, with their percent relative densities. Important species are those species with a percent relative density greater than .3 percent in any one sample. Livin g stems Dead stems Species 1 2 3 4 5 6 / 8 9 10 Achnanthes minutissima 3.1 Amphora perpusiUa 3.3 3.2 Amphora ceneta 17.3 5.6 7 2 11.2 Cyclotella meneojiiniaiui 3.0 Melosira granulata var. angustissima 3.9 S'avicula cnjptocephala var. veneta 3.4 4.9 4.2 3.7 5.4 3.3 6.1 3.5 Xaiicula graciloides 9.9 13.2 13.5 14.0 11.2 5.5 13.4 14.9 8.5 13.6 Naiicula radiosa var. tenelki 3.3 3.2 6.2 Saviciila schroeteri var. escambia 5.7 6.3 6.2 6.2 10.6 \aviciila tripunctata var. scliizonemoides .3.3 \itzschia dissipata 6.8 8.7 3.6 7.8 3.1 4.1 4.6 7.9 3.6 4.7 Xitzschia filifonnis 3.1 12.5 5.4 11.1 3.5 6.6 3.2 7.0 5.2 12.2 Xitzschia inconspicua 15.5 23.4 9.5 7.3 12.2 25.7 6.2 12.1 10.1 .Vitast7i ia hatiztsch ia no 3.1 8.7 4.6 5.4 12.3 3.1 Xitzschia holsatica 3.7 3.3 5.4 .3.9 Xitzschia palea 5.0 5.7 5.7 7.8 10.4 17.7 9.4 12.4 16.1 12.0 Xitzschia paleacea 10.6 4.5 6.8 5.0 3.3 7 7 7.4 Rlioicosph en ia cu na ta 7.7 6.2 6.6 9.4 12.2 Stephanodiscus astrea 1 var. miniitida 3.4 5.3 5.2 6.2 5.5 4.6 4.1 3.2 43 1 The most important species in each sample are indicated by boldface type. Table 3. Shannon-Wiener diversity values for the five living and five dead Phragmites australis stem sections from Provo Ba\. Utah Lake. Sample No. Livins Dead Living Dead 2.96 2.96 2.86 2.96 3.07 2.962 3.12 2.98 3.09 2.85 3.15 3.038 48 50 33 44 47 44.4 52 47 50 38 51 47.6 *.\verage number of species/substrate. An analysis of the diatom types unique to the living steins reveals that most were peri- phvtic stalk formers, whereas those imique to the dead stems were mostly periphytic mo- bile forms. The distribution of species of Gompho- nema and Navicula on Phragmites stems showed significant deviation from random. Thus, of a total of nine Gomphonema species encoimtered during this study, eight of these occurred on living stems, five of which were restricted to hving stems. Conversely, of four species that occurred on dead stems, only one was restricted to that habitat. These data sug- gest that the living stems provide a more suitable substrate for several Gomphonema species. Such species tend to be strictly epi- phytic in distribution, usuallv being attached by a gelatinous jelly stij)e (Patrick and Rei- mer 1966). Whether the preference of these species for living stems is relative to the availability of nutrients or the physical condi- tion of the substrate is yet to be determined. Some interesting distribution patterns were also observed in the 26 species of Navicula. Twenty-five of these were found on the dead stems, of which 11 were restricted to that substrate. On the other hand, onlv 15 Xav- September 1980 Grimes et al.: Diatom Assemblages 227 20 30 PERCENT SIMILARITY 40 50 60 70 —I 1—5 Living 6—10 Dead 2 10 4 Fig. 1. Cluster dendrogram showing similarities of diatoms on living and dead Phragmites australis stem sections from Provo Bav, Utah Lake. icula species were observed on the living steins and only one taxon was restricted to living stems. We believe the high number of Navicula species in our samples can be accounted for, at least in part, by the fact that many are op- portimistic, occurring on a wide variety of substrates. These opportunistic Navicula spe- cies occurred primarily on dead Phragmites stems except for one or two species that dom- inated both living and dead stems. The rea- son for this is open to speculation, but it is probably related to nutrient interaction, the physical condition of the substrate, or re- duced competition on the dead stems. The hypothesis that condition of the Phrag- mites stems had no effect on the presence or absence of Gomphonema and Naviculu was tested by chi-square analysis using a 2 X 2 contingency table. The results departed sig- nificantly from random. This supported the concept that Gomphonema and Navicula were separated on the basis of habitat type. Consistent with other Utah Lake studies, the diatoms in this study reflect the condition of the lake waters. Most of the prevalent dia- toms were either alkaphilous or alkabiontic forms and also indicators of eutrophy. Addi- tionally, many are known to have the ability to withstand elevated levels of dissolved salts. These data, together with the elevated diver- sity found at Utah Lake, support the current view that Utah Lake is a saline-eutrophic ecosystem. We recognize the preliminary nature of the present study. Even so, we believe the differences shown in communities on the liv- ing versus the dead stems are significant. Fu- ture studies are planned to expand our data 228 Great Basin \atur.\list Vol. 40, No. 3 Table 4. Alphabetical list of diatom genera found on Phragniites australis stem sections from Pro%o Bay. Utah Lake, and the occurrence of species from those genera on Hvine and dead substrates. Genus Li\ing Dead m^in Achnanthes Amphora Ationioeneis Asterionella Caloneis Cocconeis Coscinodisctis Cyclotella Cymbella Diatonia Diploneis Epithemia Fragilana Goinphonema Melosira Savicula Sitzschia Ophepbora Rhocosphenia Rhoipalodia Steplianodiscits Surirella Synedra 4 4 1 1 1 1 1 1 3 1 1 1 9 8 3 15 18 1 1 2 2 6 1 2 9 4 3 25 17 1 1 2 2 ■7 base to the other species of emergent macro- ph\1:es in Utah Lake. Furthermore, we plan studies to answer the following questions: (1) Are some epiphytes host specific? (2^ What patterns of seasonal succession are evident in the epiphytic flora? (3) \Miat impact does the epiphxtic flora have on productivity and tro- phic structure of the lake? These questions take on added significance for future re- source management in light of proposed large-scale changes in Utah Lake, such as the dikino; of Provo and Goshen bavs. LiTER-\TURE Cited Allen. H. L. 1971. Primar)- productivity, chemo-organ- otrophy. and nutritional interactions of epiphvtic algae and bacteria on macroph\tes in the littoral of a lake. Ecol. Monogr. 41(2i:97-127. GoD\v.\RD. Nf. B. 19.34. An investigation of the causal distribution of algal epiph\tes. Beih. Bot. Cen- tralbl. .52A:506-539. 19.37. .\n ecological and taxonomic investigation of the littoral algal flora of Lake Windermere. J. Ecology 25:496-568. Hough, R. A., .\nd R. G. Wetzel. 1975. The release of dissolved organic carbon from submersed aquatic macroph\tes: diel, seasonal, and communitv rela- tionships. Verb. Int. Ver. Limnol. 19:939-948. Kncdson. B. M. 1957. Ecology- of the epiphytic diatom TabeUaria floccuhsa (Roth^ Kutz. var. flocculosa in three English lakes. J. Ecolog> 45:93-112. S. P.. AND C. D. McI.vTiRE. 1974. The distribution of epiph\tic diatoms in Yaquina Estuar\'. Oregon. Bot. Mar. 17:88-99. McIntire, C. D.. and W. S. Overton. 1971. Distribu- tional patterns of assemblages of attached dia- toms from Yaquina Estuarv. Oregon. Ecologv 52:758-777. P.\TRiCK, R.. .\ND C. Reimer. 1966. The diatoms of the United States. Acad. \at. Sci. Phil.. Monograph 13. V. 1. 688 pp. Pronvse. G. a. 1959. Relationship between epiphvtic al- gal species and their macroph\tic hosts. Nature 183:1204-1205. Rlshforth. S. R.. L. L. St. Cl.\ir. J. .\. Grimes. J. R. JoH.\NSEN. A.ND M. Whitinc. The phytoplankton of Utah Lake. Great Basin Nat. Mem. 5. In press. RrziCK-\. M. 1958. Anwendung mathematisch-statistic- her methoden in der geobotanik iSvTithetische bearbeitung von aufnahmen). Biologia Bratisl. 13:647-661. Sh.vnno.n, C. E.. .and \\'. \\'iE.NER. 1963. The mathemati- cal theory of communication. University of Il- linois Press. Urbana. Sneath. R. H. .\., .\ND R. R. SoK-\L. 1963. Numerical tax- onom%': principles and practice of numerical clas- sification. \\". H. Freeman Co.. San Francisco. 573 pp. Snedecor. G. ^^'., .\ND W. CocHR.\N. 1968. Statistical methods. Iowa State Press. 593 pp. Squires. L. E., S. R. Rushforth. .\.sd J. D. Brotherso.n. 1979. Algal response to a thermal effluent: study of a power station on the Provo River. Utah. U.S.A. Hvdrobiologia 63(1): 17-32. St. Cl\ir, L. L., .\.nd S. R. Rushforth. 1977. The dia- tom flora of the Goshen Warm Springs ponds and wet meadows, Goshen, Utah. U.S..\. Nova Hed- wigia 28:35.3-425. Stowe. W. C.. .\nd J. C. GossELiNK. 1971. Community structure and production of the epiphvtic algae in the Barataria Bay area of Louisiana. Paper read at the .34th annual meeting of the .\merican Society of Limnolog)' and Oceanography, \\inni- peg. June 14-17. ^^'ETZEL. R. G. 1964. A comparative study of the pri- mary productivity of higher aquatic plants, per- iphvton. and phvtoplankton in a large, shallow lake. Int. Rev. Ges. Hvdrobiol. 49:1-61. 196.5a. Techniques and problems of primary pro- ductivity measurements in higher aquatic plants and periphvton. Mem. 1st. Ital. Idrobiol, Suppl. 18:147-165.' 1969b. Factors influencing photos\-nthesis and e.x- cretion of dissolved organic matter by aquatic macroph\tes in hardwater lakes. Verb. Int. \'er. Limnol. 17:72-85. POISONOUS PLANTS OF UTAH Jack D. Brotherson.' Lee A. Szyska,' and William E. Evenson- Abstr\ct.— a list of the major livestock-poisoning plants has been compiled for the state of Utah. Two hundred fifteen taxa representing 36 families. 119 genera, and 209 species occur within the state. Fortv-one percent are from two families, the Asteraceae and the Fabaceae. The remaining families of major imjx)rtance are: Poaceae, Ranuncu- laceae. Solanaceae. Chenopodiaceae. Brassicaceae. .\scelpiadaceae, Liliaceae. and Euphorbiaceae. Sixtv-nine percent of the genera occur with a single species. Thirty-three percent of the ta.xa are introduced to the state. .Most of the plants are insect pollinated; 57 percent are herbaceous perennials. Most livestock poisoning occurs during the spring. This is due both to concentration of toxins in emerging vegeta- tion and to the absence of more suitable forage on late winter and spring ranges. Green herbage is poisoning in about 80 percent of all taxa, seeds and fruits in about 15 percent, and the remaining 5 percent have toxic compounds confined to flower heads, sap. tubers, or roots. Disturbed or cultivated habitats and poorlv managed range harbor the greatest diversity of poisonous plants. Wetlands contain fewer poisonous taxa than do xeric or mesic areas. The predominating plant toxins are various alkaloids and glycosides. Sixteen percent of the plants have un- characterized toxins. Cattle and sheep are more susceptible to poisoning than are horses, swine, or poultrw Records document man's encounters with poisonous plants since ancient times. Thev have played both positive and negative roles in human cult\ire (Dayton 1948). This con- spicuous dualitv of poisonous plants remains a major concern for range management. Kingsbury's (1964) manual on the poisonous plants of the United States and Canada was designed to aid veterinarians and ranchers in recognizing poisonous plants and the svmp- toms they produce in poisoned livestock. Val- entine (1978^ prepared an extensive bibliog- raphv on the poisonous plants of American rangelands, and numerous works have been published dealing with local species lists and descriptions (Evers 1972, Mihalopoulus 1974, Schmutz et al. 1968. Stoddard et al. 1949. USDA 1968). The scope of this paper is twofold: to pro- \"ide a list of taxa of the major poisonous plants of Utah, and to present some general patterns observed among poisonous plants within the state. It is hoped that this anno- tated compilation and discussion will prove useful to range managers and biologists alike. NL\TERIALS .\.ND METHODS Data on poisonous taxa were gleaned from the published literature and by consultation with specialists in botany and toxicolog}-. Much of the descriptive literature on poi- sonous plants is redundant, consequently, only the more recent works are cited here. Criteria used in compiling the list of poi- sonous plants were: 1. The ta.xon had to be sufficientlv abun- dant I either native or introduced) in natural ecosystems to constitute a legiti- mate threat to livestock or wildlife. For example, some species of the genus As- tragalus are known to be toxic but are not abundant enough within the state to be considered dangerous (Williams and Bameby 1977). 2. Ornamentals were included only if they have escaped widelv from cultivation. Such plants are frequent along ecotones or in disturbed habitats. 3. Suspicions of toxicity had to be reason- abh well-founded. The genus Astra- galus, for example, is represented by more than 100 species in Utah (Welsh 1978), but only those ta.xa demonstrably to.xic were included in the present list- ing. Additional variables considered for each taxon were: life history strategy (annual. biennial, perennial), patchiness of distribu- Department of Botany and Range Science. Brigham Young University, Provo, l"tah &4602. Department of Phvsics and .\stronomv. Brigham Young University. Provo, Utah 84602. 229 230 Great Basin Naturalist Vol. 40, No. 3 tion, occurrence as a cultivar or as a common range plant, growth form (vine, forb, grass, shrub, or tree), generalized habitat require- ments including elevation, soil texture and acidity, and moisture preference, nature of the toxin and its localization within the plant, seasonality of poisoning, animals affected, specific juvenile mortality and abortifacient properties. Not all variables could be docu- mented for each species. Results Taxonomy Conservatively estimated, Utah has at least 215 major toxic taxa of plants representing 36 families, 119 genera, and 209 species. Thirty- three percent of these taxa are introduced to the state. Two of these 36 families, the Aste- raceae and the Fabaceae contain 41 percent of the total known taxa of poisonous plants (Table 1). In decreasing order of floral promi- nence, the eight next important families are: Poaceae, Ranunculaceae, Solanaceae, Che- nopodiaceae, Brassicaceae, Asclepiadaceae, Liliaceae, and Euphorbiaceae. The number of toxic taxa within a family is not related to the degree of toxicity of individual taxa: two of the most deadly plants, rosary pea or pre- catory bean {Abms precatorious) and poison hemlock {Conium maculatum) belong to the Fabaceae and Apiaceae, one major and one relatively minor family if numbers alone are considered. Table 1. Poisonous plant taxa by families. Family No. taxa Percent of taxa occurring in each family Fabaceae 50 23 Asteraceae 39 18 Poaceae 19 9 Ranunculaceae 13 6 Solanaceae 11 5 Chenopodiaceae 9 4 Brassicaceae 8 4 Ascelepiadaceae 7 3 Liliaceae 7 3 Euphorbiaceae 6 3 26 other families 46 21 Total 215 Sixteen genera (13 percent) occur in the flora with more than two taxa. Twenty-two other genera (18 percent) occur with two taxa (including Cannabis, which has two sub- species of a single species). Sixty-nine percent of the genera occur with a single species. The taxonomy of poisonous plants is not readily explained. The poisonous flora of the eastern half of the United States resembles that of Europe more than it does that of the western states (Kingsbury 1961), and Day- ton's paper (1948) on the poisonous plants of the continental United States gives a differ- ent listing of major families than that found for the state of Utah alone. Table 2. Major toxins found in Utah's poisonous plant flora. Toxin Percent of taxa in which toxin occurs Alkaloids 22 Glycosides 22 cyanogenetic glycosides goitrogenic glycosides irritant oils coumarm steroids and triterpenoids Seleniinn 11 Nitrates 5 Unknown toxins 16 Other toxins 19 amines oxalates resins and resinoids photosensitizing compounds nutritional deficiencies Mechanically injurious 4 Plant Toxins Most poisonous principles are secondary by-products of the plants' metabolism (Kings- bury 1964, Levin 1976). In Utah's flora, many of these toxic metabolites are loosely classed as alkaloids and glycosides (Table 2). These two biochemical groups are primarily arti- ficial constructs, and each consists of numer- ous toxins (only a few of which have been identified) having generally similar molecular structures or modes of activity. When the impact of poisonous range plants on livestock mortality is assessed, how- ever, many of the minor toxins become or September 1980 Brotherson et al.: Poisonous Plants 231 serious concern to ranchers. Selenium poison- ing by members of the genus Astragalus and other species is an important source of mor- tahty, as is oxalate poisoning by halogeton Halogeton glomeratus) and fivehook bassia Bassia hyssopi folia), and loss of sheep due to photosensitization by plants such as spring jarsley {Cijmoptcrus icatsonii) and St. Johns- .vort {Hypericum fowiosuui). Thus, the com- nonness of a particular plant species or toxin loes not necessarily imply high mortality. Seasonality and Specificity Most livestock losses occur in the early pring (Table 3) as animals are turned out into slowly greening range (Evers 1972, Kee- er 1978, Kingsburv 1964, Kreger and Sharp 978, Merrill and Schuster 1978). A second, mailer peak in mortality follows in the sum- ler as the more palatable vegetation withers ^ the heat and toxins are concentrated in ruits and seeds of poisonous species. Cattle are susceptible to poisoning by lore taxa than sheep, with consequently igher mortality rates and greater economic )ss (Nielson 1978). Most of the toxic taxa ommonly found on Utah's rangeland will 'oison all livestock but others are relatively pecies specific (such as Delphinium poison- ig in cattle— ranges infected with the vari- us species of this plant may be safely grazed y sheep). Other poisonous taxa may be razed in moderate amounts without harm if Iternative palatable forage is available, and inie species, for example, halogeton, can be tilized by sheep if the animals are in- oduced gradually to the plant, allowing leir rumen microflora to adapt to etoxifying large amounts of calcium oxalate ameset al. 1976). Table 3. Livestock inortalitv bv seasons. •ason Percent of reported poisonings .ring" 65 miner 17 ill 11 inter 6 Mortality especially high in the late winter and earlv spring The Ecology of Poisonous Plants The ecology of Utah's poisonous flora is highly variable, but our data permit a few general observations. Most of the plants are insect pollinated, and 57% are herbaceous perennials. Green herbage is toxic in about 80% of all taxa, .seeds and fruits in about 15%, and about 5% of the taxa have toxins con- fined to flower head, sap, tubers and roots. The relative locations and concentrations of toxins within a plant may vary through the growing season, and depend upon the par- ticular taxon being considered. Toxic species are likely to be found any- where in the state, although disturbed habi- tats and poorly managed rangelands are espe- cially prone to harbor dangerous species. The predominance of poisonous plants in these habitats reflects both a bias in the reported literature and the very real dangers of over- grazing in the western states. Dry desert soils have more taxa of poisonous plants than mes- ic or hydric ones; however, some of the indi- vidually most deadly plants occur in wet- lands. Data on elevation, soil texture and acidity were insufficiently complete to per- mit any valid conclusions. Discussion and Summary The benefit to the plant of manufacturing and maintaining toxic substances is unknown. A few phytotoxins such as abrin and cicuto- xin appear to function specifically as verte- brate poisons (Kingsbury 1961, 1964) and may have evolved in response to herbivore pressures (Laycock 1978). Selenium accu- mulators are toxic due to characteristics of the soil, although primary accumulators may actually require trace amounts of selenium for proper growth (Kingsbury 1964). There does not appear to be any univer- sally applicable method for managing range- lands infested with poisonous plants. Species that occur in dense clumps or thickets may be individually irradicated by chemical or mechanical means. More commonly, how- ever, vast areas will be infested, often as a re- sult of overgrazing. Wise management will include aspects of the following practices: 1. Recognition of poisonous plants and an accurate assessment of their potential 232 Great Basin Naturalist Vol. 40, No. 3 lethality. Unless forced by hunger, live- stock generally will ignore poisonous plants in favor of more palatable for- age. 2. Removal of susceptible animals. 3. Provision of sufficient alternate forage if animals must be turned onto ranges in early spring before palatable plants are abundant. 4. Scheduling of range use around live- stock susceptibility patterns. There are several good reviews of manage- ment practices (Evers 1972, Keeler 1978, Knieger and Sharp 1978, Merrill and Schus- ter 1978), and the range literature abounds in articles dealing with specific plants and their effects (see Valentine 1978 for a com- prehensive listing). The purpose of the pres- ent paper is not to review management tech- niques in detail, but primarily to provide an updated listing of Utah's dangerous flora. Further research needs to be done not only in management but in toxicology and pharma- cology so that control of poisonous plants will be a matter of understanding instead of irradication and vast areas of range can again be utilized and productive. POISONOUS PLANTS OF UTAH The present list of plant taxa was gleaned from published literature and by consultation with experts in botany and toxicology as cited above. Certain plant characteristics are desig- nated for each taxon in parentheses immedi- ately following the taxon name. The abbre- viations are as follows: p Perennial B Biennial A Annual N Native I Introduced T Tree S Shnib F Forb G Crass R Rush The following list of families, genera, spe- cies, and varieties is arranged in alphabetical order for ease of reference. Plant synonymy follows Manual of the Plants of Colorado (Harrington 1964) and Utah Plants (Welsh and Moore 1973). Amaranth ACE AE Amaranthus retroflexus L. (AIF) Common name: pigweed, carelessweed, redroot, redroot amaranth. Toxin: nitrates, under conditions of over- fertilization with too little water. Habitat: common garden and field weed; waste places. Animals affected: livestock. Reference: Brakenridge 1956, Kingsbury 1964. Anacardiaceae Toxicodendron rydbergii (Small) Greene (PNS) Common name: poison ivy. Toxin: 3-n-pentadecylcatechol. Habitat: moist areas at lower elevations. Animals affected: humans (dermatitis). Reference: McNair 1923, Kingsbury 1964. Apiaceae Cicuta douglasii (DC.) Coult. & Rose (PNF) Common name: Douglas waterhemlock. Toxin: alcohol (cicutoxin). Habitat: swampy or wet habitats along streams and in marshes. Animals affected: livestock, humans. Reference.: Alberts 1931, Kingsbury 1964. Conium tnaculatum L. (BIF) Common name: hemlock, poisonous hem- lock, spotted hemlock, California or Nebras- ka fern. Toxin: alkaloids (conine, N-methyl conine, conhydrine, lambda-coniceine, pseudoconhy- drine). Habitat: weed of roadsides, ditches, edges of cultivated fields and other waste areas. Animals affected: livestock, humans. Reference: Penny 1953, Kingsbury 1964. Daucus carota L. (BIF) Common name: wild ;parrot. Queen Ann's lace. Toxin: nitrates, under conditions of over- fertilization with too little water. Habitat: waste places. Animals affected: horses, cattle. Reference: Volker 1950, Kingsbury 1964. Pastinaca sativa L. var. sylvestris DC. (BIF) Common name: common parsnip. Toxin: unknown. September 1980 Brotherson et al.: Poisonous Plants 233 Habitat: widely naturalized weed. Animals affected: humans (dermatitis). Reference: S. L. Welsh (personal commu- nication, 1980). Slum suave Walt. (PNF) Common name: water parsnip, hemlock water parsnip. Toxin: unknown. Habitat: marshy lands and wet soils. Animals affected: hogs, cattle. Reference: Fyles 1920, Kingsbury 1964. Apocynaceae Apocynum androsaemifolium L. (PNF) Common name: spreading dogbane. Toxin: resins, glycosides. Habitat: common weed of open places, in coarse soils along streams, meadows, and wooded hillsides. Animals affected: cats, dogs. Reference: Moore 1909, Kingsbury 1964. Note: The closely related species A. me- dium Greene and A. sibericwn Jacq. niav show similar effects. Apocynum cannabinum L. (PNF) Common name: Indian hemp, dogbane, hemp dogbane. Toxin: resins, glycosides. Habitat: common weed of open places, in coarse soils along streams. Animals affected: cats, dogs. Reference: Finnemore 1909, Kingsbury 1964. Nerium oleander L. (PIS) Common name: oleander. Toxin: glycosides. Habitat: cultivated greenhouse plant, street plant in St. George. Animals affected: livestock, humans. Reference: West 1957, Kingsbury 1964. Asclepiadaceae Asclepias asperula (Decne) Woodson (PNF) Common name: asper milkweed, spider antelopehorn. Toxin: resinoids, glycosides and an alka- loid. Habitat: open dry soils, flats, desert swales, sandy or rocky hillsides with pinyon, juniper or oak. Animals affected: sheep, cattle, goats, horses, poultry. Reference: Huffman 1956, Kingsbury 1964. Asclepias fascicularis Decne ex DC. (PNF) Common name: Mexican whorled milk- weed. Toxin: resinoids, glycosides and an alka- loid. Habitat: dry hillsides and roadsides; pas- tures, moist streamsides. Animals affected: sheep, cattle, goats, horses, fowl. Reference: Schmutz et al. 1968. Asclepias incarnata L. (PNF) Common name: swamp milkweed. Toxin: resinoids, glycosides and an alka- loid. Habitat: marshes. Animals affected: sheep, cattle, horses, poultry. Reference: Hansen 1924, Kingsbury 1964. Asclepias labriformis Jones (PNF) Common name: labriform milkweed. Toxin: resinoids, glycosides and an alka- loid. Habitat: in sandy soils along old stream beds. Animals affected: sheep. Reference: Holmgren 1945, Kingsbury 1964. Asclepias latifolia (Torr.) Raf. (PNF) Common name: broadleaf milkweed. Toxin: resinoids, glycosides and an alka- loid. Habitat: dry plains in sandy soils. Animals affected: sheep. Reference: Schmutz et al. 1968, Kingsbury 1964, Shrift 1958. Asclepias speciosa Torr. (PNF) Common name: showy milkweed. Toxin: resinoids, glycosides, and an alka- loid. Habitat: prairies and open areas. Animals affected: sheep. Reference: Fleming 1920, Kingsbury 1964. Asclepias subverticillata (Gray) Vail (PNF) Common name: whorled milkweed, west- ern whorled milkweed. Toxin: resinoids, glycosides and an alka- loid. 234 Great Basin Naturalist Vol. 40. No. 3 Habitat: dry plains and foothills; spreads rapidlv along waterways and irrigation canals, forming dense stands; prefers sandy soils. .\niinals affected: sheep. Reference: Glover 1917, Kingsbiir\- 1964. .\steraceae Achillea miJlefolium L. iPNF) Common name: \arrow. To.xin: alkaloids and glycosides. Habitat: various. Animals affected: livestock. Reference: Hurst 1942, Kingsbury 1964. Ambrosia tomentosa Nutt. (PNF^ Common name: white ragweed, skeleton leaf bursage To.xin: nitrates, imder conditions of over- fertihzation with too little water. Habitat: dr\" plains, hills, waste ground and fields. Animals affected: livestock. Reference: Huffman 1956, Kingsbury 1964. Anthemis cotula L. (AIF) Common name: dog fennel, mayweed. ma\Aveed camomile. Toxin: acrid substance irritating to mucous membranes. Habitat: weedy plant of disturbed soils, fields and waste places; common weed in hay. .\nimals affected: poultry". Reference: Los Angeles Count}- Livestock Department 1938, Kingsbur>- 1964. Artemisia fiUfoUa Torr. (PNS^ Common name: sand sagebrush, old man sagebrush. Toxin: volatile oils. Habitat: sandy soils. Animals affected: horses. Reference: Beath 1953, Kingsbur>- 1964. Artemisia spinescens (DC.) Eaton PNS) Common name: bud sagebrush. Toxin: volatile oils. Habitat: dry plains and hills. Animals affected: hvestock. Reference: Sampson 1942, Kingsbur\- 1964. Aster chilensis Nees ssp. adscendens (Lindl.) Cronq, PNFi Common name: pacific aster. Toxin: secondary- selenium accumulator. Habitat: widely scattered in moist habitats. Animals affected: livestock. Reference: Trealease and Beath 1949. Kingsbury 1964. Aster glaucodes Blake i^PNF^ Common name: gray aster. Toxin: secondary- selenium accumulator. Habitat: moimtains. Animals affected: sheep. Reference: Trelease and Beath 1949, KingsbuT)" 1964. Aster laevis L. (PNF) Common name: smooth aster. Toxin: secondarv selenium accumulator. Habitat: widely scattered in drv to moist habitats. Animals affected: livestock. Reference: Trelease and Beath 1949. Kingsbur) 1964. Aster occidentalis (Nutt.) Torr. 6c Grav iPNF) Common name: western aster. To.xin: secondary- selenium acctunulator. Habitat: mountain meadows at moderate elevations. Animals affected: livestock. Reference: Trelease and Beath 1949, Kingsbury 1964. Aster pauciflorus Nutt. (PNF) Common name: fewhead aster. Toxin: secondary- selenium accumulator. Habitat: widespread in saline soils. Animals affected: livestock. Reference: J. D. Brotherson (personal com- munication. 19801 Bahia oppositifolia (Nutt.) DC. iPNF) Common name: bahia. plains bahia. Toxin: cvanogenetic glvcoside. Habitat: dr)' soils; plains and hills. Animals affected: cattle, sheep. Reference: Deem et al. 1939. Kingsburv 1964. Baileya midtiradiata Har\-. & Gray (BNF) Common name: desert baileya. cloth of gold, desert marigold. Toxin: unknown. Habitat: sandy and gravelly soils in dr\' areas. Animals affected: sheep, goats. Reference: Mathews 1933, Kingsburv 1964. September 1980 Brotherson et al.: Poisonous Plants 235 Baileya pleniradiata Har\ . & Gray (ANF) Common name: desert marigold baileya. Toxin: unknown. Habitat: mesas and deserts of southeastern Utah. Animals affected: sheep, goats. Reference: Schmutz et al. 1968. Centaurea repens L. (PIF) Common name: Russian knapweed. Toxin: unknown; produces nigropallidal encephalomalacia. Habitat: fields, roadsides and waste places. ^Animals affected: horses. Reference: Mecke 1979. Centaurea sohtitialis L. (AIF) Common name: yellow star thistle, vellow centaurea. Toxin: imknown; produces nigropallidal encephalomalacia; also mechanically in- jurious. Habitat: waste places, fields and roadsides. Animals affected: horses. Reference: Mettler and Stem 1963, Kings- bur>- 1964. Chrysothamnus nauseosus (Pall.) Britton (PXS) Common name: rubber rabbitbrush. Toxin: unknown. Habitat: dr\. open places at moderate and low elevations. Animals affected: livestock. Reference: Sampson 1942, Kingsbury 1964. Gritidelia squarrosa (Pursh) Dunal (BXF) Common name: gum weed, gumplane, cur- lycup gumweed. To.xin: secondarv selenium accumulator. Habitat: dr\ open places; prairies, plains, roadsides and fields. Animals affected: livestock. Reference: Trelease and DiSomma 1960, Kingsbury 1964. Heleiiium autumnaJe L. (PXF^ Common name: sneezeweed. bitterweed. Toxin: unknown acrid substance. Habitat: moist low ground in lowlands and foothills. Animals affected: sheep and cattle. Reference: Kingsbury 1964. Helenium hoopesii Gray (PNF) Common name: sneezeweed. orange sneezeweed. Toxin: glycoside (dugaldine). Habitat: high mountain slopes and vallevs, often forming dense stands in moist, sunnv, undisturbed localities. Animals affected: sheep and cattle. Reference: Marsh et al. 1921, Kingsbury 1964. Heleomeris longifolia Rob. & Greenm. var. annua (Jones) Yates (AXF) Common name: annual goldeneye, resin- weed, talloweed. Toxin: unknown. Habitat: ranges, hills, plains, and river bot- toms. Animals affected: cattle. Reference: Schmutz et al. 1968, Kingsbury 1964. Hymenoxys richardsonii (Hook.) Cockerel! iPXF Common name: pingue, Colorado rubber- weed, pingue hymeno-xys, rubberweed. Toxin: unknown; may be associated with mineral imbalance. Habitat: dry. rocky or clay soils of plains and mountain slopes from 1500 to 12,000 feet. Animals affected: sheep, cattle, goats. Reference: Aanes 1961, Kingsbury 1964. Oxytenia acerosa Xutt. PXS) Common name: copperweed; prickly oxy- tenia. Toxin: unknown. Habitat: alkaline soils in draws or stream- beds of desert ranges and foothills. Animals affected: cattle, sheep. Reference: Throp et al. 1940, Kingsburv 1964. Psathyrotes annua (Nutt.) Gray i.\XF) Common name: annual psathyrotes. Toxin: unknown. Habitat: dry, sandy, often alkaline soils, es- pecially of creek beds and dr)- washes. .\nimals affected: sheep. Reference: Binns et al. 1962, Kingsbur\ 1964. Psilostrophe sparsiflora (Gray) A. Nels. (PXF) Common name: greenstem paperflower. Toxin: unknown; induces kidney damage. Habitat: dr\-, open range. 236 Great Basin Naturalist Vol. 40, No. 3 Animals affected: sheep. Reference: Huffman 1956, Kingsbury 1964. Rudbeckia occidentalis Nutt. (PNF) Common name: western coneflower, nig- gerheads. Toxin: unknown. Habitat: streambanks and woodlands. Animals affected: generally unpalatable to livestock; affects hogs and sheep in feeding trials. Reference: Pammel 1911, Kingsbury 1964. Senecio integerrimus Nutt. (PNF) Common name: groundsel, senecio, lambs- tongue groimdsel. Toxin: alkaloids. Habitat: dry or moist open woods and slopes, from valleys to near timberline. x\nimals affected: livestock, humans (?). Reference: Clawson 1933, Kingsbury 1964. Senecio longilobus Benth. (PNF) Common name: wooly groundsel, thread- leaf groundsel. Toxin: pyrrolizidine alkaloids. Habitat: dry slopes, mesas and dry washes. Animals affected: cattle, horses, sheep, goats. Reference: Clawson 1933, Kingsbury 1964. Senecio spartioides Torr. & Gray (PNF) Common name: broom groundsel. Toxin: pyrrolizidine alkaloids. Habitat: valleys, plains; open areas and pine forests. Animals affected: cattle, horses, sheep, goats, humans(?). Reference: Clawson 1933, Kingsburv 1964, Davis 1958. Senecio vulgaris L. (AIF) Common name: common groundsel. Toxin: pyrrolizidine alkaloids. Habitat: weed of gardens and waste places. Animals affected: cattle, horses, sheep, goats, humans(?). Reference: Steyn 1934, Kingsbury 1964. Solidago parryi (Gray) Greene (PNF) Common name: Parry goldenweed. Toxin: unknown; causes milk-sickness or trembles. Habitat: mountains, coniferous forests. Animals affected: cattle. Reference: Schmutz et al. 1968, Kingsburv 1964. Tanacetum vulgare L. (PIF) Common name: common tansv. Toxin: abortifacient. Habitat: weed along roadsides, waste areas, ditchbanks and other moist areas. Animals affected: cattle, humans(?). Reference: Cress 1935, Kingsbury 1964. Tetradymia canescens DC. (PNS) Common name: spineless horsebrush, gray horsebrush. Toxin: photosensitizing compounds. Habitat: dry desert and sagebrush ranges. Animals affected: sheep. Reference: Kingsburv 1964, Schmutz et al. 1968. Tetradymia glabrata Gray (PNS). Common name: littleleaf horsebrush, spring rabbitbrush, coaloil brush. Toxin: photosensitizing compounds. Habitat: dry desert and sagebrush ranges. Animals affected: sheep. Reference: Kingsburv 1964, Fleming et al. 1922. Tetradymia ntittallii T. & G. (PNS) Common name: Nuttall horsebrush. Toxin: photosensitizing compounds. Habitat: dry desert and sagebaish ranges. Animals affected: sheep. Reference: Kingsbury 1964. Tetradymia spinosa T. and G. var. long- ispina Jones (PNS) Common name: longspine horsebrush. Toxin: photosensitizing compounds. Habitat: dry desert and sagebrush ranges. Animals affected: sheep. Reference: S. L. Welsh (personal commu- nication, 1980). Tetradymia spinosa T. and G. var. spinosa (PNS) Common name: spinv horsebrush. Toxin: photosensitizing compounds. Habitat: dry desert and sagebrush ranges. Animals affected: sheep. Reference: S. L. Welsh (personal commu- nication, 1980). Xanthium strumarium L. (AIF) Common name: spiny clotbur, spiny cock- lebur. September 1980 Brotherson et al.: Poisonous Plants 237 Toxin: hydroquinone. Habitat: fields and wastelands; along shores of ponds, rivers and in flood plains. .\nimals affected: livestock, fowl, hogs, hu- mans (dermatitis). Reference: Forrest 1938, Kuzel and Miller 1950, Kingsbury 1964. Xanthocephalum microcephahim (DC.) Gray (PNS) Common name: broomweed, perennial snakeweed, slinkweed, turpentine weed, threadleaf snakeweed, matchweed, resin- weed. Toxin: saponin. Habitat: dry stony plains, slopes and mesas. Animals affected: cattle, sheep, goats, swine, chicks, rabbits. Reference: Dollahite 1957, Kingsburv 1964. Xanthocephalum sarothrae (Pursh) Britt. and Rushy (A\F) Common name: broom snakeweed, snake- weed, matchbnish. Toxin: saponin. Habitat: dry stony plains, slopes and mesas. Animals affected: cattle, sheep, goats, swine, chicks, rabbits. Reference: Dollahite 1962, Kingsburv 1964. BORAGINACEAE Amsinckia intermedia Fisch, & Mey (AXF) Common name: tarweed, fiddleneck, fire- weed fiddleneck. Toxin: imknown; potentially lethal nitrate levels, pyrrolizidine alkaloids (?). Habitat: dry open cultivated ground or waste areas. Animals affected: horses, hogs, cattle. Reference: McCulloch 1940, Kingsburv 1964. Note: The closelv related species A. tesse- lata Gray and A. retrorsa Suksd. may show similar effects. Cynoglossiim officinale L. (BIF) Common name: houndstongue. Toxin: unknown. Habitat: waste places of plains and hills. Animals affected: livestock. Reference: Kingsbury 1964, S. L. Welsh (personal commimication, 1980). Brassicace.\e Brassica hirta Moench. (AIF) Common name: white mustard. Toxin: cyanogenetic glycoside. Habitat: cultivated weed, escaped to waste areas. Animals affected: cattle, sheep. Reference: Eaton 1941, Kingsbury 1964. Brassica kaber Wheeler (.\1F) Common name: charlock, wild mustard. Toxin: cyanogenetic glycoside. Habitat: common weed of grain crops and in waste areas. Animals affected: cattle, hogs, sheep. Reference: Thomson and Sifton 1922, Kingsbm-y 1964. Descurainia pinnata (Walt.) Britt. (ANF) Common name: tansv mustard, pinnate tansy mustard. Toxin: unknown. Habitat: heavy stands on dry, sandy soils. Animals affected: cattle. Reference: Hershev 1935, Kingsburv 1964. Erysimum cheiranthoides L. (ANF) Common name: wormweed mustard, treacle wallflower. Toxin: cvanogenetic glycoside. Habitat: weed of cultivation, roadsides, meadows; moist waste areas in valleys and canyons. Animals affected: hogs. Reference: Thomson and Sifton 1922, Kingsbur) 1964. Stanleya integrifolia James (PXS) Common name: wholeleaf desert prince's plume. Toxin: primarv selenium accumulator. Habitat: dry plains and hills. Animals affected: Not observed to be eaten bv livestock. ' Reference: Beath et al. 1953, Kingsbur\' 1964. Stanleya pinnata (Pursh) Britt. (PNS) Common name: prince's plume, desert prince's plume. Toxin: primary selenium accumulator. Habitat: desert soils, dry plains and mesas. Animals affected: experimental; normally unpalatable. Reference: Beath et al. 1953, Kingsbury 1964. 238 Great Basin Naturalist Vol. 40, No. 3 Stanletja viridiflora Nutt. (PNF) Common name: greenflower prince's plume. Toxin: primary selenium accumulator. Habitat: dry plains and hills. Animals affected: Not observed to be eaten by livestock. Reference: Beath et al. 1953, Kingsbury 1964. Thlaspi arvense L. (AIF) Common name: fanweed, field penny- cress. Toxin: cyanogenetic glycoside. Habitat: Common weed of cultivated and waste places. Animals affected: livestock. Reference: Thomson and Sifton 1922, Kingsbury 1964. C.wnabinaceae Cannabis sativa L. ssp. sativa (AIF) Common name: marijuana, hemp. Toxin: narcotic element contained in tetra- hydrocannabinol. Habitat: waste places. Animals affected: humans, livestock. Reference: Steyn 1934, Kingsbury 1964, Small and Cronquist 1976, Welsh 1980. Cannabis sativa L. ssp. indica (Lam.) Small & Cronq. (AIF) Common name: marijuana, hemp. Toxin: narcotic element contained in tetra- hydrocannabinol. Habitat: waste places. Animals affected: humans, livestock. Reference: Small and Cronquist 1976, Welsh 1980. Caprifoliaceae Sambucus coerulea Raf. (PNS) Common name: blue elderberry. Toxin: unknown. Habitat: moist soils of plains and hills. Animals affected: cattle, children (?) Reference: Schmutz et al. 1968. Sambucus racemosa L. (PNS) Common name: red elder. Toxin: unknown; concentrated in root. Habitat: moist forests, 7,500-10,000 ft. Animals affected: cattle, children(?) Reference: Schmutz et al. 1968. Caryophyllaceae Saponaria officinalis L. (PIF) Common name: bouncing bet, soapwort. Toxin: saponin. Habitat: fields, waste places; cultivated and escaping. Animals affected: sheep. Reference: Kingsbury 1964. Chenopodiaceae Atriplex gardneri Moq. (PNS) Common name: Nuttall saltbush, Gardner saltbush. Toxin: secondary selenium accumulator. Habitat: saline plains and hillsides. Animals affected: livestock. Reference: Fleming 1920, Kingsbury 1964. Bassia hyssopifolia (Pall.) Volk (AIF) Common name: fivehook bassia, smoth- erweed. Toxin: oxalates. Habitat: dry, saline soils. Animals affected: sheep. Reference: Pammel 1911. Chenopodium ambrosioides L. (AIF) Common name: wormseed goosefoot. Toxin: antihelminthic oil. Habitat: weed of waste places. Animals affected: geese, humans. Reference: Bamford 1951, Kingsbury 1964. Chenopodium album L. (AIF) Common name: lambsquarter. Toxin: nitrates, under conditions of over- fertilization with too little water. Habitat: weed of waste places. Animals affected: livestock. Reference: Case 1957, Kingsbury 1964. Chenopodium glaucum L. (AIF) Common name: oakleaf goosefoot. Toxin: nitrates, under conditions of over- fertilization with too little water. Habitat: weed of waste pmces. Animals affected: livestock. Reference: Case 1957, Kingsbury 1964. Halogeton glomeratus (Bieb.) C. A. Mey (AIF) Common name: halogeton, barilla. Toxin: oxalates. Habitat: dry saline plains and alkaline soils; roadsides. September 1980 Brotherson et al.: Poisonous Plants 239 Animals affected: livestock, especially sheep. Reference: Cook and Stoddart 1953, Kings- bury 1964. Kochia scoparia L. Schrad, (AIF) Common name: summer cypress, burning bush, Mexican fireweed. Belvedere summer cypress. Toxin: photosensitizing compounds. Habitat: dry soils, roadsides and waste places. Animals affected: cattle, sheep, horses. Reference: Rottgardt 1944, Kingsbury 1964, Schmutz et al. 1968. Salsola iberica Sennen & Pau (AIF) Common name: Russian thistle. Toxin: nitrates (?), possible oxalates (?). ' Habitat: dry soils of plains and foothills. Animals affected: livestock. Reference: Huffman et al. 1956. Sarcobatus vermiculutus (Hook.) Torr. (PNS) Common name: greasewood, black grease- wood. Toxin: oxalates. Habitat: dense stands confined to alkaline flats or saline soils of low and lower middle elevations. Animals affected: sheep, sometimes cattle. Reference: Kouch 1922, Kingsbury 1964. Cyperaceae Scirpus pungens Vahl (PNR) Common name: bulrush, three-square, American bulrush. Toxin: unknown; suspected of producing pulmonary emphysema. Habitat: wet or moist ground. Animals affected: cattle. Reference: Beath et al. 1953, Kingsburv 1964. Equisetaceae Equisetum arvense L. (PNF) Common name: horsetail, foxtail, rush, marsh horsetail. Toxin: alkaloids. Habitat: sandy or gravelly soils along streams and in moist fields and meadows. Animals affected: horses. Reference: Gussow 1912, Kingsbury 1964. Equisetum laevigatum A. Br. (PNF) Common name: smooth horsetail. Toxin: thiaminase. Habitat: marshes, alluvial thickets, sandy banks; weed of cultivation. Animals affected: horses. Reference: Samp.son and Malmsten 1942, Kingsbury 1964. Ericaceae Kalmia microphylla (Hook.) Heller (PNS) Common name: pale laurel, bog laurel, al- pine kalmia. Toxin: resinoids, andromedotoxin. Habitat: wet meadows and bogs of high elevation. Animals affected: sheep, calves, goats. Reference: Kingsbury 1964. Ledum glandulosum Nutt. (PNS) Common name: western Labrador tea. Toxin: resinoids, andromedotoxin. Habitat: wet meadows and bogs of high elevation. Animals affected: sheep, cattle. Reference: Kingsbury 1964. Euphorbiaceae Croton longipes Jones (PNF) Common name: croton. Toxin: croton oil (caustic). Habitat: roadsides, fields, and dry stream- beds; artemisia and pinyon belts. Animals affected: livestock. Reference: Schmutz et al. 1968, S. L. Welsh' (personal communication, 1980). Croton texensis (Klotz.) Muell. Arg. ex DC. (ANF) Common name: Texas croton. Toxin: croton oil (caustic). Habitat: roadsides, fields, and dry stream- beds; artemisia and pinyon belts. Animals affected: livestock. Reference: Volker 1950, Kingsbury 1964, Schmutz et al. 1968. Euphorbia cyparissias L. (PIF) Common name: cypress spurge, graveyard weed. Toxin: unknown acrid principle. Habitat: cultivated and escaping to road- sides and waste places. 240 Great Basin Naturalist Vol. 40, No. 3 Animals affected: cattle. Reference: Muenscher 1935, Kingsbury 1964. Euphorbia esula L. (PIF) Common name: leafy spurge. Toxin: unknown acrid principle. Habitat: field weed of roadsides and waste places. Animals affected: horses, sheep. Reference: Johnston and Peake 1960, Kingsbury 1964. Reverchonia arenaria Gray (ANF) Common name: reverchonia, sand rever- chonia. Toxin: unknown. Habitat: uncommon; plains and hillsides, sandy areas, Kane Co. Animals affected: sheep. Reference: Schmutz et al. 1968, Kingsbury 1964. Ricinus communis L. (AIF) Common name: castor bean. Toxin: ricin (a phytotoxin). Habitat: cultivated as an ornamental. Animals affected: livestock, humans. Reference: Clarke 1947, Kingsbury 1964. F.\BACEAE Acacia greggii Gray (PNT) Common name: catclaw acacia. Toxin: cyanogenetic glycoside. Habitat: plains and dry canyons; forms thickets along Beaver Dam Wash, Washing- ton Co. Animals affected: sheep. Reference: Schmutz et al. 1968, Kingsburv 1964. Astragalus asclepiadoides Jones (PNF) Common name: milkweed milkvetch. Toxin: primary selenium accumulator. Habitat: saline desert areas. Animals affected: livestock. Reference: Welsh 1978. Astragalus hisulcatus (Hook.) Gray (PNF) Common name: two-grooved poisonvetch, two-grooved milkvetch. Toxin: primary selenium accumulator. Habitat: plains and bottom lands, sage- brush zone. Animals affected: livestock. Reference: Welsh 1978, Kingsbury 1964. Astragalus convallarius Greene var. con- vallarius (PNF) Common name: timber poisonvetch, lesser rushy milkvetch. Toxin: produces locoism. Habitat: dry hillsides, desert shrub to lower montane zones. Animals affected: livestock. Reference: Muenscher 1951, Kingsbury 1964. Astragalus drummondii Dougl. ex Hook. (PNF) Common name: Drummond milkvetch. Toxin: produces locoism. Habitat: plains and hillsides, brushy places. Animals affected: livestock. Reference: Welsh 1978, Kingsbury 1964. Astragalus eastwoodae Jones (PNF) Common name: Eastwood poisonvetch, Eastwood milkvetch. Toxin: primary selenium accumulator. Habitat: dry hillsides. Animals affected: livestock. Reference: Welsh 1978, Kingsbury 1964. Astragalus flavus Nutt. ex Torr. & Gray var. argillosus (Jones) Barneby (PNF) Common name: yellow milkvetch. Toxin: primary selenium accumulator. Habitat: dry plains and hillsides, salt desert areas. Animals affected: livestock. Reference: Welsh 1978. Astragalus flavus Nutt. ex Torr. & Gray var. candicans Gray (PNF) Common name: Canada yellow milkvetch. Toxin: primary selenium accumulator. Habitat: dry plains and hillsides, shales and clays of southern Utah. Animals affected: livestock. Reference: Welsh 1978. Astragalus flavus Nutt. ex Torr. & Gray var. flavus (PNF) Common name: yellow milkvetch. Toxin: primary selenium accumulator. Habitat: dry plains and hillsides, saline silts and clays in saline desert areas, south-central Utah. Animals affected: livestock. Reference: Welsh 1978, Kingsbury 1964. September 1980 Brotherson et al.: Poisonous Plants 241 Astragalus iselyi Welsh (PNF) Common name: Isely milkvetch. Toxin: primary selenium accumulator. Habitat: dry hillsides, salt desert areas. Animals affected: livestock. Reference: Welsh 1978. Astragalus lentiginosus Dougl. ex Hook. var. araneosus (Sheld.) Barneby (PNF) Common name: spider locoweed, cobweed milkvetch. Toxin: produces locoism. Habitat: dry hillsides in sagebrush. Animals affected: livestock. Reference: Welsh 1978, Kingsbury 1964. Astragalus lentiginosus Dougl. ex Hook. var. palans (Jones) Jones (PNF) Common name: straggling milkvetch. Toxin: produces locoism. Habitat: salt desert areas, dry hillsides and canyons, mixed desert shrub communities. Animals affected: livestock. Reference: Welsh 1978, S. L. Welsh (per- sonal communication, 1980), Kingsbury 1964. Astragalus lentiginosus Dougl. ex Hook. var. wahweapensis Welsh (PNF) Common name: Wahweap loco, Wahweap milkvetch. Toxin: produces locoism. Habitat: dry hillsides in sagebrush, sandy soils, Kane Co. Animals affected: livestock, esp. horses. Reference: Welsh 1978, Kingsbury 1964. Astragalus miser Dougl. ex Hook. var. ob- longifolius (Rydb.) Cronq. (PNF) Common name: timber milkvetch, Ryd- berg weedv milkvetch. Toxin: miserotoxin. Habitat: widely scattered in lower mon- tane zones. Animals affected: livestock. Reference: Williams 1969, Welsh 1978. Astragalus moencoppensis Jones (PNF) Common name: Moenkopi poisonvetch, Moenkopi milkvetch. Toxin: primary selenium accumulator. Habitat: heavy soils, salt desert through pinyon-juniper areas. Animals affected: livestock. Reference: Welsh 1978, Kingsbury 1964. Astragalus mollissimus Torr. (PNF) Common name: Thompson woolly loco- weed. Toxin: produces locoism. Habitat: dry plains and hillsides. Animals affected: livestock. Reference: Welsh 1978, Kingsbury 1964. Astragalus pattersonii Gray ex Brand. (PNF) Common name: Patterson locoweed, Pat- terson milkvetch. Toxin: primary selenium accumulator. Habitat: dry plains and hillsides. Animals affected: livestock. Reference: Welsh 1978, Kingsbury 1964. Astragalus praelongus Sheldon (PNF) Common name: stinking milkvetch. Toxin: primary selenium accumulator. Habitat: dry plains and hillsides, clay and seleniferous soils. Animals affected: livestock. Reference: Welsh 1978. Astragalus preussii Gray (PNF) Common name: Preuss milkvetch. Toxin: primarv selenium accumulator. Habitat: dry plains and hillsides, seleni- ferous clays and silts. Animals affected: livestock. Reference: Welsh 1978, Kingsbury 1964. Astragalus pubentissimus Torr. & Gray (PNF) Common name: Green River locoweed. Green River milkvetch. Toxin: produces locoism. Habitat: canyons, mountainsides. Animals affected: livestock, mainly sheep. Reference: Buck 1961, Kingsbury 1964. Astragalus racemosus Pursh var. treleasi Por- ter (PNF) Common name: alkali milkvetch. Toxin: primary selenium accumulator, causes "alkali disease" or "blind staggers." Habitat: Uinta and Duchesne River forma- tions. Animals affected: cattle. Reference: Welsh 1978. Astragalus rafaelensis Jones (PNF) Common name: San Rafael milkvetch. Toxin: primary selenium accumulator. Habitat: seleniferous clays and silts, salt desert shrub zones, Emery Co. 242 Great Basin Naturalist Vol. 40, No. 3 Animals affected: livestock. Reference: Welsh 1978. Astragalus sabuhsus Jones (PNF) Common name: straightstem poisonvetch, Cisco milkvetch. Toxin: primary selenium accumulator. Habitat: salt desert shrub zone on shales. Animals affected: livestock. Reference: Welsh 1978, Marsh 1935. Astragalus saurinus Barneby (PNF) Common name: dinosaur milkvetch. Toxin: primary selenium accumulator. Habitat: salt desert shrub and pinyon-juni- per zone, Uintah Co. Animals affected: livestock. Reference: Welsh 1978. Astragalus tetrapterus Gray (PNF) Common name: fourwing poisonvetch, four-wing milkvetch. Toxin: produces locoism. Habitat: plains, dry hillsides. Animals affected: cattle, sheep. Reference: Marsh 1920, Welsh 1978. Astragalus toanus Jones (PNF) Common name: Toano milkvetch. Toxin: primary selenium accumulator. Habitat: clay soils, salt desert shrub zone. Animals affected: livestock. Reference: Welsh 1978, Kingsbury 1964. Astragalus woodruffii Jones (PNF) Common name: Woodruff milkvetch. Toxin: primary selenium accumulator. Habitat: desert shnib zone on sandy and sandy-silt soils. Animals affected: livestock. Reference: Welsh 1978. Lathyrus latifolius L. (PIF) Common name: perennial peavine, per- ennial sweet pea. Toxin: alpha, gamma-diaminobutyric acid. Habitat: rangeland. Animals affected: rats (experimental), sheep (lathyrism). Reference: Lewis 1949, Kingsbury 1964, S. L. Welsh (personal commvmication, 1980). Lathyrus sylvestris L. (PNF) Common name: everlasting sweetpea, flat pea. Toxin: alpha, gamma-diaminobutyric acid. Habitat: rangeland. Animals affected: sheep, rats (experimen- tal). Reference: Lewis 1948, Kingsbury 1964. Lupinus alpestris A. Nels. (PNF) Common name: mountain lupine. Toxin: alkaloids. Habitat: mountains. Animals affected: cattle, sheep. Reference: Beath et al. 1953, Kingsbury 1964. Lupinus arbustus Dougl. ex Lindl. var. cal- caratus (Kellogg) Welsh (PNF) Common name: Douglas spurred lupine, spur lupine. Toxin: alkaloids. Habitat: hillsides, dry soils. Animals affected: cattle, horses, sheep. Reference: Clawson 1931, Kingsbury 1964. Lupinus argenteus Pursh (PNF) Common name: silvery lupine. Toxin: alkaloids. Habitat: dry flats and slopes in woods, plains and hills. Animals affected: sheep, cattle, horses, goats, hogs, deer. Reference: Marsh 1916, Kingsbury 1964. Lupinus caudatus Kell. (PNF) Common name: Kellogg's spurred lupine, tailcut lupine. Toxin: alkaloids. Habitat: exposed hillsides. Animals affected: cattle. Reference: Animal Disease and Parasite Research Division USDA 1958, Kingsbury 1964. Lupinus leucophyllus Dougl. (PNF) Common name: woolly-leaved lupine, vel- vet lupine. Toxin: alkaloids. Habitat: dry soil, foothills. Animals affected: sheep, cattle, horses, goats, hogs, deer. Reference: Marsh 1916, Kingsbury 1964. Lupinus pusillus Pursh (ANF) Common name: low lupine, rusty lupine. Toxin: alkaloids. Habitat: dry plains, foothills. Animals affected: sheep. Reference: Sampson 1942, Kingsbury 1964. September 1980 Brotherson et al.: Poisonous Plants 243 Lupinus sericeus Pursh (PNF) Common name: silky lupine. Toxin: alkaloids. Habitat: dry hillsides and valleys. Animals affected: sheep, cattle, horses, goats, hogs, deer. Reference: Binns and James 1961, Kings- bury 1964. Medicago sativa L. (PIF) Common name: alfalfa, lucerne. Toxin: saponin. Habitat: cultivated and escaping. Animals affected: cattle, chicks, hogs, sheep. Reference: Walter 1954, Kingsbury 1964. Melilotus alba Desr. (BIF) Common name: white sweetclover. Toxin: dicoumarin. Habitat: waste places and fields, escaped from cultivation. Animals affected: cattle. Reference: Roderick 1931, Kingsbury 1964. Melilotus officinalis L. Lam. (BIF) Common name: yellow sweetclover. Toxin: dicoumarin. Habitat: waste ground, fields; used for for- age and fertilizer. Animals affected: cattle, sheep, horses. Reference: Roderick 1931, Kingsbury 1964. Oxytropis lambertii Pursh (PNF) Common name: white loco, white paint loco, silky crazyweed. Toxin: unknown alkaloids; produces locoism. Habitat: prairies and mountains, usually in drier areas, lower to middle elevations. Animals affected: livestock. Reference: Couch 1929, Kingsbury 1964. Oxytropis sericea Nutt. ex Torr. & Gray (PNF) Common name: white paint loco, silky cra- zyweed. Toxin: alkaloids; produces locoism. Habitat: open gravelly or well-drained slopes and hills at lower to middle elevations. Animals affected: livestock. Reference: Porter 1951, Kingsbury 1964. Visum sativum L. (AIF) Common name: garden pea. Toxin: unknown. Habitat: cultivated. Animals affected: sheep, cattle. Reference: Whiting et al. 1957, Kingsbury 1964. Poinciana gilliesii Hook. (PIS) Common name: bird of paradise. Toxin: unknown; green seed pods are gas- trointestinal irritants. Habitat: cultivated ornamental, small pop- ulation established in Washington Co. Animals affected: humans, livestock. Reference: Cann and Verhulst 1958, Kings- bury 1964. Prosopis glandulosa Torr. (PNT) Common name: mesquite. Toxin: unknown; may cause a nutritional deficiency. Habitat: dry ranges, washes and draws at low elevations, especially along streams where the water table is high. Animals affected: cattle. Reference: Adler 1949, Kingsbury 1964. Robinia pseudoacacia L. (PIT) Common name: black locust. Toxin: unknown. Habitat: escaped from cultivation; around dwellings or along fencerows. Animals affected: horses, cattle, sheep, poultry, humans. Reference: Power 1901, Kingsbury 1964. Thermopsis montana Nutt. (PNF) Common name: goldenpea, mountain ther- mopsis, yellow pea. Toxin: alkaloids. Habitat: common in pastures. Affected animals: cattle. Reference: Schmutz et al. 1968. Trifolium hybridum L. (PIF) Common name: .\lsike clover. Toxin: photosensitizing compound. Habitat: cultivated; escaped to roadsides and meadows. Animals affected: horses, hogs, sheep, cattle. Reference: Fincher and Fuller 1942, Kings- bury 1964. 244 Great Basin Naturalist Vol. 40, No. 3 Trifolium praetense L. (PIF) Common name: red clover. Toxin: unknown. Habitat: cultivated and escaping along roadsides and ditches. Animals affected: cattle, horses, sheep. Reference: O'Dell 1959, Kingsbury 1964. Trifolium repens L. (PIF) Common name: white clover. Toxin: cyanogenetic. Habitat: cultivated and escaping. Animals affected: newborn pigs. Reference: Garner 1957, Kingsbury 1964. Vicia villosa Roth (PIF) Common name: hairy vetch, winter vetch. Toxin: photosensitizing compound. Habitat: cultivated; occasionally escaping. Animals affected: cattle, horses. Reference: Claughton and Claughton 1954, Kingsbury 1964. Haemodoraceae Iris missouriensis Nutt. (PNF) Common name: wild iris, blue flag, fleur- de-lis, western blue flag. Toxin: unknown. Habitat: moist soils along stream banks, in marshes or moist mountain meadows. Animals affected: calves, laboratory ani- mals. Reference: Beath et al. 1953, Kingsbury 1964. Hypericaceae Hypericum formosum H.B.K. (PNF) Common name: southwestern St. Johns- wort. Toxin: photosensitizing compounds. Habitat: moist soils of plains and hills. Animals affected: cattle, sheep, horses, goats. Reference: Harris 1951. Fagaceae Quercus gambelii Nutt. (PNT) Common name: Gambel oak. Toxin: tannins. Habitat: throughout the state, often form- ing dense thickets. Animals affected: cattle, sheep, and goats. Reference: Boughton 194.3, Kingsbury 1964. FUMARIACEAE Corydalis aurea Willd. (ANF) Common name: Golden corydalis. Toxin: alkaloids. Habitat: woods and well-shaded mountain slopes. Animals affected: sheep, cattle. Reference: Sperry 1955, Kingsbury 1964. Gentianaceae Centaurium calycosum (Buckl.) Fern. (ANF) Common name: Buckley centaury, moun- tain pink, arizona centaury. Toxin: unknown. Habitat: moist soil, river valleys. Animals affected: sheep, goats. Reference: Dollahite and Allen 1962, Kingsbury 1964. JUNCAGINACEAE Triglochin concinna Davy (PNF) Common name: arrowgrass, goosegrass, sourgrass, podgrass, Utah arrowgrass. Toxin: hydrocyanic acid. Habitat: salty marshes and ponds. Animals affected: sheep, cattle. Reference: Schmutz et al. 1968. Triglochin debilis L. (PNF) Common name: arrowgrass, weak arrow- grass. Toxin: hydrocyanic acid. Habitat: damp soils, marshes and sloughs; usually where the soil is alkaline or the water calcareous or brackish. Animals affected: sheep, cattle. Reference: Schmutz et al. 1968, Kingsbury 1964. Triglochin maritima L. (PNF) Common name: arrowgrass, seashore ar- rowgrass, shore arrowgrass. Toxin: hydrocyanic acid. Habitat: damp soils, marshes and sloughs; usually where the soil is alkaline or the water calcareous or brackish. Animals affected: sheep, cattle. Reference: Beath et al. 1933, Kingsbury 1964. September 1980 Brotherson et al.: Poisonous Plants 245 Lamiaceae Lamium amplexicaule L. (AIF) Common name: henbit, dead nettle. Toxin: unknown. Habitat: occasional weed of fields and waste places. Animals affected: sheep, horses, cattle. Reference: Hurst 1942, Kingsbury 1964. LiLIACEAE Allium schoenoprasm L. (PNF) Common name: chives. Toxin: unknown. Habitat: cultivated; river bars, lake shores, wet meadows. Animals affected: horses. Reference: Kobayashi 1950, Kingsbury 1964. Asparagus officinalis L. (PIF) Common name: asparagus. Toxin: unknown. Habitat: cultivated and widely escaped. Animals affected: cattle and dairy cows. Reference: Los Angeles County Livestock Department 1938, Kingsbury 1964. Ornithogalum umbellatum L, (PIF) Common name: star-of-Bethlehem, snow- drop. Toxin: cholchicine alkaloid. Habitat: weed of grasslands and thickets. Animals affected: sheep, cattle, children. Reference: Reynard and Norton 1942, Kingsbury 1964. Veratrum californicum Durand (PNF) Common name: false hellebore, corn-lily, skunk cabbage. Toxin: alkaloids. Habitat: bogs and wet meadows from 7500 to 9500 feet. Animals affected: cattle, sheep, fowl, hu- mans. Reference: Schmutz et al. 1968, Kingsbury 1964. Yucca L. (PNS) Note: members of this genus have been re- ported to contain saponins, salicylic acid, the alkaloid imperialin, and several resins. None of the species in which these toxins have been identified are found in Utah. Reference: Pammel 1911. Zigadenus elegans Pursh (PNF) Common name: death camas, mountain death camas. Toxin: alkaloids. Habitat: prairies, meadows. Animals affected: cattle, horses, hoes, fowl humans. Reference: Marsh et al. 1915, Kingsbury 1964. Zigadenus paniculatus (Nutt.) Wats. (PNF) Common name: death camas, foothill death camas, sandcom. Toxin: alkaloids. Habitat: dry soils; hills and plains. Animals affected: cattle, horses, hogs, hu- mans. Reference: Fleming et al. 1921, Kingsbury 1964. Zigadenus venenosus Wats. (PNF) Common name: death camas, meadow death camas. Toxin: alkaloids. Habitat: moist, grassy meadows. Animals affected: sheep, cattle, horses, hogs, humans. Reference: Cameron 1952, Kingsbury 1964. PiNACEAE Pinus ponderosa Doug, ex Laws (PNT) Common name: western yellow pine, pon- derosa pine. Toxin: unknown. Habitat: coniferous forest at moderate ele- vations; dry hillsides, plateaus, slopes, valleys and mesas. Animals affected: cattle. Reference: MacDonald 1952, Kingsbury 1964. POACEAE Avena fatua L. (AIG) Common name: wild oats. Toxin: mechanically injurious. Habitat: cultivated land and waste places. Animals affected: livestock. Reference: Pammel 1911. Avena sativa L. (AIG) Common name: cultivated oats. Toxin: nitrates, photosensitizing com- pounds, grass tetany. 246 Great Basin Naturalist Vol. 40, No. 3 Habitat: open ground, grasslands, waste places; lawns and golf courses. Animals affected: cattle, horses, hogs, tur- keys, goats, sheep and wild ruminants. Reference: Newsom et al. 1937, Kingsbury 1964. Bromus rigidus Roth (AIG) Common name: ripgut brome. Toxin: mechanical injury from mature awns. Habitat: common weed. Animals affected: cattle and sheep. Reference: Range Plant Handbook 1937. Bromus rubens L. (AIG) Common name: foxtail chess, red brome. Toxin: mechanical injury from mature awns. Habitat: common weed, dry and saline soils. Animals affected: cattle and sheep. Reference: Davis 1952. Bromus tectorum L. (AIG) Common name: cheatgrass, downy cheat. Toxin: mechanical injury from mature awns; may also be implicated in ergot poi- soning. Habitat: common weed, especially in dry places; plains and foothills. Animals affected: cattle. Reference: Pammel 1911. Cynodon dactylon L. Pers. (PIG) Common name: Bermuda grass. Toxin: photosensitizing compound. Habitat: open ground, grasslands, waste places; lawns and golf courses. Animals affected: cattle. Reference: Gibbons 1953, Kingsbury 1964. Eragrostis cilianensis (All.) Link (AIG) Common name: lovegrass, stinkgrass, stick grass. Toxin: unknown. Habitat: cultivated ground, gardens and waste places; weed in fields and along road- sides. Animals affected: horses. Reference: Gates 1930, Kingsbury 1964. Festuca arundinacea Schreb. (PIG) Common name: fescue, tall fescue, alta fes- cue, goar fescue. Toxin: alkaloids. Habitat: unimproved pastures; wet, heavy soils of high organic content. Animals affected: cattle. Reference: Maag and Tobiska 1956, Kings- bury 1964. Glyceria striata Lam. Hitch. (PNG) Common name: fowl mannagrass. Toxin: cyanogenetic. Habitat: wet areas. Animals affected: cattle. Reference: Reynard and Norton 1942, Kingsbury 1964. Hilaria rigida (Thurb.) Benth ex Scribn. (PNG) Common name: galleta grass, big galleta, dixie grass. Toxin: unknown. Habitat: dry lands and desert ranges to 4000 feet. Animals affected: cattle. Reference: Quortrup and McFarland 1956, Kingsbury 1964. Holcus lanatus L. (PIG) Common name: velvet grass, mesquite grass, Yorkshire velvet grass. Toxin: unknown. Habitat: open ground, meadows and moist places; occasionally cultivated. Animals affected: livestock. Reference: Couch 1932, Kingsbury 1964. Hordeum jubatum L. (PNG) Common name: squirreltail grass, foxtail grass, wild barley. Toxin: mechanically injurious. Habitat: weed in open ground, meadows, prairies, along streams, ditches and waste places. Animals affected: sheep, cattle, horses. Reference: Fleming and Peterson 1919, Kingsbury 1964. Hordeum vulgare L. (AIG) Common name: cultivated barley. Toxin: mechanically injurious. Habitat: cultivated for grain and along road shoulders; sometimes spontaneous in waste places but not persistent. Animals affected: hogs, dogs, humans, poultry. Reference: Christensen and Kernkamp 1936, Kingsbury 1964. September 1980 Brotherson et al.: Poisonous Plants 247 Setaria lutescens (Wiegel) Hubb. (AIG) Common name: yellow bristle grass, foxtail grass, pigeon grass. Toxin: mechanically injurious. Habitat: common weed of cultivated and waste areas. Animals affected: livestock. Reference: Bankowski et al. 1956, Kings- bury 1964. Sorghum halpense L. Pers. (PIG) Common name: Johnson grass. Toxin: hydrocyanic acid, nitrates. Habitat: weed of cultivated fields, waste places and along irrigation ditches and stream bottoms. Animals affected: cattle, sheep, horses. Reference: Slade 1903, Kingsbury 1964. Sorghum vulgare Pers. (AIG) Common name: grain sorghum. Toxin: cyanogenetic glycoside. Habitat: cultivated. Animals affected: cattle, sheep, horses. Reference: Slade 1903, Kingsbury 1964. Stipa commata Trin. & Rupr. (PNG) Common name: needle-and-thread grass. Toxin: mechanically injurious. Habitat: dry plains and hillsides, sandy soil. Animals affected: livestock. Reference: Pammel 1911. Stipa neomexicana (Thurb.) Scribn. (PNG) Common name: New Mexican feather- grass. Toxin: mechanically injurious. Habitat: common in dry rocky canyons and mesas. Animals affected: livestock. Reference: Pammel 1911. Zea mays L. (AIG) Common name: com, maize. Toxin: nitrates, under conditions of over- fertilization with too little water. Habitat: cultivated for grain, forage or si- lage. Animals affected: livestock, humans. Reference: Brady et al. 1955, Kingsbury 1964. POLYGONACEAE Beta vulgaris L. (AIF) Common name: beet, sugar beet, fodder beet, mangel-worzel, mangold. Toxin: oxalates; nitrates, under conditions of overfertilization with too little water. Habitat: cultivated. Animals affected: livestock. Reference: Baker and Eden 1954, Kings- bury 1964. Rheum rhaponticum L. (PIF) Common name: rhubarb. Toxin: oxalic acid, oxalates. Habitat: cultivated and persisting. Animals affected: livestock, humans. Reference: Hansen 1930, Kingsbury 1964. Rumex acetosella L. (PIF) Common name: sheep sorrel, dock. Toxin: oxalates. Habitat: common weed of acid or sterile, gravelly soils of pastures and meadows; waste places. Animals affected: sheep. Reference: Connor and Adams 1951, Kingsbury 1964. Rumex crispus L. (PIF) Common name: curly dock. Toxin: oxalates. Habitat: moist fields and waste places. Animals affected: sheep. Reference: Connor and Adams 1951, Kingsbury 1964. POLYPODIACEAE Dryopteris felix-mas (L.) Schrott (PNF) Common name: male fern. Toxin: thiaminase. Habitat: mountains; damp soils, deep shaded ravines, in cliffs or tallus. Animals affected: horses. Reference: Harvey et al. 1944, Kingsbury 1964. Pteridium aquilinum (L.) Kuhn (PNF) Common name: Bracken fern. Toxin: thiaminase and its coenzymes. Habitat: upland pastures, a.spen zone. Animals affected: livestock. Reference: Carpenter 1950, Kingsbury 1964. PORTULACACEAE Portulaca oleracea L. (AIF) Common name: purslane, pusley. Toxin: oxalates. 248 Great Basin Naturalist Vol. 40, No. 3 Habitat: common weed of garden and cul- tivated areas. Animals affected: sheep. Reference: Mathams and Sutherland 1952, Kingsbury 1964. Ranunculaceae Aconitum columbianum Nutt. (PNF) Common name: western monkshood. Toxin: alkaloids. Habitat: mountains from 5000 to 10,000 feet; along streams and wet meadows; moist places and thickets. Animals affected: livestock, humans. Reference: Stern 1960, Kingsbury 1964. Actaea arguta Nutt. (PNF) Common name: baneberry, western bane- berry. Toxin: irritant oil. Habitat: mountains in rich, moist soil. Animals affected: livestock, humans. Reference: Schmutz et al. 1968. Caltha leptosepala DC. (PNF) Common name: elkslip marshmarigold. Toxin: glycoside (protoanemonin). Habitat: wet mountain soils. Animals affected: livestock. Reference: Schmutz et al. 1968. Delphinium andersonii Gray (PNF) Common name: Anderson larkspur. Toxin: alkaloids. Habitat: subsaline soils of plains and hills. Animals affected: cattle, sheep. Reference: Miller 1923, Kingsbury 1964. Delphinium barbeyi Huth. (PNF) Common name: barbey, larkspur, tall larkspur. Toxin: alkaloids. Habitat: mountains; meadows and open woods, summer ranges; common under aspen and along streams. Animals affected: cattle, occasionally sheep. Reference: Cook 1951, Kingsbury 1964. Delphinium nuttallianum Fritz. (PNF) Common name: Nuttall larkspur, low larkspur. Nelson larkspur. Toxin: alkaloids. Habitat: moist soils, hills, foothills, and sagebrush deserts. Animals affected: cattle, sheep. Reference: Ewan 1945, Kingsbury 1964. Delphinium occidentale Wats. (PNF) Common name: duncecap larkspur, tall larkspur. Toxin: alkaloids. Habitat: mountain summer ranges, com- mon under aspen and along streams; moun- tain meadows. Animals affected: cattle, occasionally sheep. Reference: Couch 1936, Kingsbury 1964. Ranunculus acris L. (PIF) Common name: tall field buttercup, tall buttercup. Toxin: protoanemonin. Habitat: common pasture weed. Animals affected: livestock. Reference: Tehon et al. 1946, Kingsbury 1964. Ranunculus cymbalaria Pursh (PNF) Common name: alkali buttercup; trailing buttercup. Toxin: glycosides (protoanemonin). Habitat: muddy banks along brackish streams and marshes. Animals affected: livestock. Reference: Fleming 1920, Kingsbury 1964. Ranunculus flammula var. filiformis (Michx.) Hook. (PNF) Common name: creeping spearwort but- tercup. Toxin: glycosides (protoanemonin). Habitat: marshy ground of lakes, streams and ditches. Animals affected: livestock. Reference: Hill and van Heyningen 1951, Kingsbury 1964. Ranunculus repens L. (PIF) Common name: creeping buttercup. Toxin: protoanemonin. Habitat: meadows and marshes at lower elevations; wet pastures. Animals affected: livestock. Reference: Gilkey 1958, Kingsbury 1964. Ranunculus scleratus L. (PNF) Common name: celeryleaf crowfoot. Toxin: glycosides (protoanemonin). Habitat: borders of lakes, ponds and streams. September 1980 Brotherson et al.: Poisonous Plants 249 Animals affected: livestock. Reference: Fleming 1920, Kingsbury 1964. Ranunculus testiculatus Crantz (AIF) Common name: burbuttercup; testiculate buttercup. Toxin: glycosides (protoanemonin). Habitat: general in the intermountain re- gion. Animals affected: livestock. Reference: Schmutz et al. 1968. ROSACEAE Cercocarpus montanus Raf. (PNS) Common name: mountain mahogany, birchleaf mountain mahogany, true mountain mahogany. Toxin: cyanogenetic glycoside. Habitat: stony hills and slopes. Animals affected: livestock. Reference: Burke 1960, Kingsbury 1964. Prunus armeniaca L. (PIT) Common name: apricot. Toxin: cyanide. Habitat: cultivated and persisting. Animals affected: livestock, humans. Reference: Hurst 1942, Kingsbury 1964. Prunus persica Batsch. (PNT) Common name: peach. Toxin: cyanide. Habitat: cultivated. Animals affected: livestock. Reference: Reynard and Norton 1942, Kingsbury 1964. Prunus virginiana L. (PNT) Common name: choke cherry. Toxin: cyanogenetic glycoside. Habitat: common in hills, mountains, along streams, thickets, fencerows and edges of woods. Animals affected: sheep, cattle. Reference: Pijoan 1942, Kingsbury 1964. Santalaceae Comandra umbellata (L.) Nutt. (PNF) Common name: bastard toadflax. Toxin: alkaloids, glycosides, secondary se- lenium accumulator. Habitat: common weed, found in various habitats. Animals affected: livestock. Reference: Trelease and Beath 1949, Kingsbury 1964. SOLANACEAE Datura meteloides Dunal (ANF) Common name: datura, stramonium, thornapple, Jimson weed, Jamestown weed, apple of Peru, tolgaudia, sacred datura, In- dian apple. Toxin: alkaloids (atropine, hyoscvamine, hyoscine). Habitat: plains, dry hills and valleys; culti- vated and escaping. Animals affected: humans, horses, cattle, sheep, hogs, mules, chickens. Reference: Hansen 1924, Kingsbury 1964. Datura stramonium L. (ANF) Common name: Jimsonweed, sacred da- tura. Toxin: alkaloids (atropine, hyoscyamine, hyoscine). Habitat: waste areas, rich soils of barn- yards, heavily used portions of pastures. Animals affected: horses, cattle, sheep, hogs, mules, chickens, humans. Reference: Hansen 1924, Kingsbury 1964. Hyoscyamus niger L. (BIF) Common name: black henbane, henbane. Toxin: alkaloids (hyoscyamine, hyoscine, atropine). Habitat: widespread dry soils of roadsides and waste areas. Animals affected: humans, fowl, livestock. Reference: Long 1917, Kingsbury 1964. Lycium halmifolium Mill. (PIS) Common name: matrimony vine, tea vine. Toxin: unknown. Habitat: cultivated and escaping around homesites and cemetaries. Animals affected: calves, sheep. Reference: Hansen 1927, Kingsbury 1964. Nicotiana attenuata Torr. ex S. Wats. (.\NF) Common name: wild tobacco, coyote to- bacco. Toxin: nicotine. Habitat: dry, sandy stream beds and flats. Animals affected: horses, pigs, livestock, humans. Reference: Marsh et al. 1927, Kingsbury 1964. 250 Great Basin Naturalist Vol. 40, No. 3 Nicotiana trigonophylla Dunal ex DC. (ANF) Common name: wild tobacco, desert to- bacco. Toxin: nicotine. Habitat: dry desert soils. Animals affected: horses, pigs, livestock, humans. Reference: Marsh et al. 1927, Kingsbury 1964. Solarium dulcamara L. (PIF) Common name: European bittersweet, climbing nightshade, bitter nightshade. Toxin: glycoalkaloids. Habitat: woods, thickets and waste places; cultivated and escaping. Animals affected: cattle, horses, sheep, hu- mans. Reference: Craig and Kehoe 1925, Kings- bury 1964. Solanum eleagnifolium Cav. (PNF) Common name: silverleaf nightshade, white horsenettle, trapillo. Toxin: glycoalkaloids. Habitat: serious weed of prairies, open woods and disturbed soils; dry ground; barn- yards. Animals affected: cattle, sheep. Reference: Buck et al. 1960, Kingsbury 1964. Solanum nigrum L. (AIF) Common name: black nightshade. Toxin: glycoalkaloids. Habitat: common weed of fields and waste places. Animals affected: livestock, humans. Reference: Carey 1955, Kingsbury 1964. Solanum rostratum Dunal (ANF) Common name: buffalo bur, Kansas thistle, Texas thistle, buffalobur nightshade. Toxin: glycoalkaloids. Habitat: plains, roadsides, barnyards. Animals affected: hogs. Reference: Simic 1943, Kingsbury 1964. Solanum triflorum Nutt. (ANF) Common name: three flowered nightshade, cutleaf nightshade. Toxin: glycosides. Habitat: prairies, fields and waste places; weed of cultivation and disturbed soils. Animals affected: horses, cattle. Reference: Pammel 1921, Kingsbury 1964. Typhaceae Typha latifolia L. (PNF) Common name: cattail. Toxin: unknown. Habitat: common in moist soils, marshes and ponds. Animals affected: horses. Reference: Hansen 1930, Kingsbury 1964. Zygophyllaceae Tribulus terrestris L. (AIF) Common name: puncture vine, caltrap. Toxin: nitrates, photosensitizing com- pound. Habitat: dry soils of waste lands, roadsides and deserts. Animals affected: sheep. Reference: Durrell et al. 1952, Kingsbury 1964. Literature Cited Aanes, W. a. 1961. Pingue (Hijmenoxys richardsonii) Poisoning in sheep. Am. J. Vet. Res. 22:47. Adler, H. E. 1949. Indigestion from an unbalanced kiawe (mesquite) bean diet. J. Am. Vet. Med. As- soc. 115:263. Alberts, H. W. 1931. Poisoning of livestock by plants. Alaska Agr. Expt. Sta., Rept., 19.30:35. Animal Disease and Parasite Research Division. 1958. Sixteen plants poisonous to livestock in the western states. USDA Farmers' Bull. 2106. Baker, C. J. L., and A. Eden. 1954. Studies on the oxa- late content of the leaves of certain varieties of Beta vulgaris. J. Agric. Sci. 44:394. Bamford, F. 1951. Poisons, their isolation and identi- fication. Blakiston Company, Philadelphia. 3d ed. Bankowski, R. a., R. W. Wichmann, and E. E. Stuart. 1956. Stomatitis of cattle and horses due to yel- low bristle grass (Setaria hitescens). J. Am. Vet. Med. Assoc. 129:149. Beath, O. a., J. H. Draize, and H. F. Eppson. 1933. Ar- row grass, chemical and physiological consid- erations. Wyoming Agri. Expt- St. Bull. 193. Beath, O. A., C. S. Gilbert, H. F. Eppson, and I. RosENFELD. 1953. Poisouous plants and livestock poisoning. Wyoming Agr. Expt. Sta. Bull. 324. BiNNs, W., AND L. F. James. 1961. A congenital defor- mity in calves, similar to "crooked calf disease, " has been experimentally produced by feeding heifers lupine and lead. Proc. Am. Soc. Anim. Prod. (Western Sect.) 12(66): 1. BoucHTON, I. B. 1943. Oak bmsh poisoning. Texas Vet. Bull. 5(4):2. September 1980 Brotherson et al.: Poisonous Plants 251 Brady, D. E., J. E. Comfort, J. F. Lasley, and VV. H. Pfander. 1955.-Forage poisoning in Missouri due to excessive amounts of nitrate. Missouri Agr. Expt. Sta., Bull. 652:19. Br.\kenridc;e, D. T. 1956. Nitrate poisoning caused by turnips and redroot. New Zealand Vet. J. 4:165. Buck, W. B., J. W. Dollahite, and T. J. Allen. 1960. Silver-leafed nightshade poisoning. J. Am. Vet. Med. Assoc. 137:348. Buck, W. B., L. F. James, and W. Binns. 1961. Changes in senim transaminase activities associated with plant and mineral toxicity in sheep and cattle. Proc. Am. Col. Vet. Toxicol. 1961:13. Burke, J. M., J. Marchisotto, J. J. A. McLaughlin, AND L. Provasoll 1960. Analysis of the toxin produced by Gonijaulax catenelUi in axenic pure culture. Ann. N.Y. Acad. Sci. 90:8.37. Cameron, K. 1952. Death camas poisoning. Northwest Med. 1952:682. Cann, H. M., and H. L. Verhulst. 1958. Poisonous plants. Natl. Clearinghouse Pois. Contr. Cent., September 1958:2. Carey, J. C. 1955. Black nightshade poisoning in swine. N. Am. Vet. 36:466. Carpenter, K. J., A. T. Phillipson, and VV. Thomson. 1950. Experiments with dried bracken {Pteris acftiiUna). British Vet. J. 106:292. Case, A. .\. 19.57. Some aspects of nitrate intoxication in livestock. J. Am. Vet. Med. Assoc. 1.30:.323. Christensen, J. J., AND H. C. H. Kernkamp. 1936. Stud- ies on the toxicity of blighted barley to swine. Minnesota Agr. Expt. Sta., Tech. Bull. 113. Clawson, a. B. 1933. Additional information con- cerning larkspur poisoning. Supplement to USDA,Varmers' Bull. 988. Clawson, A. B. 1933. The American groimdsels, species of senecio as stock-poisoning plants. Vet. Med. 28:105. 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Wild tobaccos (Xicotiana trigonophijUa Dunal and Xicotiana attentiata Torrey) as stock-poison- ing plants. USDA, Tech. Bull. 22. Mathams, R. H., and a. K. Sutherla.nd. 19.52. The oxa- late content of some Queensland pasture plants. Queensland J. .\gr. Sci. 9:317. M.ATHEws, F. P. 1933. The toxicity of Baileija multi- radiata for sheep and goats. J. .\m. Vet. Med. .^s- soc. 83:673. McCulloch, E. C. 1940. The experimental production of hepatic cirrhosis by the seed of Amsinckia in- tennedia. Science 91:95. McNair, J. B. 1923. Rhus dermatitis, its pathology and chemotherapy. University of Chicago Press. Mecke, M. B. 1979. Poisonous range plants of Wyom- ing—their importance and management. Annual Meeting of Society for Range Management, Cas- per, Wyoming. 14 pp. Unpublished manuscript. Merrill, L. B. and J. L. Schuster. 1978. Grazing man- agement practices effect on livestock loss from poisonous plants. J. Range Manage. 31(5): .351-354. Mettler, F. a., and G. M. Ster.n. 1963. 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The occur- rence of nitro-toxins in North .\merican Astra- galus (Fabaceae). Brittonia 29:310-326. THE SUCCESSIONAL STATUS OF CUPRESSUS ARIZOMCA Albert J. Parker .\bstbact.— Several investigations isize-class analysis, age-determination inquiries, and germination tests"* suggest that Cupressus arizonica of southeastern .\rizona is a pioneer species. The tree requires disturbance to remove or reduce soil litter, which other\%-ise inhibits the reproduction of the species. Reduction of light intensity caused by canopy closure appears to be less important than litter accumulation in restricting C. arizonica reproduction. Fol- lowing disturbance, successful establishment of seedlings may occur over an e.xtended period ,50 to 100 years I as Utter graduallv accumulates. The absence of C. arizonica seedlings in present populations suggest that fire suppres- sion policies on federal lands where C. arizonica occurs have altered fire frequency, and consequently have fostered a short-term reduction in C. arizonica establishment. Only in floodplain en\ironments. where flooding disturbs the soil surface, has much reproduction occurred in recent years. The long-term population pattern of C. arizonica appears stable, due to the great longevity of the species. Rough-barked Arizona c^'press {Cupressus arizonicu Greene; all taxonomy after Kearney and Peebles I960' is a tree species of local occurrence in the mountain ranges of south- eastern .\rizona, southwestern New Me.xico, western Texas, and northern Mexico. Though it has topically been characterized as a mois- tiu-e demanding species of riparian associ- ation iWolf 1948a. \Miittaker and Xiering 1965. BrouTi and Lowe 1974', recent findings (Parker 1980 > demonstrate that C. arizonica may occupy relatively xeric landscape posi- tions—including south-facing slopes, ridge tops, and convex slope configurations— at higher elevations laroimd 2000 ml In the first pubUshed report mentioning Cupressus arizonica regeneration. Posev and Goggans - 1967' obser\ed little reproduction anxAvhere in the Southwest, although they collected no data to substantiate or in- vestigate this phenomena. Thev did express concern that the limited natural range of the species, coupled with its apparent inability to reproduce in many natural circtmistances, could indicate that its existence was threat- ened. This paper investigates the population d\Tiamics, reproductive strategy, and role in species replacement processes of C. arizonica in the upland forests of southeastern Arizona. Study Are.a.s Populations of Cupressus arizonica were sampled in the Chiricahua Moimtains, the Department of Geography. Univeran- of Georgia, .Mhens, Georgia 30602. Blue Range north of Clifton, the Dragoon Mountains, and the Santa Catalina Moun- tains, all located in .\rizona (Fig. l\ Cupr- essus arizonica is most common in the wood- land zone of these mountain ranges (1300-1900 m; Wliittaker and Xiering 1965), a zone characterized by the frequent co-do- minance of alligator juniper Juniperus dep- peana\ pinyons [Pinus cembroides and P. edulisl and a nmnber of oak species, includ- ing silverleaf oak (Quercus hypoleucoides), netleaf oak [Q. rugosa), and .\rizona white oak Q. arizonica"!. Cupressus arizonica is pro- gressively more restricted to riparian habitats where the woodland zone grades into the shrub-dominated desert and semidesert vege- tation types below. It is also found in stands at higher elevations (1600-2300 mi with other conifers, notably Arizona pine (Pinus ponderosa var. arizonica^ Apache pine {P. engehiannii), Chihuahua pine P. leiophylla), and Douglas-fir iPseudotsuga menziesii); these forested stands often have closed ca- nopies. The climate of the woodland zone is sub- humid and mild, with about 50 cm annual precipitation and a mean annual temperature of 12.5 C. The wettest periods of the year are middle to late summer and midwinter (Bry- son and Lown.- 1955 1. The isolated mountain ranges of this region of basin and range topography are roughly linearlv aligned, northward trending, and 254 September 1980 P.\rker: Arizona Cypress Succession iDO ARIZONA /^C AT A LIN A ^ MTS. M3 i"''*"/"„ aV" •■;/'-$, f.ii-i/'//^ •'/; % ''^*i; i V"'.. -:. km. '-/y,...„.? ARIZONA MEXFCO" CARTOGRAPH.C LASGRATOfiY. uMVERSiTV Gf W.SGG\S;f4 - V.ADISOM Fig. 1. Study area uith the species range inset. The location of stands included in this study are noted by number. Open circles in the inset represent the three extensive areas of nonriparian C. ari:onica occuTTence in southeastern .\rizona: the Chiricahua Mountains, the Blue Range, and the Santa Catalina Mountains. 256 Great Basin Naturalist Vol. 40, No. 3 separated by extensive sediment-filled low- lands. The bedrock core of these ranges in- clude both igneous and sedimentary rocks (Fenneman 1931). Methods Environmental and vegetational measure- ments were obtained from 19 internally homogeneous stands of C. arizonica. Within each stand, three 4 by 25 m quadrats were oriented so that their central long axes were normal to a contour line bisecting the stand, with intervals of 10 m separating each quad- rat. The following measurements were re- corded within each quadrat: the species and circumference at breast height (1.37 m) of each tree, the number of saplings of all tree species, and the number of C. arizonica seed- lings. Any stem of at least 20 cm circum- ference at breast height was considered a tree. A sapling was defined as the stem of any tree species which exceeded breast height, but which was less than 20 cm in circum- ference at that height. All stems of tree spe- cies less than breast height were counted as seedlings. The depth of the litter layer in each stand was measured at 1 m intervals along the central long axis of each quadrat. A total of 60 litter depth measurements was made per stand. In two stands, both in the Chiricahua Mountains, all C. arizonica trees with a cir- cumference at breast height exceeding 30 cm were cored with an increment borer at breast height. These two stands were chosen for their differing size structures. One stand was characterized by trees few in number but large in diameter, but the other stand had a great number of individuals, particularly in the smaller size-classes. These two stands were selected as representative of 14 of the 19 stands sampled. The other five stands has a size structure intermediate in character be- tween those chosen for coring. In total, 79 trees were cored. Cores were aged in the laboratory, al- though the aging was complicated by two factors. First, several of the C. arizonica trees possessed rotted xylem tissue that caused fragmentation of some cores and loss of re- cord from the damaged segments. Second, C. arizonica has been shown to produce false an- nual rings (Bannan 1954). To circumvent the first difficulty, the length of each core was measured and the following formula was used to compute a "tree ring" age for each sample: 0.95 (^)(g) = t where: t = extrapolated tree ring age (yr) c = circumference of the tree (cm) g = number of annual rings per unit length of core (yr*cm-i; this value is later referred to as "mean time required per unit of radial growth") 0.95 = a coefficient to adjust for bark thickness With respect to the second complication. Clock and Agerter (1963) were able to utilize microclimatic records from a plantation of C. arizonica near Lubbock, Texas (500 km from the species' natural range) to investigate the production of false annual rings in this spe- cies. They reported that multiple false rings may be produced by an individual in a single year, and that the number of false rings is not consistent from tree to tree, or even from place to place on the same tree. Because of these difficulties, precise absolute aging was not possible, and tree ring ages reported in this study may be as much as two times larger than the actual tree age. Nevertheless, the ring counts and extrapolated ages provide an indication of the relative ages of individ- uals and thus allow identification and inter- pretation of the approximate age structures of the stands. Cones of C. arizonica were collected for the purpose of determining the mean number of seeds per cone, as well as to provide seeds for use in germination tests. All 250 cones collected were from the second year foliage of a number of C. arizonica individuals lo- cated in the Blue Range. Only closed cones were collected. Fifty of tht«e cones were ran- domly selected, and seeds from each cone were counted. Following this, the other cones were emptied of seeds, and all seeds were then sorted by size, the larger seeds being kept for use in germination tests. A series of germination tests were per- formed on lots of 100 seeds selected ran- domly from the seed source. These tests in- September 1980 Parker: Arizona Cypress Succession 257 vestigated the relationship of C. arizonica germination success to the following factors: freezing, flooding, fire, litter buildup, and light intensity. The physical conditions in which the germination tests were performed followed those of Wolf (1948b) in general (see Table 4). Results Stands were segregated into three groups, according to the size-class frequency distribu- tion of individuals of Cupressus arizonica. The eight stands of the first group display a generally logarithmatic decrease in the num- ber of individuals in successively larger size- classes, with the exception that the seedling layer is virtually absent (Fig. 2). Populations with a size stnicture similar to that of the first group are often characterized as stable through time, with losses of canopy individ- uals being balanced by replacements from the sapling layer (Daubenmire 1968). The six stands of the third group, in contrast, lack a logarithmatic trend. Instead, the number of individuals in successively larger size-classes is more constant (Fig. 2), suggesting that a past period of reproduction has ended. The 400-1 350- Group 1 1 — \ — \ — ] — I — r-1 i—\ — I — \ — I — I T* I ' (DraiDirjinLnioinminTTTTTT ■ ID £ r^ CO CNJ CD in to h- (DO) o>ai^CT)Oio>ojoi a> Group 2 " I 1 1 "" 1 — I I I I I ID ininiDuSininioiT) *- CNJ to ^ *0 to ^ 60- 40- 20- Group 3 Fig. 2. Composite size-class structure histograms for stands of C. arizonica. See text for a definition of each of the three groups. 258 Great Basin Naturalist Vol. 40, No. 3 second group includes five stands inter- mediate in character between the other groups. Even in those cases which appear to pos- sess a steady-state population above the sap- ling layer, the almost universal absence of C. arizonica seedlings in stand understories (Table 1) is conspicuous, and agrees with the observation of Posey and Goggans (1967). Conditions in the two stands which do con- tain a relatively large number of seedlings are significant; in both cases, disturbance has exposed bare mineral soil, in one case by nat- ural flooding and in the other by logging ac- tivities that mechanically stripped off the lit- ter from a portion of the forest floor. The behavior of other tree species in C. anzontca -dominated commimities fall pri- marily into two types, according to stand size-class information (Table 2). A first group consists of intolerant pioneer species, which reproduce episodically in association with disturbance events, and probably depend on wide dispersal of seeds to maintain represen- tation on a given site. These species display an even aged structure in most C. arizonica stands. The common upland conifers of the region, such as Apache pine, Arizona pine, and Douglas-fir fit this behavior pattern. A second group consists of tolerant species with stable populations that reproduce either veg- etatively of by widespread dispersal from ad- jacent locales. These display continues repro- Table 1. The seedling class. No. seedlings Mean Recently flooded stand Recently logged stand All other stands (17) Total 20 36 17 73 1.0 3.7 duction in C. arizonica-dominated stands, with a maximum number of stems occurring in recent cohorts. This group includes the typical pinyon/oak woodland dominants- Mexican pinyon, silverleaf oak, netleaf oak, and Arizona white oak. Core analysis indicated that the mean tree ring age of C. arizonica in the stand with few but large individuals was 317.1 years, with ages ranging from 197.6 to 456.7 years (Table 3). The majority of tree ring ages were be- tween 250 and 375 years. No seedlings, sap- lings, or trees of less than 30 cm circum- ference at breast height were found in the stand. Thus, the tree ring age of the youngest C. arizonica individual was almost 200 years. The mean tree ring age of C. arizonica trees in the stand with the greater density of small- er trees was 116.9 years, with an age range from 62.4 to 178.1 years. However, there were also a large number of saplings and trees too small to core, and these size-classes probably extended the age range down to about 25 years. The conspicuous absence of C. arizonica seedlings in the imderstory of this stand (Table 3) suggested that reproduc- tion has been inhibited during about the last quarter century. To determine the degree to which under- story individuals of C. arizonica were sup- pressed by the overstory, a linear correlation and regression analysis was performed on the mean time required for unit growth (yr*cm-i) for each tree against tree radius. A negative correlation coefficient would be predicted in conditions of understory suppression, because small trees would require longer periods of time to produce an amount of radial growth equal to that produced on the larger, more rapidly growing trees in the canopy in a shorter period of time. The results suggest that suppression of understory individuals Table 2. Composite size-class structnres for upland conifers and pinyon-oak species. The figures heading each size-class category are the smallest possible circumference in that class, and size-classes include all stems up to the next larger value. Upland conifers include Arizona pine, ponderosa pine, Apache pine, and Douglas fir. Pinyons in- clude Mexican pinyon and two-leafed pinyon. Oaks include silverleaf oak, netleaf oak, and Arizona white oak. sapl 20 30 40 Size-class 50 (cm 60 circumference) 70 80 90 100 110 120 Upland conifers 3 4 4 7 4 3 8 4 4 5 9 Pinyons Oaks 63 111 27 27 18 18 13 24 4 18 7 8 2 4 2 3 1 1 1 1 September 1980 Parker: Arizona Cypress Succession 259 Table 3. Characteristics of cored stands. First stand Second stand Tree density (ha"^) Total basal area (dm2-ha-i) Mean basal area of C. arizonica trees (dm^'tree"^ Number of C. arizonica cored 'Mean time required per unit growth of C. arizonica (yr-cnr') Mean extrapolated tree age of C. arizonica (yr) Standard deviation of extrapolated tree age (yr) Oldest C arizcmica extrapolated age (yr) Yoimgest C. arizonica extrapolated age (yr) Number of C. arizonica saplings and trees too small to core (20-30 cm) Number of C. arizonica seedlings Correlation coefficient for suppression test 1333 3733 11901 5854 9.51 1.49 21 58 24.2 16.6 317.1 116.9 63.0 26.4 456.7 178.1 197.6 62.4 81 -0.833 -0.673 'Calculated by dividing the number of annual rings per core by the length of that core, it expresses the number of years required for an individ- ual to add one cm of radial growth and is the reciprocal of the growth rate. The great number of seeds produced does not necessarily insure abundant reproduction. Sudworth (1915) noted that seeding of C. arizonica was best on moist, bare mineral soils. Field observations made during the course of data collection confirm this sugges- tion, in that C. arizonica reproduction was generally restricted to areas within 2 of the 19 stands, and then only on exposed mineral soil associated with recent floods or logging. Moreover, the weighted average of litter depth where C. arizonica seedlings were en- countered (1.74 cm) was only half the mean depth of litter for all stands in this study (3.42 cm), indicating that successful reproduction is associated with reduced litter depth. Germination tests also indicated that a lit- ter layer sharply reduced the germination and survival of seedlings. Onlv 3.7 percent of the seeds germinated on the litter-covered substrate, in contrast to 10.0 percent on min- eral soil controls (Table 4). Of the other fac- tors tested, freezing of seeds prior to germi- nation, reduced light intensity, and immersion of seeds in water did not alter ger- mination success. Simulated exposure of seed- bearing cones to canopy fire conditions, while reducing germination success (6.0 per- was evident in both stands (Table 3). The de- gree of suppression was greater in the older stand, as a consequence of the longer period of time over which competition was acting. These results indicate that small C. arizonica trees may persist beneath a canopy, and sug- gest that the species is shade tolerant. Evidence from core aging also .suggests that C. arizonica possesses potentially great longevity. Based on the rates of growth de- termined in this study (Table 3), and account- ing for multiple annual rings, some of the larger individuals encountered in canyon bot- toms (exceeding 1 m dbh) may be 300 to 500 years of age. Not only does the species achieve old ages, but also it produces great numbers of seed. The mean number of seeds per cone was 104.2, with a standard deviation of 19.8. It was estimated that healthy trees of moderate size (50 cm circumference) may produce from 103 to 10* cones per year, therefore re- sulting in an annual seed crop per tree of from 10^ to 10'^ seeds. Table 4. Germination test results. The soil in each pot was composed of a mixture of 50 percent sand and .50 percent silt loam. The surface was without litter cov- er. Pots were supplied with adequate water regularly and were grown under high levels of simlight. The pots were 15 cm in diameter and allowed free drainage be- low. The air temperature was between 20 and 24 C throughout the duration of the tests. Composite Number of germination Treatment test pots rates (%) Litter cover over mineral substrate 6 3.7 Seeds exposed to freezing before planting 4 8.8 Seeds immersed in agitated water 2 10.0 "Seeds exposed to canopy fire conditions 1 6.0 °° Seeds exposed to groimd fire conditions 1 0.0 Seeds grown under reduced light levels 2 8.5 °° "Controls 10 10.0 •—Cones filled with seeds were exposed to 80 C for eight minutes. ■-Cones filled with seeds were exposed to 315 C for eight minutes. *— Control conditions. 260 Great Basin Naturalist Vol. 40, No. 3 cent), did not destroy all seeds. In contrast, exposure to simulated ground fire conditions, which are much hotter, killed all seeds within the cones. Thus, C. arizonica seeds enclosed in cones on branches may remain viable after fires, and have the capability to germinate on the mineral seedbeds exposed by burning. Discussion The absence of C. arizonica reproduction under adults of the same species in natural stands is striking. Two interpretations of this observed absence of young individuals are possible. First, successful reproduction may be temporally sporadic and linked to occa- sional optimal climatic conditions that foster waves of C. arizonica reproduction. If such optimal conditions have not occurred during the last 20 or 30 years, the paucity of C. arizonica seedlings would be explained. How- ever, successful reproduction of C. arizonica was encountered in two recently disturbed habitats, and this indicates that recent climat- ic conditions are not inimical to the tree's re- production. The environmental conditions as- sociated with these cases of successful reproduction do support, however, a second interpretation of the absence of C. arizonica seedlings, namely, that disturbance is neces- sary to create conditions favorable for rege- neration, and that factors associated with a closed forest inhibit seeding by the tree. Stand size-class analyses would seem to both siipport and contradict this character- ization of C. arizonica as a species requiring disturbance for reproduction. Stands of the first group suggest that the species is tolerant, able to maintain itself under a closed canopy through time. Stands of the second and third size-class groups imply that the species is in- tolerant, with a period of establishment, fol- lowed by the cessation of C. arizonica repro- duction. Therefore, the latter groups support the hypothesis that the species is opportunis- tic, capable of invading open habitats, but not able to compete effectively with other species as successional processes proceed. Inspection of core analyses resolve this ap- parent paradox. The first stand cored, repre- senting those stands that are dominated by large individuals and which suggest that the species is a pioneer, possesses no individuals with a tree ring age of less than 200 years. The trees from this stand exhibit a tendency to clump in the 250- to 375-year tree ring age range. Lack of successful C. arizonica regeneration over an extended period, com- bined with a tendency toward clumping of ages, suggests that, following a lengthy peri- od of C. arizonica establishment, reproduc- tion is virtually eliminated. The second stand cored, representing those stands with a mixed size structure and suggesting that the species is a climax-type, possesses trees ranging in tree ring age from 25 to 150 years. This stand possesses, however, a conspicuous absence of reproduction during the last quarter century. These data, in the same manner as those from the first stand, suggest that following a lengthy period of colonization and reproduc- tion of C. arizonica, perhaps 50 to 100 years in length, reproduction subsequently ceases. The population structures of these two stands, therefore, differ only in the elapsed times since disturbance, rather than in some fundamental difference in the ecological be havior of the species. In both cases, C. arizo- nica acts as an intolerant species, requiring disturbance to create invadable habitats. In the denser, more youthful C. arizonica popu- lation of the second stand, disturbance was more recent than in the older population of C. arizonica in the first stand. The initial in- terpretation of stand structures of the first type— that the species is a climax-type, with a stable, self-perpetuating population— is shown to be incorrect. Stands of the first group attain this pseudoclimax type structure only temporarily as a consequence of the ex- tended period following disturbance when C. arizonica reproduction is possible. Thus, the three groups identified by differing size structures may be interpreted as stages of a temporal continuum, with each stage reflect- ing a longer period of development since a disturbance event. Other results yield evidence favoring the interpretation of C. arizonica as an intolerant species. The longevity of C. arizonica is char- acteristic of many pioneer species, as it in- creases the allowable length of time between periodic disturbances, thus enhancing the probability that perturbation will occur on or near the site while mature seed trees are available for colonization. Seed counts and estimated cone crop sizes indicate that C. September 1980 Parker: Arizona Cypress Succession 261 arizonico is a prolific seeder, also a character- istic of the reproductive strategy of a pioneer species, because it increases the probability that a germule will be dispersed to an open habitat. The inverse relationship of litter depth to C. arizonica seedling number in- dicates that partial or complete removal of litter on the forest floor is necessary to pro- mote reproduction. The inhibition of litter on germination in test pots further corroborates the negative effects of litter accumulation on this tree's regeneration. This need for mineral seedbeds, created by natural disturbance (i.e., fire or flooding) or by certain human activi- ties (i.e., logging) is typical of pioneer ele- ments. Finally, the ability of seeds to remain viable following a canopy fire is an opportu- nistic character often favored by intolerant species that must colonize open habitats fol- lowing such fires. In contrast to the characteristics suggesting that C. arizonica is a pioneer species, the tree's abilities to germinate in low light con- ditions and to tolerate shaded sites by sup- pression of growth are characteristics more commonly associated with climax-type than with pioneer species. These responses to light are puzzling and deserve closer inspection in the future. In general, however, it is con- cluded that C. arizonica is a pioneer species, requiring periodic disturbance to open new sites for colonization. A unique characteristic of C. arizonica stand dynamics is the long period, as long as a century, during which colonization and re- production remain possible after the disturb- ance event. Jenny et al. (1949) have demon- strated that, in ponderosa pine forests at 1220 to 2220 m elevation in California, 100 to 200 years are required to reach a near-equilib- rium steady-state condition of litter thickness, in which mineralization of organic matter balances the addition of litter produced by the vegetation. Such a long period of litter accumulation is characteristic of moist, cool montane conifer forest ecosystems, and fits well with the observed time scale of C. arizo- nica establishment on a site. This peculiarly long period of colonization appears to be re- lated to the shade-tolerant character of the species. Most pioneer species, being intoler- ant, rapidly cease reproduction as light levels decrease under the species' own canopy. The ability of C. arizonica to reproduce in shade allows it to continue reproducing for a much longer period than most colonizing species. This period is later terminated by the more gradual process of litter accumulation. Thus, C. arizonica may be considered to be a pioneer species that is intolerant of litter accumulation under a closed canopv, even though it is tolerant of low light levels fol- lowing colonization of disturbed sites. Addi- tionally, its habit of seed persistence in the cone is critical to its maintenance on most sites. It may not depend, then, on seed dis- persed from distant sources to colonize a giv- en disturbed area. Using the Noble and Slatyer (1977) model of successional processes (from Cattelino et al. 1979), this study suggests three tvpes of species that exhibit fundamentally different responses to disturbance events. First, the up- land conifers are DI species, characterized bv widespread dispersal capabilities (D) and in- tolerance to a closed canopy (I). These trees depend on colonizing disturbed sites with seed from an off-site source. Second, the pin- yon oak group are DT/VT species, which persist on a disturbed site by colonization from an off-site source (D— pinyon) or by vegetative reproduction (V— oaks). These spe- cies maintain themselves by virtue of their tolerance of closed canopy conditions (T). Cupresstis arizonica, the third type, is a CI (DI) species. As described above, it is intoler- ant (I) of closed forest conditions and gener- ally is maintained locally by seed persistence in cones from the canopy (C), which shelter viable seeds through disturbance events. Be- cause of its limited range, C. arizonica is less commonly maintained by dispersal from adja- cent poulations (D), except perhaps in ripa- rian stands, where surface water wash may supply seeds to wash environments from sur- rounding upland slopes. These three types of species, in com- bination with differential longevity patterns, yield three possible pathways of community development (Fig. 3). The initial composition of each case is represented by at least one stand sampled in this study. The first and sec- ond cases diagram multiple successional pathways in communities which possess rep- resentatives of the three behavioral types. In the first case, the model assumes a greater 262 Great Basin Naturalist Vol. 40, No. 3 Case 1 b Case 2 ' \a_c I Case 3 N \ a»^ / ^"^ T Fig. 3. Hourly distribution of individual flower vis- itors to Penstemon cijaniinthtis (dotted line) and P. ea- tonii (solid line) throughout the observation period. Fig. 4. Hourlv distribution of insect families that vis- ited Penstemon ri/anantlius (dotted line) and P. eatonii (solid line) throughout the observation period. September 1980 Bateman: Comparative Floral Biology 271 longed to three hymenopteran families. Dur- ing tlie heat of the day, when the greater numbers of insects were actively foraging, 4 or 5 attempted to enter the narrow corolla tube. Few were successful in their efforts. Once during the observation period, a broad- tailed hummingbird visited the plant at 0930, before the temperature had risen above 24 C (75 F). The bird moved systematically down the canyon, stopping at every P. eatonii indi- vidual within 10 or 20 m of either side of the path. Upon reaching the observation plant. the bird sampled five or six flowers on four of the six stalks, pausing less than one second at each flower. It hovered in the air .slightly be- low each flower (Figs. 3 and 4). Penstemon cyananthu.s averaged 66.7 per- cent fruit set per plant, and P. eatonii showed a much higher 82.4 percent (Table 2). Discussion The results of this study clearly indicate that the two penstemons considered have de- Table 1. Activity and presence of each family throvigho\it the observation period. The first (upper) n\imber in- dicates the number of individual visitors; the second (lower) indicates the total number of flowers visited. Hour observed Family 8 9 10 11 12 1 2 3 4 5 6 Total Penstenion cyanantlnis .\pidae' Chrysididae' 1 2 1 3 3 10 3 12 1 3 3 8 6 22 Halictidae' Hesperiidae- Megachilidae' 'Nymphalidae Pieridae- Syrphidae' Vespidae' Total 1 3 1 6 2 4 2 7 3 9 10 25 2 11 6 17 10 27 14 36 5 4 3 2 2 1 21 6 4 5 6 15 2 3 2 1 1 29 13 25 / 16 10 10 18 2 75 6 4 39 26 15 47 5 183 6 6 2 2 1 1 1 1 11 1 1 3 3 8 2 8 2 3 25 22 6 26 9 15 89 r; 28 26 21 24 7 153 s (.3 66 46 61 23 370 Penstenwn eatonii Formicidae' Halictidae' Megachilidae 'Trochilidae' Total 1 15 2 16 2 9 5 17 1 15 2 1 2.3 5 2 54 Key to the orders: 1, Hymenoptera; 2, Lepidoptera^ 3. Diptera; 4, Apodiformes {'Selasphorus platycercus). 272 Great Basin Naturalist Vol. 40, No. 3 Table 2. Percent fruiting success per plant of Penste- mon cyananthus and P. eatonii (Number of fruit/spent flowers/plant). (Difference significant at the .05 level) Plant P. cyananthus P. eatonii % % 1 84.7 45.5 2 47.5 93.3 3 59.8 73.1 4 47.6 68.5 5 74.8 89.7 6 75.5 90.0 7 69.7 77.4 8 79.3 9 90.8 10 86.1 11 95.4 12 82.4 13 100.0 Average % 66.7 82.4 veloped different ways of attracting pollina- tors. The narrow red corolla tube of P. eaton- ii physically excludes all but a few small in- .sect visitors, and the absence of odor appears to minimize attractiveness to insect visitors. The absence of any sort of a landing pad hin- ders the ability of nonhovering visitors to successfully work the flowers. The only ob- vious diurnal pollinator, a hummingbird, is le.ss frequent but more systematic and specif- ic. That the job of pollination is done more efficiently by such a specific pollinator is sug- gested by the higher fruit set. The more promiscuous P. cyananthus also enjoys a fairly high fruit set. It does this, however, with larger, more accessible blos- soms and with no assurance that its pollina- tors will be species specific. The flowers are, of necessity, displayed so as to form a con- venient landing pad for approaching hyme- nopterans. Limitations Although the initial implications are clear, these data constitute only preliminary results. Data were collected during daylight hours late in the flowering season, when flowers of both species contained little or no nectar. Only one hummingbird was observed directly during the study, but my presence may have frightened usual avian visitors away. No at- tempt was made to observe early evening or nocturnal pollinators. The degree to which either species is capable of self-pollination is unknown. Literature Cited Bond, H. W., and W. Brown. 1978. The exploitation of floral nectar in Eucalyptus incrassata by honey- eaters and honeybees. Unpublished manuscript. BoRROR, D. L., and R. E. white. 1970. A field guide to the insects. Houghton Mifflin Co., Boston. Faegri,K., and L. van der Pijl. 1971. The principles of pollination ecology. 2d ed. Pergamon Press, Ox- ford, England. Free, J. B. 1970. Effect of flower shapes and nectar guides on the behavior of foraging honeybees. Be- havior 37:269-285. Grant, K. A. 1966. A hypothesis concerning the preva- lence of red coloration in Californian humming- bird flowers. American Nat. 100:85-98. Levin, D. A. 1969. The effect of corolla color and out- line on interspecific pollen flow in Phlox. Evolu- tion 23:444-445. Proctor, M., and P. Yeo. 1972. The pollination of flow- ers. Taplinger Publishing, New York. Raven, P., R. F. Evert, and H. Curtis. 1976. Biology of plants. 2d ed. Worth Publishers, New York. Bobbins, C. S., B. Bruun, and H. Zim. 1966. Birds of North America. Western Publishing Co., New York. Welsh, S. L., and G. Moore. 1973. Utah plants: Tracheophyta. Brighain Young University Press, Provo, Utah. DIFFERENTIAL HABITAT UTILIZATION BY THE SEXES OF MULE DEER Michael M. King' and H. Duane Smith' Abstract.- Habitat segregation trends have been observed and published for the sexes of mule deer {Odocoilem hcmionus) based on elevation and slope exposure. Despite these brief descriptions, quantitative studies on habitat segregation by the sexes of mule deer are lacking. Results of research conducted in central Utah indicated no signifi- cant difference in elevation positions used by males, but did show significant difference in utilization of studv sites based on slope exposure, relative percentage forb cover, and relative percentage hiding cover. Males were most com- mon at sites characterized by low forb abundance and hiding cover, and on south-facing exposures. Females were most common at sites characterized by high forb abundance and hiding cover, and on north-facing exposures. Pos- sible advantages of habitat separation to both sexes and management implications are discu.ssed. Mule deer {Odocoileus hemionus), like many other ungulates, seem to exhibit habitat partitioning between sexes (Darling 1937, Estes 1974, Geist 1974, 1977, Gest and Pe- tocz 1977, Hirth 1977, Leuthold 1978). DeVos et al. (1967) indicated that male and female mule deer are separated throughout the year with the exception of the breeding season. Dasmann and Taber (1956) found that males occupied more open south-facing slopes and females occupied densely vegeta- ted north-facing slopes. Several workers have suggested that males prefer higher altitudes and ridge tops more than do females (Cowan 1956, Miller 1970). The same trend was ob- served in a Nevada mule deer herd by Robin- ette et al. (1977) where subalpine and alpine conditions were prevalent. Males were found predominantly above 3000 m elevation, whereas females were more often below 2500 m elevation. Although habitat separation by male and female mule deer has been report- ed, little attempt has been made to quantify differential habitat use or to describe site dif- ferences other than to suggest slope exposure and elevational differences. The objectives of this study were: (1) to de- termine quantitatively if male and female mule deer differentially utilize habitat, (2) to suggest possible advantages to habitat separa- tion by sexes of mule deer, and (3) to identify critical management problems related to dif- ferential resource utilization between male and female mule deer. Study Area The study was conducted on the Bighorn Ranch, a privately owned ranch in the Nebo Range of the Wasatch Mountains, Utah. The study area was approximately 1130 ha in size, ranging from 2200 to 2500 m elevation. Human access is restricted, thus providing a relatively undisturbed area for observation of mule deer behavior, distribution, and habitat utilization. Ridge tops, south-facing slopes, and other well-drained areas were dominated by Gambel oak {Quercus gamhelii) and big sagebrush {Artemisia tridentata) communities with little herbaceous growth. Drainage bot- toms, north-facing slopes, and well-watered areas were dominated by quaking aspen {Pop- ulus tremuloides). Rocky Mountain maple (Acer glabrum), and chokecherry {Prttnus vir giniana) communities, with numerous forb species in the understory. Methods Preliminary observations made in Septem- ber 1977 to determine deer distribution in the various watersheds of the ranch indicated habitat segregation between male and female mule deer. Based on that survey, the follow- ing spring nine study sites (Fig. 1) were se- lected where deer numbers were relatively high. Other areas had equally as many deer, but excessive area or distance from access roads made observation unrealistic. 'Department of Zoology, Brigham Young University. Provo, Utah 84602. 273 274 Great Basin Naturalist Vol. 40, No. 3 THE BIGHORN RANCH O Study Sites — Ranch Border Nephi and Fountain Green Highway 89 N Highway 89 Fountain Green Fij;. 1, Map ot the Bit^honi Ranch showing the approximate location of study sites. September 1980 King, Smith: Deer Habital Utilization 275 Each study site was observed weekly from 1 June 1978 to 1 September 1978. Observa- tions were made from established observation points or by vehicle from sunrise until late morning or from early afternoon until dark. Ob.servation time for each site was alternated weekly between morning and evening so ap- proximately equal observation time was spent at each site during each time period. A variable 15-45X spotting scope and 12X bi- noculars were used for daytime observation, and a 200,000 candlepower spotlight oper- ated through the electrical system of the ve- hicle, along with spotting scope and binocu- lars, was used for observation at night. Total observation time for the study exceeded 900 hours. Observed deer were recorded according to sex, slope position (Fig. 2), and slope expo- sure. A 2X4 contingency analysis (Zar 1974) was performed to determine significant dif- ferences in utilization of slope positions by males and females. Total numbers of males and females recorded at north- and south-fac- ing exposures were also subjected to contin- gency analysis to determine slope exposure usage differences. To characterize study sites two critical parameters, relative abundance of forb and hiding cover, were examined at each site. Forbs were defined as succulent, low-growing, nonwoody vegetation, and hid- ing cover as vegetation more than 2 m in height. Both estimates were determined by a line-point transect method for determining relative abundance of vegetation (Kershaw 1973). Simple correlation procedures relating rel- ative abundance of forb and hiding cover with the corresponding male/female ratio for each site were used to determine if utiliza- tion of sites by males and females differed significantly based on forb abundance and hiding cover (Zar 1974). The maximum prob- ability accepted for statistical significance was 0.05; probabilities less than 0.01 were considered highly significant. Results and Discussion Analysis of slope positions used by males and females (Table 1, Fig. 3) showed no sig- nificant difference in slope position utiliza- tion by sex (P = 0.06). The data, though not significant at the 0.05 level, approach signifi- SLOPE POSITIONS ^^^^jM-i^-jt-^is^aN. Fig. 2. Diagram of the four slope positions at each site; slope position 1 = canyon bottom to '4 slope, slope position 2=1/2 slope to '/2 slope, slope position 3 = '/2 slope to % slope, and slope position 4 = \ slope to ridgetop. 276 Great Basin Naturalist Vol. 40, No. 3 Table 1. Total number of male and female mule deer at each slope position for all study sites. 2X4 contingency analysis indicates a nonsignificant difference in utilization of slope position by male and female mule deer (X- = 7.42, df = 3, P = 0.06). Numbers in parentheses are expected values. Slope position Sex 1 2 3 4 Total Males 74 (86.9) 63 (56.4) 42 (39.9) 14 (9.8) 193 Females 157 (144.1) 87 (93.4) 64 (66.1) 12 (16.2) 320 Total 231 150 106 26 513 cance and indicate support for elevational segregation observed in other areas. Further categorical analysis to determine usage of in- dividual slope positions by males and females shows that considerable differences exist in utilization of slope positions 1 and 4 by males and females (Fienberg 1977), with more fe- males than males at slope position 1, and more males than females at slope position 4 (position 1 male = -.208, position 1 females = + .208; position 4 males = -I- .245, posi- tion 4 females = -.245; positive values in- dicate most usage at slope position). Analysis of slope exposure use by males and females UJ < O 111 < 100 -I 75 - ^ 50 - + FEMALES • MALES Ik O O o u 85 25 T 1 r 1 2 3 SLOPE POSITION Fig. 3. Percentage of total males and total females observed at each slope position. September 1980 King, Smith: Deer Habital Utilization 277 showed a significant difference (P< 0.005), with males most often at south-facing slopes and females most often at north-facing slopes (Table 2, Fig. 4). Calculated male /female ratios for each site correlated with corresponding relative per- centages of forb and hiding cover (Table 3, Fig. 5) showed a highly significant negative correlation between forb abundance and male/female ratios (r = -.89, df = 7, P< 0.005) and a significant negative correla- tion between male/female ratios and hiding cover (r= -.69, df = 7, P<0.05). Therefore, as forb and hiding cover increased, the male/female ratios decreased, indicating that females select areas characterized by rela- tively high forb and hiding cover densities, but males select areas characterized by low forb and hiding cover densites. Table 2. Total number of male and female mule deer at each slope exposure for all study sites. 2X2 contin- gency analysis indicates a significant difference in utili- zation of slope exposure bv male and female mule deer (X^ = 29.3, df = 1, P< 0.005). Numbers in parentheses are expected values. Slope exposure Sex North-facing South-faciiiu T..i,,l Males 64 (93.7) 129 (99.3) 193 Females 185 (155.3) 135 (164.7) 320 Total 249 264 513 To comprehend implications of this pat- tern of spatial separation, po.ssible advantages accrued by males and females in their spring- summer habitats should be examined. During the spring-summer season perhaps the most critical events to females are production and 100 -1 3 o Q. X Q. o < < ui < z < UJ < 75 - 50 - 67 33 25 - NORTH-FACING EXPOSURE SOUTH-FACING EXPOSURE MALES FEMALES Fig. 4. Percentage of total males and total females observed at each slope exposure. 278 Great Basin Naturalist Vol. 40, No. 3 § U flO O Ik > < Ul > O o z o z Ul > 50 - 40 - 30 - 20 - 10 - r= -.89 P<0.005 50 - 40 - = 30 - 20 - 10 - »'=-.69 P<0.05 T" 1.0 2.0 1^ 3.0 "T" 4.0 male/female ratios Fig. 5. Correlation of relative percentage fort) cover and relative percentage hiding cover with corresponding male/female ratio for each study site. September 1980 King, Smith: Deer Habital Utilization 279 Table 3. Relative percentage forb cover, relative percentage hiding cover, total number of male and female mule deer, and male/female ratios for each study site. Site No. Relative % forb cover Relative hiding cover Males Females M/F ratios 40.50 19.00 7 63 0.11 32.25 14.00 6 39 0.15 41.25 19.(X) 12 61 0.20 46.50 22.(X) 2 25 0.08 37.75 16.00 7 46 0.16 16.50 15.00 52 29 1.80 19.00 13.00 32 23 1.40 20.75 8.00 38 15 2.53 19.75 17.00 37 19 1.95 rearing of offspring. This implies that consid- erable energy is apportioned to gestation, parturition, and lactation beyond normal body maintenance requirements (Nelson 1975, Stebbins 1977). If these requirements are not met through diet resources, body re- serves are utilized, thus reducing offspring vigor and survival. Since energy demands for offspring production exceed normal energy requirements, it is important that females oc- cupy areas where nutritious, high-quality for- age is readily available. Adequate nutrition insures successful offspring production and facilitates proper lactation. Research in- dicates that, during the time period critical to fawn production, high moisture content, ease of digestion, and increased nutrition content (Smith 1952, Short 1966, Short and Reagor 1970, Boeker et al. 1972) make the forbs preferred diet items of mule deer (Smith 1952, Morris and Schwartz 1957, Lo- vass 1958, Anderson et al. 1965, Crouch 1966, Dasmann et al. 1967, Nelson 1975). It has also been shown that deer herds having a variety of succulent forage in their diets have greater herd productivity and vigor than those that utilize a greater percentage of woody vegetation (Biswell 1961, Julander et al. 1961, Boeker et al. 1972, Nelson 1975, Pe- derson and Harper 1978). We assume, there- fore, that forb abundance is related to site quality and that areas of high forb abundance are considered high quality, whereas areas of low forb abundance are low quality. Deer that select forb-rich areas would have survi- val advantages because of availability of choice forage. It follows that females should select high-quality sites to assure adequate nutrition and energy for fawn production and survival. However, it seems strange that males would occupy areas of low forb abun- dance when selection of high-forb areas would more readily insure adequate energy for increasing body size, accumulating fat re- serves, and developing antlers. We suggest as a partial explanation that it is more advanta- geous for the sexes to be separated to reduce chances of energy-expensive agonistic expres- sion between males and females (Geist and Petocz 1977, McCullough 1979). This allows energy allocation to gestation, parturition, and lactation rather than to stresses of haras.s- ment. Females on ranges uncontested by males should leave more offspring, and those behavioral traits responsible for habitat parti- tioning should be selected to increase the population. Males that did not compete for resources necessary for fawn production, though occupying lower-quality sites, should likewise leave more offspring to succeeding generations than males whose behavior bring them into competition with their offspring (Wilson 1975, Geist and Petocz 1977, McCullough 1979). Differential habitat use patterns could also confer advantages to males and females by increasing odds for predator avoidance. Ecol- ogists working on vmgulates have suggested a theory of predator avoidance based on the relative degree of habitat openness and group size. Ungulates that inhabit densely vegetated areas usually occur as solitary animals or in small groups that use hiding as a mechanism for predator avoidance, whereas ungulates that occupy open areas are primarily herding animals that rely on the use of collective senses and high mobility for predator escape 280 Great Basin Naturalist Vol. 40, No. 3 (Dasmann and Taber 1956, Kitchen 1974, Hirth 1977). Life history studies of mule deer (Linsdale and Tomich 1953, Robinette et al. 1977), along with personal observations, give sup- portive evidence to this hypothesis. Males and females form different-sized groups in their preferred habitats. During spring and summer months females seek isolation in areas where hiding cover is relatively abun- dant (Fig. 5) and tolerate few deer other than their offspring of the year. This partial. soli- tary existence has advantages from a predator avoidance standpoint in that females can se- cret themselves and their offspring in dense vegetation during periods of high vulnera- bility to potential danger. After the young are bom, maternal duties restrict the mobil- ity and escape efficiency of females. It would, therefore, be advantageous for fe- males with fawns to avoid open habitats where predators can detect and capture them or their fawns more easily. In contrast, males on the Bighorn Ranch during the same time period were often ob- served to form fraternal groups in areas with relatively low abundance of hiding cover (Fig. 5). Male groups that inhabit open areas of high visibility can collectively monitor their surroundings and take advantage of rap- id flight when escape is necessary. This is better than hiding in dense cover, where ef- fectiveness of concealment would be reduced by large groups and would increase chances of detection by predators. Furtlier supportive evidence for the pre- dator avoidance theory is provided by the differential use of slope positions 1 and 4 by males and females. More females than males occupied slope position 1, the lower position of the slope where the greatest abundance of hiding cover was located. This suggests the importance of cover to females. More males than females used slope position 4, the open ridges, suggesting preference by males for the areas of high visibility. In this study habitat separation by males and females is primarily based on slope expo- sure, forb abundance, and hiding cover, with some evidence, though not significant, for elevational segregation. We do not detract from the importance of elevational segrega- tion as it has been observed frequently in other areas, but suggest to wildlife managers that there are several habitat separation pos- sibilities, depending on characteristics of mule deer range in a given area. We encour- age that further research delineating segrega- tion characteristics, advantages, and mecha- nisms be initiated so that management implications can be evaluated. Knowledge of habitat separation between the sexes of mule deer will have considerable influence on several critical management problems. Three important problems as we view them are now discussed. First, as deer populations are being censused in various areas and habitats, a prime concern is the de- termination of an accurate sex ratio. If man- agers are not aware of site-specific habitat separation by male and female deer, biases favoring one sex over the other will arise in calculated ratios depending on the area sam- pled. Failure to determine accurate sex ratios will allow faulty plans to be devised and im- plemented. Second, through recognition of specific habitat requirements of the sexes, it is possible that habitat can be manipulated through appropriate techniques to create conditions favorable to either sex. This will allow sex ratio manipulation depending on management needs. Third, critical areas to females and offspring as well as males must be protected from detrimental commercial, industrial, and recreational development. De- struction of important fawning areas through development will force females to occupy suboptimal habitats and result in reduced fawn production and survival. Development in areas occupied predominantly by males will restrict fall hunting and ultimately re- duce herd productivity if adequate male/female ratios are not maintained. An understanding of how male and female mule deer partition the habitant and how hab- itats preferred by females differ from those most frequented by males will undoubtedly improve abilities to effectively manage mule deer habitat. Proper use of knowledge re- garding differential habitat and resource uti- lization by the sexes of mule deer can in- crease the efficiency with which agencies manage the deer resource. September 1980 King, Smith: Deer Habital Utilization 281 Literature Cited Anderson, A. E., W. A. Snyder, and G. W. Brown. 1965. Stomach content analysis related to condi- tion in mule deer, Guadalupe Mountains, New Mexico. J. VVildl. Manage. 29: .352-.366. BiswELL, H. H. 1961. Manipulation of chamise brvi.sh for deer range improvement. California Fish and Game 47:128-144. BoEKER, E. L., V. E. Scott, H. G. Reynolds, a.nd B. A. Donaldson. 1972. Seasonal food habits of mule deer in southwestern New Mexico. J. Wildl. Manage. .36:56-6.3. Cowan, I. M. 1956. Life and times of the coast black- tailed deer. Pages 52.3-617 in W. P. Tavlor, ed.. The deer of North America. Harrishurg: Stack- pole Co. 668 pp. Crouch, G. L. 1966. Preferences of black-tailed deer for native forage and Douglas-fir seedlings. J. Wildl. Manage. .30:471-475. Darling, F. F. 1937. A herd of red deer. London: Ox- ford University Press. 215 pp. Dasmann, R. F., and R. D. Taber. 1956. The behavior of Columbian black-tailed deer with reference to population ecology. J. Mammal. 37:143-164. Dasmann, W., R. Hubb.\rd, VV. G. MacGrecor, and A. E. Smith. 1967. Evaluation of the wildlife results from fuel breaks, browsewavs, and tvpe con- versions. Proc. 7th Tall Timbers Fire Ecol. Conf. 7:179-193. Devos, a., P. Brokx, and V. Geist. 1967. A review of social behavior of North American cervids during the reproductive period. Amer. Midi. Natur. 77:390-417. Estes, R. D. 1974. Social organization of the .\frican bo- vids. Pages 166-205 in V. Geist and F. Walther, eds.. The behavior of ungulates and its relation to management. Vol. 1. Morges, Switzerland: lUCN Publ. No. 24. 511 pp. FiENBERG, S. E. 1977. The analysis of cross-classified cat- egorical data. Cambridge: MIT Press. 151 pp. Geist, V. 1974. On the relationship of social evolution and ecologv in ungulates. .\mer. Zool. 11:205-220. ' 1977. A comparison of social adaptations in rela- tion to ecology in gallinaceous bird and ungulate societies. Ann. Rev. Ecol. Syst. 8:19.3-207. Geist, V., and R. G. Petroczy. 1977. Bighorn sheep in winter: do rams maximize reproductive fitness bv spatial and habitat segregation from ewes? Can. J. Zool. ,55:1802-1810. Hirth, D. H. 1977. Social behavior of white-tailed deer in relation to habitat. Wildl. Monogr. 53:1-55. Julander, O., W. L. Robinette, and D. A. Jones. 1961. Relation of summer range condition to mule deer herd productivity. J. Wildl. Manage. 25:54-60. Kershaw, K. A. 1973. Quantitative and dynamic plant ecology. 2d ed. New York: American Elsevier Publishing Co. Inc. .308 pp. Kitchen, D. W. 1974. The social behavior and ecolog>' of the pronghorn. Wildl. Monogr. ,38:1-96. Leuthold, W. 1978. .\frican ungulates: a comparative review of their ethology and behavioral ecology. New York: Springer- Verlag. .307 pp. Linsdale, J. M., and p. Q. Tomich. 1953. A herd of mule deer. Berkeley: University of California Press. 567 pp. LovASs, A. L. 1958. Mule deer food habits and range use. Little Belt Mountains, Montana. J. Wildl. Manage. 22: 275-283. McCullough, D. R. 1979. The George Reserve deer herd. Ann Arbor: University of Michigan Press. 271 pp. Miller, F. L. 1970. Di.stribution pattern of black-tailed deer in relation to enviromneut. J. Mammal. .52: 248-259. Morris, M. S., and J. E. SvHwartz. 1957. Mule deer and elk food habits on the National Bison Range. J. Wildl. Manage. 21:18.3-189. Nelson, J. R. 1975. Forest fire and big game in the Pa- cific .Northwest. Proc. 15th Tall Timbers Fire Ecology Conf. 15:85-102. Pederson, J. C, AND K. T. Harper. 1978. Factors in- fluencing productivity of two mule deer herds in Utah. J. Range Manage. 31:105-110. Robinette, W. L., N. V. Hancock, and D. A. Jones. 1977. The Oak Creek mule deer herd in Utah. Salt Lake Citv: Utah State Div, Wildl. Res. Publ. No. 77-15. 148 pp. Short, H. L. 1966. Effects of cellulose levels on appar- ent digestability of feeds eaten bv mule deer. J. Wildl. Manage.' .30: 163- 167. Short, H. L., and J. G. Reagor. 1970. Cell wall diges- tability affects forage value of woody twigs. J. Wildl. Manage .34:964-967. Smith, A. D. 1952. Digestability of some native forages for mule deer. J. Wildl. .Manage. 16:.309-,312. Smith, J. G. 1952. Food habits of mule deer in Utah. J. Wildl. Manage. 16:148-155. Stebbins, L. L. 1977. Energy requirements during repro- duction of Peromyscus tnanictilattts. Canadian J. Zool. 55:1701-1704. Wilson, E. O. 1975. ScxiobiologV'. Cambridge: Belknap Press of Harvard University Press. 697 pp. Zar, J. H. 1974. Biostatistical analysis. Englewood Cliffs: Prentice-Hall Inc. 620 pp. TEMPORAL ACTIVITY PATTERNS OF A DIPODOMYS ORDII POPULATION Clive D. Jorgensen,' H. Diiane Smith,', and James R. Garcia' Abstract.— Temporal activity patterns for Dipodomijs ordii were generally bimodal during the summer, with the highest peak occurring during early predawn hours when conditions were optimum for water conservation. Removal of dominant members in the population resulted in a substantial shift in the activity pattern to increased activity during the evening hours. Ecologists studying small mammals must contimially attempt to resolve the problems of inadequate methods to study daily activity patterns in natural environments, especially when studying secretive and/ or nocturnal species that require trapping procedures. Usefulness of temporal activity data has been demonstrated in simulation trapping studies (Burnham and Overton 1969, Manly 1970, Jorgensen et al. 1972), population estimator studies (Scott et al. 1978), and energetics studies (Kenagy 1973), among others. Al- though methods for obtaining data under field conditions have not been well devel- oped, some have been reported (Jorgensen and Hayward 1965, Eider 1968, Marten 1973). Generally, their results are deficient in one or more of the following: (1) numbers of recorded activity events per day, (2) timing of the observed or measured activity event, (3) ability to assign an activity event to a spe- cific individual, and (4) correlation between the measured activity under laboratory con- ditions with analogous activity in natural en- vironments. Harling (1971) attempted to relieve some of the difficulty in recording activity by de- veloping a trap that could be continually monitored during a trapping period. His traps were electrically wired to a central communications console. Using walkie-talkie communications between someone on the grid and another at the console, Harling (1971) was able to obtain the precise time when an animal was caught as well as when it was released. We extended Harling's (1971) methods to include an entire grid of traps comparably wired (Garcia et al. 1974) to study a popu- 'Department of Zoology. Brigham Voting University, Provo, Utah 84602. lation of Dipodomys ordii under field condi- tions. Our objectives were to determine the optimum foraging times for D. ordii in the salt desert shrub community of west-central Utah, and illustrate activity patterns that could assist in interpreting trapping data from other studies that include this species. Study Site and Methods Data were collected from two sites at the Desert Range Experiment Station, Millard Co., Utah, from 21 August to 3 September 1971 and 25 June to 27 August 1973. Al- though D. ordii was the species trapped most frequently during this study, Perognathus longimernbris was common and Peromyscus maniculatus and Onychomys leucogaster were trapped occasionally. Both sites were sandy and dominated by Oryzopsis hyme- noides, Chrysothamnus nauseosus, and Sal- sola kali, although Ambrosia acanthicarpa. Astragalus spp., Atriplex canescens, Gilia hiitchinsifolia, and Hilaria jamesii also were present. The trap design and surveillance methods were described in detail by Garcia et al. (1974). A 10 X 10 (100 traps) grid was wired to a central communications console, where one researcher recorded traps as they were "set off." He then informed an assistant working on the grid where trapped animals were. Animal data were radioed from the as- sistant to the researcher at the console by walkie-talkie. Data were then recorded and the trap reset. Animals seldom were detained in traps for more than a few minutes. Data collected in our study included: species, rela- tive age (juvenile, subadult, adult), sex, repro- 282 September 1980 Jorgensen et al.: Dipodomys Activity Patterns 283 ductive condition, and identification mark (toe clip). Supportive data, used as the inde- pendent variables in our analyses and collect- ed each time an animal was captured, were: ambient temperature, soil temperature, wind speed, cloud cover and lunar events (sub- jectively assessed), light intensity, barometric pressure, and relative humidity. Absolute hu- midities (gm/m^) were determined using the methods described by Piatt and Griffiths (1964), i.e., X = 217(RH) (^'J/IOOT, where T is degrees Kelvin and e^ is vapor pressures in air saturated with water. The period between sundown and sunrise was divided into 20 subperiods, which were used as the time units while plotting activity. Since activity periods changed slightly as day lengths changed, it was necessary to establish a standard set of subperiods before data for different periods could be pooled. Stepwise regression methods were used to assess the ef- fects of independent variables on activity rates among the pooled data for all activity subperiods. Differences in activity among sexes, ages, and reproductive condition class- es were determined using Chi-square tests of independence. An opportunity to assess possible effects of socially dominant individuals in the D. ordii population was noted after three weeks of ac- tivity data had been gathered in 1973. An an- imal was considered dominant if it was the only adult captured within the area pre- scribed when its capture points had been connected, or if it was the only adult repeat- edly caught in a specific trap. Eleven domi- nant individuals among the 42 individuals re- corded on the grid were removed from the poulation and activity data gathered for an additional five days. Temporal activitv pat- terns of the populations before and after the selected removals were compared. Results and Discussion Activity for the intact population of D. or- dii was essentially bimodal, with the greatest activity occurring during the predawn hours c 0) u 0) Q. 9.0 n so- ld T3 •i 70 H a; Q. "I 6.0 CD 0) 5.0- " 4.0-1 < 3.0 2.0- 1.0 - Dipodomys ordii (Dominants included) Dipodomys ordii (Dominants removed ) I I I ■ ■ 7 9 11 13 15 Activity Subperiods 17 19 21 Fig. 1. Activity patterns for Dipodomys ordii pooled among 20 subperiods over a 24-hoiir activity period. 284 Great Basin Naturalist Vol. 40, No. 3 (Fig. 1). These observations agree with Jor- gensen and Hayward (1965), although our early evening peak is not as distinct. Since their data were gathered over the entire year from trapping rates per hour, we question whether their results are directly comparable with ours. Although we have no season-spe- cific data to demonstrate seasonal variations in activity patterns, any variation would pro- duce composite patterns difficult to interpret and compare. Precipitation (11.4 cm) was unusually low in 1971 and rather high (20.5 cm) in 1973. I Q. a o as well as their intra- and interspecific ecologi- cal relationships . Wiens 1976'. Commonly, investigations of arthropod populations are limited to faunistic invento- ries e.g.. Allred et al. 1965. Beck and Allred 1968'. or to broad scale comparisons e.s;.. Al- hed 1973. .\lh-ed and Gertsch 1976, Gertsch and .\llred 1965 1. The present report repre- sents a preliminar\" analysis of one part of an ongoing investigation of lizard ecology" in the Great Basin Desert. Here I will deal with the arthropod food base at a single location, fo- cusing upon the relative success of two sam- pling schemes, seasonal changes in arthropod abundance and diversity", and the apparent degree of local spatial and temporal hetero- geneity' in arthropod populations. Study Site and Methods The research site is in Fairsiew \'allev, Ne- vada, a relatively flat basin ranging in eleva- tion from 1370 to 1500 m. Vegetation within the valley is dominated by Atriplex confer- tifolia, Sarcobatus baileyi, and Oryzopsis hymenoides and generally t\pifies the shad- sc-ale zone common to much of interior Ne- vada (Billings 1949 1. Average monthlv tem- peratures for the site varv" from about C to just over 23 C; average monthly precipitation varies from about .5 cm to just over 1.5 cm: mean growing season is 142 days. Data were analyzed from an insect sam- pling plot estabhshed appro.ximately 9.6 km N of Frenchman, Churchill County, where 49 pitfall traps were arrayed along cardinal compass directions at intervals of 5 m Fig. 1l Traps were randomly located with respect to the vegetation; the total linear distance of each line was 120 m. Each pitfall trap mea- sures 98 mm in diameter by 144 mm in depth: a funnel insert prevents escape of ar- thropods once captured. A Malaise trap Townes 1972) was placed at appro.ximately the center of the two lines iFig. L. Malaise and pitfall traps were opened for a period of 48 hours, followed by a closed peri- od of generally equal duration. This schedule was maintained from 13 May to 24 August 1978 and yielded a new sample at roughly four-day intervals. Each trap contained a standardized amount of 5 percent formalde- hyde solution. The captures of each "arm" of the pitfall traplines for each trapping period were combined. Thus, each of the N-, S-, and W-arm samples contained the contents of 12 traps, and the E-arm sample contained those of 13 traps. Contents of NIalaise trap samples were maintained separately. To examine the major seasonal patterns in the prospective arthropod food base, samples were analvzed for four trapping periods des- ignated 13 May. 12 June. 14 July, and 12 292 September 19S0 Pietruszka: Season.\i, V.\riation of Arthropods 293 Fig. 1. Pitfall and Malaise trap placements. Dots ref>- resent pitfall traps; M represents Malaise trap. Au2;iist. These spanned the major part of the surface-active season for lizards in 1978. Arachnids caught were identified to the or- dinal level: insects caught were identified to family or superfamily where practicable, us- ing Borror. DeLong, and Triplehom J976\ Borror and \Miite (1970), and Chu (1949*. Samples from each arm of the pitfall trap- ping grid were analyzed separately, as were Malaise samples. Total counts of identified groups were then determined for each sample. Arthropod diversity within a sample was assessed using B = 1 — pf. where p, equals the proportion of individuals in cate- gory i ^MacArthur 1972). The relative de- grees of spatial and temporal heterogeneity were assessed using a similarity inde.x, S, where S = l-4(Z|p^-p,.,|) , and p^^, and p,, are the proportions of sam- ples X and y in category i ^Schoener 1970). Results Trapping success.— For the four trapping periods analyzed, the two methods employed amassed a total catch of 7176 arthropods. The vast majoritv of these. 6117. were col- lected along pitfall traplines: 1059 arthro- pods were collected bv Malaise trapping. On a per-trapping period basis. Malaise trapping yielded an average of 265 captures. This is a substantially lower capture rate (up to 50 percent lower than when these traps are used in forested habitats Matthews and Mat- thews 1971). Pitfall traps also \ielded a high- er number of captiu-es per trapping period, averaging 382 per trapline arm, or just over 1500 captures per trapping grid. However, capture rates for the two methods are not di- rectly comparable due to the greater "at risk" area for pitfall traps. The composition of collections obtained bv the t^vo methods also differed substantially. Pitfall collections were dominated bv mites (Acarina>, which comprised almost half the total collection. These were followed, in or- der of numerical importance, by H\Tnenoi>- tera (the vast majority- of which were ants), Coleoptera. and Diptera. In all, largely or completely terrestrial forms comprised ap- proximately 88 percent of the arthropods col- lected. Malaise trap collections, by c-ontrast were dominated by Diptera. which formed over 70 percent of the total collection. H\"menoptera, Homoptera. Lepidoptera. and Coleoptera combined to form just over one- quarter of the total collection Table 1). These data are consistent with those from Malaise traps used in forested areas in which Diptera, H\"menoptera. Hemiptera including Homoptera'. and Lepidoptera constitute at least 90 percent of each collection (Matthews and Matthews 1971). Seusorwl clianges in abundance and diver- sity.— Arthropod abundance appeared to peak during mid-June, approximately 1.5 months after the last of the spring rains. Numbers of arthropods declined rapidly thereafter to moderate levels. This seasonal trend is closely reflected in collections from pitfall traps but not from Malaise trapping (Fio. 2.\\ The low number o* captures on 12 June mav reflect an actual decrease in aerial insects, but it is more Ukely that this is a re- flection of local changes in wind conditions, to which this technique is highly susceptible ^Matthews and Matthews 1971). This inter- pretation is strengthened by the observation that aerial insects occurred in approximately equal numbers in the 12 June and 14 July Malaise samples. 294 Great Basin Naturalist Vol. 40, No. 3 Table 1. Siiminary of arthropods collected in pitfall and Malaise trap samples. Pitfall trap Malaise trap Acarina Araneida Scorpion ida Solpiigida Coleoptera Carabidae Collembola Diptera Hemiptera Homoptera Taxa Number of specimens Relative abundance Number of specimens Relative abundance 3051 0.4988 Anthribidae Buprestidae Curculionidae Dascillidae Histeridae Leiodidae Melyridae Nitidulidae Pedilidae Staphylinidae Tenebrionidae Sminthuridae Poduridae Anthomyiidae Bibionidae Bombyliidae Calliphoridae Cecidomyiidae Chironomidae Conopidae Dolichopodidae Empididae Lauxaniidae Muscidae Mycetophilidae Pipunculidae Psychodidae Ptychopteridae Sarcophagidae Sciaridae Simuliidae Syrphidae Tachinidae Therevidae Tipulidae Xylophagidae Acalypterate muscoids Diptera larvae Lygaeidae Miridae Nabidae Pentatomidae Tingidae Aphididae Cercopidae Cicadellidae Coccoidea Nymph /larvae 104 0.0170 16 0.0026 2 0.0003 30 0.0049 17 0.0028 5 0.0008 2 0.0003 5 0.0008 2 0.0003 244 0.0399 1 0.0002 1 0.0002 3 0.0005 243 0.0397 38 0.0062 7 0.0011 24 0.0039 1 0.0002 3 0.0005 1 0.0002 37 0.0060 18 0.0029 1 0.0002 8 0.0013 4 0.0007 0.0002 1 0.0002 3 0.0005 45 0.0074 1 0.0002 18 0.0029 1 0.0002 42 0.0069 2 0.0003 84 0.0137 18 0.0029 2 0.0003 2 0.0003 1 0.0002 10 0.0016 5 0.0008 102 0.0167 2 0.0003 1 0.0002 4 29 3 5 197 5 1 4 11 3 9 41 203 21 1 1 3 248 1 60 11 0.0009 0.0057 0.0019 0.0009 0.0019 0.0009 0.0038 0.0274 0.0028 0.0047 0.1860 0.0047 0.0009 0.0038 0.0104 0.0028 0.0085 0.0009 0.0387 0.1917 0.0198 0.0009 0.0009 0.0028 0.2342 0.0028 0.0038 0.0009 0.0567 0.0104 September 1980 Pietruszka: Seasonal Variat ION OF Arthropods 295 Table 1 continued. Pitfall trap Malaise trap Hymenoptera Andrenidae 6 0.0010 Apidae 21 0.00.34 1 0.0009 Braconidae 13 0.0123 Chalcidoidea 51 0.0083 87 0.0822 Chrysididae 6 0.0010 Dryinidae 2 0.0003 Fonnicidae 1577 0.2578 13 0.012.3 Halictidae 66 0.0108 5 0.0047 Ichneumon idae 1 0.0002 1 0.0009 Mutillidae 5 0.0008 Pom pil idae 4 0.0007 2 0.(K)19 Sphecidae 46 0.0075 3 0.0028 Lepidoptera Cosmopterygidae 19 0.0031 38 0.0359 Lycaenidae 1 0.0002 Pyralidae 5 0.0008 4 0.00,38 Unidentified larvae 12 0.0020 Neuroptera Chrysopidae 1 0.0009 Coniopterygidae 3 0.0005 1 0.0009 Henierobiidae 18 0.0029 1 0.0009 Myrmeleontidae 17 0.0028 Unidentified larvae 3 0.0005 Orthoptera Acrididae 2 0.0003 Blattidae 2 0.0003 Gryllacrididae 18 0.0029 1 0.0009 Mantidae 1 0.(XK)2 Thysanoptera Heterothripidae 4 0.(XK)7 6 0.0057 Phlaeothripidae 7 0.0011 Thripidae 9 0.0015 Unidentified larva (campodeiform) 1 0.(K)02 Isopoda 1 0.0002 Among the major arthropod groups occur- ring in seasonal samples, mites most closely follow the general trend. It is quite likely, in fact, that mite populations are the major fac- tor underlying the observed seasonal pattern. The much greater abundance of mites tends to mask other groups, such as the Hymenop- tera, Diptera, and Coleoptera, which tend to remain at low to moderate levels of abun- dance throughout the season (Fig. 2B). Diversity values based on pitfall trapping and Malaise trapping, respectively, were sub- stantially different from one another during all trapping periods of the 1978 season (Fig. 3). Among pitfall samples arthropod diversity is generally low due to the high abundance of both mites and ants (see Table 1). The trend toward increa.sing diversity reflects the rela- tive decrease in mite abundance in late .sea- son samples. Among Malaise samples diver- sity values reflect, in part, the lower total catch afforded by this method. Trapping dates with the highest diversity values, 12 June and 12 August, had catches that were approximately 28 and 8 percent, respective- ly, of the catches for the remaining trapping periods. These samples contained fewer taxa more equitably represented, yielding greater apparent diversity. It is likely that the values for 12 June and 12 August Malaise samples are inordinately high due to the sensitivities of the technique mentioned earlier. Never- theless, that there should be relatively greater diversity of aerial insects seems reasonable, if for no other reason than their greater mobility. Spatial and temporal heterogeneity of ar- thropods— As mentioned above, spatial and temporal variation in arthropod abundance may affect not only characteristics of individ- ual consumer behavior, but also the ecologi- cal relationships within and between species. As an approach to spatial variation on a rela- tivelv small scale (minimum area effect of the 296 Great Basin Naturalist Vol. 40, No. 3 8-1 Hymenoptera 13 May 12 June 14 July 12 August Fig. 2. Seasonal changes in arthropod abundance: a, as reflected by pitfall and Malaise trap samples; b, sea- sonal changes in the major taxa of these samples. trapping grid is probably on the order of 1.4 ha), I calculated similarity values for all pos- sible combinations of grid arms for each trapping period. An average similarity value (S) was then obtained as a measure of the overall spatial heterogeneity over the trap- ping grid. Immediately apparent from this analysis is the high degree of similarity (low heterogeneity) between grid arms at all trapping periods (range = .730-.893, Fig. 4). Yet, there does appear to be a trend toward increasing arthropod patchiness with decreas- ing abundance levels. The trend is not statis- tically significant, however, based upon these data. 13 May 12 June 14 July 12 August Fig. 3. Seasonal changes in arthropod diversity as re- flected in pitfall and Malaise trap samples. Pitfall diver- sities represent the average diversity for the four grid arms at each sampling date. Though evidence for spatial variation in arthropod numbers during 1978 is meager, temporal variation is much more apparent. Average similarity values, based upon all pos- sible comparisons of each grid arm over all trapping dates, were substantially lower than for spatial variation: N-arm = .621; E-arm = .648; W-arm = .672; S-arm = .692. These data indicate a substantial change in the arthropod fauna throughout the active season over a relatively small area. Com- parisons of Malaise trap collections support this interpretation (S = .384), but, as men- tioned above, the collections for two of the dates may be suspect. Discussion Desert habitats are characterized by both cyclic and unpredictable climatic changes on micro- as well as macrogeographic scales (Cloudsley-Thompson 1968, Logan 1968). As a result, these habitats are typified by periods of pulsed production. It is particularly note- worthy that in the first four months of 1978 Fairview Valley received 255 percent greater than normal rainfall; average temperatures between April and August were below nor- mal (U.S. Weather Bureau data). Such a com- bination of climatic events may have provid- ed for a longer than normal production pulse, resulting in a marked increase in arthropod abundance throughout the season. The nu- merical dominance by mites and ants of col- September 1980 Pietruszka: Seasonal Variation of Arthropods 297 12 Aug 14 July 13 May 0.5 15 25 Arthropods/ Trapping Period ( x 10^) Fig. 4. Spatial heterogeneity of arthropods as reflect- ed in the average similarity, S, between pitfall trap grid arms at each sample date. lections spanning the entire season seems to argue for this possibiHty. Moreover, it has been suggested that at extremely high popu- lation levels habitat patchiness will be re- duced, and localized areas may even become unifonn in their species distributions (Wiens 1976). Indeed, this seems to be what occurred in 1978. If this hypothesis is correct, then ar- thropod patchiness would be predicted to be more apparent during dry years when abun- dance levels are low. The data to test this prediction have been gathered (for 1979, a substantially drier year) but have not yet been analyzed. Finally, it is clear from these analyses that to effectively monitor arthropod populations no single methodology is sufficient. The com- binati ai of pitfall and Malaise trapping pro- vides a reasonable balance of terrestrial and aerial forms from desert habitats. Never- theless, specific situations and goals will ulti- mately determine the techniques to be used. Literature Cited Allred, D. M. 1973. Scorpions of the National Testing Station, Idaho. Great Ba.sin Nat. 33:251-254. Allred, D. M., and VV. J. Gertsch. 1976. Spiders and scorpions from northern Arizona and southern Utah. J. Arachnol. 3:87-99. Allred, D. M., D. E. Johnson, and D. E. Beck. 1965. A list of some beeflies of the Nevada Test Site. Great Ba.sin Nat. 25:5-11. Beck, D. E., and D. M. Allred. 1968. Faunistic inven- tory-BYU ecological studies at the Nevada Test Site. Great Basin Nat. 28:132-141. Billings, W. D. 1949. The shadscale vegetation zone of Nevada and eastern Galifornia in relation to cli- mate and soils. .\mer. .Midi. Nat. 42:87-109. BoRROR, D. J., D. .M. DeLonc, and C. A. Triplehgrn. 1976. .\n introduction to the study of insects. 4th ed. Holt, Rinehart and Winston, New York. 852 pp. BoRROR, D. J., and R. E. White. 1970. A field guide to the insects of .\merica north of Mexico. Hough- ton Mifflin Go., Bo.ston. 404 pp. Ghu, H. F. 1949. How to know the immature insects. Wm. G. Brown Go. Publishers, Dubuque, Iowa. 234 pp. Gloudsley-Thompson, J. L. 1968. The Merkhivat Je- bels: a desert community. Pages 1-20 in G. VV. Brown, ed.. Desert biology. .Academic Press, .New York. Fautin, R. W. 1946. Biotic communities of the Northern Desert Shrub Biome in western Utah. Ecol. Mon- ogr. 16:251-310. Gertsch, W. J., and D. M. .-Vllred. 1965. Scorpions of the Nevada Test Site. Brigham Young Universitv Sci. Bull., Biol. Ser. 6(4): 1-15. Gill, F. B., a.nd L. L. Wolf. 1977. Nonrandom foraging by sunbirds in a patchy environment. Ecolog\' 58:1284-1296. Logan, R. F. 1968. Gauses, climates, and distribution of deserts. Pages 21-50 in G. W. Brown, ed.. Desert biology. .Academic Press, New York. Mac.\rthur, R. H. 1972. Geographical ecolog)'. Harper and Row, New York. Mac.\rthur, R. H., and E. R. Pianka. 1966. Optimal use of a patchy environment, .^mer. Nat. 100:60.3-609. Matthews, R. W., and J. R. Matthews. 1971. The .Ma- laise trap: its utility and potential for sampling insect populations. Michigan Entomologist. 4:117-122. Schoener, T. W. 1970. Nonsynchronous spatial overlap of lizards in patchy habitats. Ecology 51:408-418. Townes, H. 1972. A light-weight Malaise trap. Ent. News 83:2.39-247. Wiens, J. A. 1976. Population responses to patchy envi- ronments. .\nn. Rev. Ecol. Svst. 7:81-120. NOTICE TO CONTRIBUTORS Original manuscripts in English pertaining to the biological natural history of western North America and intended for publication in the Great Basin Naturalist should be directed to Brigham Young University, Stephen L. Wood, Editor, Great Basin Naturalist, Provo, Utah 84602. Those intended for the Great Basin Naturalist Memoirs should be similarly directed, but these manuscripts are not encumbered by a geographical restriction. Manuscripts. 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Reprint Schedule of the Great Basin Naturalist 2 pp. 4 pp. 6 pp. 8 pp. 10 pp. 12 pp. Each 100 copies $20 $24 $28 $32 $36 $40 additional 200 copies 28 32 36 40 44 48 2 pp. 300 copies 36 40 44 48 52 56 $4 Great Basin Naturalist Memoirs ). 1 The birds of Utah. By C. L. Hayward, C. Cottam, A. M. Woodbury, H. H. Frost. $10. ). 2 Intermountain biogeography: a symposium. By K. T. Harper, J. L. Reveal et al. $15. ). 3 The endangered species: a symposium. $6. ). 4 Soil-plant-animal relationships bearing on revegetation and land reclamation in Nevada deserts. $6. TABLE OF CONTENTS Spatiotemporal variation in phenolog)' and abundance of floral resources on short- CTrass prairie. V. J. Tepedino and N. L. Stanton 197 Dog owners and hvdatid disease in Sanpete County, Utah. Peter M. Schantz and Ferron L. Andersen 216 New grass distribution records for Arizona, New Mexico, and Texas. Stephan L. Hatch 221 A comparison of epiphvtic diatom assemblages on living and dead stems of the com- mon grass Phragmites atistralis. Judith A. Grimes, Larry L. St. Clair, and Sam- uel R. Rushforth 22.3 Poisonous plants of Utah. Jack D. Brotherson, Lee A. Szyska, and William E. Even- son 229 The successional status oi Cupressus arizvnica. Albert J. Parker 254 A self-pollination experiment in Pintis edulis. Ronald M. Lanner 265 Comparative floral biology of Penstemon eatonii and Penstemon cyananthus in cen- tral Utah: a preliminar\^ study. Lucinda Bateman 268 Differential habitat utilization by the sexes of mule deer. Michael M. King and H. Duane Smith 2/ -3 Temporal activitv patterns of a Dipodomys ordii population. Clive D. Jorgensen, H. Duane Smith, and James R. Garcia 282 New records of western Trichoptera with notes on their biolog). Bernard G. Sweg- man and Leonard C. Ferrington, Jr 287 Observations on seasonal variation in desert arthropods in central Nevada. Robert D. Pietniszka 292 HE GREAT BASIN NATURALIST ^ lume 40 No. 4 December 31, 1980 Brigham Young University MUS. COMP. ZOO' LIBRARY ^MM f^ GREAT BASIN NATURALIST Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Life Science Museum; Stanley L. Welsh, Botany; Clayton M. White, Zoology. Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agricul- tural Sciences; Ernest L. Olson, Director, Brigham Young University Press, University Editor. The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been published from one to four times a year since then by Brigham Young University, Provo, Utah. In gener- al, only previously unpublished manuscripts of less than 100 printed pages in length and per- taining to the biological and natural history of western North America are accepted. The Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natu- ral history longer than can be accommodated in the parent publication. The Memoirs appears irregularly and bears no geographical restriction in subject matter. Manuscripts are subject to the approval of the editor. Subscriptions. The annual subscription to the Great Basin Naturalist is $12 (outside the United States $13). The price for single numbers is $4 each. All back numbers are in print and are available for sale. All matters pertaining to the purchase of subscriptions and back num- bers should be directed to Brigham Young University, Life Science Museum, Provo, Utah 84602. The Great Basin Naturalist Memoirs may be purchased from the same office at the rate indicated on the inside of the back cover of either journal. Scholarly Exchanges. Libraries or other organizations interested in obtaining either journal through a continuing exchange of scholarly publications should contact the Brigham Young University Exchange Librarian, Harold B. Lee Library, Provo, Utah 84602. Manuscripts. All manuscripts and other copy for either the Great Basin Naturalist or the Great Basin Naturalist Memoirs should be addressed to the editor as instructed on the back cover. 5-81 650 50104 ISSN 0017-3614 The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 Volume 40 December 31, 1980 IMPACT OF THE 1975 WALLSBURG FIRE ON ANTELOPE BITTERBRUSH {PURSHIA TRIDENTATA) Fred J. W'agstaff .\bstract.— Antelope bitterbrush {Pur.sliia tiidentcita) is a preferred browse species that is susceptible to decreases ill population density due to fire. The reduction in density of this species due to fire was determined bv sampling areas within and adjacent to the burn. The 1975 burn caused a significant reduction in the population density of bitterbmsh. It \yas also determined that rate of growth was lower for plants within the burn. In the summer of 1975 a fire burned sever- al himdred acres of mule deer winter range in Wasatch County, Utah. The burned area includes the area from the junction of the Wallsburg road southeast to the crest of the west Daniels Canyon ridge and to the north- east along Highway 40 to near the Midway Junction. Deer Creek Reservoir is just across the highway to the northwest of the burned area. The study plots are near the northeast cor- ner of the burn in an area where a population density of bitterbrush was great enough to permit quantitative analysis of the response of this species to fire. Burned and imburned areas were studied along with some islands that escaped burning. Bitterbrush {Purshia tridentata) was selected as an indicator spe- cies because of its status as a preferred brow.se plant on mule deer winter ranges. Any factor that causes significant changes in the structure of the plant communities on winter ranges is of concern. One of the most significant agents known is fire. Fire has oc- curred naturally since time began and is a major factor in determining the .structure of many plant communities. In other commu- nities, man-caused fire has introduced an agent of change that has modified vegetation over large areas. In a situation where prefire structure is so important, the impact of fire on structure of the plant community should be known. Will plants be killed? How long will the impacts last? Will value of the area as mule deer win- ter range be completely lost? Can deer move to another winter range? These and other questions occur and need to be answered if the impacts of fire are to be understood. The area chosen for study has been burned several times in the last one hundred years. Fire has occurred at irregular times and over different portions of the area. This has led to a mosaic of vegetation types and age struc- tures. The 1975 fire was much larger than most of the past fires and affected a signifi- cant portion of mule deer winter range in the area. With passage of four years, the in- ception of the postfire plant succession should be identifiable. Three major hypotheses were fornuilated to determine some of the relative impacts of the 1975 fire. First, the density of bitterbmsh had been significantly reduced by the fire. Secondly, use by mule deer is less in the burned area. Third, the shift in deer u.se has 'U.S. Forest Service Interniountain Forest and Range Experiment Station Sl.nib Sciences Laboratory, 735 North 500 East. Provo. Utah 84601. 299 3()0 Great Basin Naturalist Vol. 40, No. 4 had a detrimental effect on surviving bitter- bnish plants. Literature Review Literature pertaining to bitterbrush is ex- tensive, with over 200 references identified. The following references support the results of the study. FiirsJiid tridcntata (antelope bitterbrush) is highly desirable as browse on deer winter range (Bissell et al. 1955, Giunta et al. 1978, Hoskins and Dalke 1955, Julander 1952, Leach 1956, Longhurst et al. 1952, Mace 1957, Reynolds 1960, Smith et al. 1954, Smith 1952). Since bitterbmsh is so highly preferred, it can be used as an indicator spe- cies for use on an area bv game animals and game winter range conditions. There have been numerous articles written about the impacts of fire on bitterbrush (Blaisdell 1950, 1953, Blaisdell and Mueggler 1956, Countryman and Cornelius 1957, Fer- guson and Basile 1966, Komarek 1965, Miller 1963, and Pechanec et al. 1954). They have determined that browse production of bitter- brush plants that have been burned has lagged behind miburned control plants for several years. Blaisdell (1950) also showed that relative densities of bitterbrush in burned and imbiuned areas differed signifi- cantly. Nord (1965) developed data that dem- onstrates the existence of fire-resistant eco- types where most plants in the population resprout after fire. Blaisdell (1953) and others have shown that variables of fire intensitv, fuel loading, and soil moisture affect re- sprouting. Even nonsprouting types will have some survivors, particularly where the fire does not burn intensely. Results The following data were collected from six 100 ft- quadrats in the burned and adjacent unburned area. Two of the quadrats were at the lower edge of the burn in the sagebrush- bitterbrush tvpe and four near the middle of the burn in the oak-sagebrush tvpe. The rela- tive density of live bitterbrush plants is shown in Table 1. There is a striking difference between the nimiber of living plants in the burned and un- burned areas. None of the burned plots had anv surviving old plants because the fire was intense enough to kill the tops of all bitter- brush plants. It appears the fire mav have been hotter at the lower part of the burn be- cause there were no relic bitterbnish plants. In the midslope plots, relics were found for most shrubs of various species. All the young bitterbrush plants in the burned area were from resprouting crowns. There were voung plants in all the unburned plots, indicating the species is successfully re- producing in the study area. In Table 2, the data collected on the current-year twig growth are presented. Twenty twigs per plant on 20 plants (200 twigs in burned areas and 200 in unbiu'iied) were measured to determine if there was anv difference in current-year growth. The differ- ence in twig growth was found to be statistic- ally significant at the 90 percent level. In other words, the burned plants were growing at a slower rate. For each of the six plots, mule deer fecal pellet groups were counted. All pellet groups were counted without regard to pellet age. Clearly, the unl)urned areas have received heavier use than the adjacent burned areas, as shown in Table 3. Table L Relative density of bitterbrush plants on six plots. Burned Slope location Bitte rbrush plants Quadrat New Old Dead Total 1 Yes Mid 2- 22' 24 2 Yes Mid f-i- 22' 28 3 No Mid (i 15 1 22 4 No Mid 9 15 1 24 5 Yes Lower 2- 2 6 No Lower 3 10 13 Includes relics probably killed by fire. Includes resprouting after fire. December 1980 Wagstaff: Impact of Fire ox Bitterbrush 301 Takle 2. 1979 lcn<4tli (if tuitrs on 20 liitterhrush plants. Table 3. Number of mule deer fecal pellet groups for each of six plots. Unhiirned area plants Aveniije Plant twig length mmiber (inches) Burned area plants Average Plant twig length number (inches) 9.5 1 9.3 2 9.4 3 7.05 4 10.15 5 10.05 6 9.8 7 6.55 8 8.45 9 9.9 10 9.02 y Discu: SSION 9.2 7.75 9.55 7.6 6.95 7.45 8.45 9.1 6.8 7.15 8.0 There are several obvious differences be- tween the burned and unburned plant popu- lations. It is evident the fire had a detrimen- tal impact on the density of bitterbrush plants and production of this component of the plant community. A relatively low per- centage of the bitterbrush plants were able to resprout and, therefore, most burned area plants were eliminated from the community. Sagebnish was also largely absent in the burned areas, but in adjacent unburned areas it was a significant part of the community. The marked change in the community spe- cies composition caused by the fire is still very much in evidence. Also the size of the browse plants has been altered to the point that much of the burned area vegetation would be totally covered by several inches of snow. This physical barrier would limit use of the burned area even though considerable forage may occur there. Another obvious factor is the difference in occurrence of fecal pellet groups between burned and adjacent unburned areas. Since the measurements were taken in adjacent areas, the reason for the significant difference seems easiest to explain on the basis of physi- cal availability of browse diuing the winter. If there are no significant differences in nu- tritive value or availability, one would expect essentially equal use near the boundary of the two areas. The difference in use must occur because of an absolute difference in the amount of browse available, which was prob- ably compounded by snow coverage. Plot Slope position Number of pellet groups Mid Burned 8 Mid Burned 5 Mid I'nburned 28 Mid Unburned .36 Low Burned 4 Low Unburned 16 All the bitterbrush plants that were exam- ined during this study exhibited substantial twig growth and appeared to be vigorous and healthy. Growth form of the old plants in the unburned areas showed a fairlx' open fcjriu that indicates little use by sheep and/or deer. These plants did not exhibit the clubhead form indicative of sustained heavy use; nei- ther was there evidence of browsing on large- diameter twigs. Growth of the Ijitterbrush plants in un- burned areas was greater, as has been docu- mented in other areas by Blaisdell and Mueggler (1956). This difference in the growth rate is expected to continue for sever- al years. In terms of total production of hit- terbnish forage, the burned area has lapsed considerably since the fire and will most likely continue to do .so for a long time. The density of plants has been reduced, as well as the size of plants. There are fewer plants of smaller size in the burned areas than were there prior to the fire. Since no evidence of new seedlings could be foinid, it is reasonable to conclude the burned area production will lag for nianv years. Deer herd populations are often directly tied to winter range availability and condi- tion because it is the element most often in shortest supplv. It is not known whether this is the case with the mule deer using the study area. There is need for concern because deer numbers are increasing and winter range in the area is decreasing due to changing land ase. Both of these trends cannot continue in- definitelv without deer niunhers reaching the limit of available winter range. CONCLL SIO.NS It is easv to conclude that the 197.5 fire was detrimental to bitterbnish, but it is not 302 Great Basin Naturalist Vol. 40, No. 4 easy to conclude that the numbers of deer have been reduced because of it. This is due to populations of deer being within the ca- pacity of the remaining winter range. The area is neither producing the winter forage for deer that it was prior to the burn nor are deer numbers anywhere near historic high levels. If this area is indeed a critical winter range when population numbers are larger and other factors holding populations down are temporary, additional concerns arise. Thought should be given to introducing a fire-tolerant ecotype of bitterbnish with the hope of hybridizing this trait into the com- munity, and effective means of reducing fire occurrence and spread should be developed. Literature Cited BissELL, H. D., B. Harris, H. Strong, and F. James. 1955. The digestibility of certain natural and arti- ficial foods eaten by deer in California. Calif. Fish and Game 41(1)1.57-78. Blaisdell, J. p. 1950. Effects of controlled burning on bitterbni.sh on the upper Snake River plains. U.S. For. Serv., Int. For. and Range Exp. Sta. Res. Pap. 20, .3 pp. 19.53. Ecological effects of planned burning of sagebnish-grass range on the upper Snake River plains. U.S.D.A. Tech. Bull. 1075, .39 pp., illus. Blaisdell, J. P., and W. F. Mueggler. 1956. Sprouting of bitterbnish (Purshia tridentata) following burn- ing or top removal. Ecology 37:.365-370, illus. Countryman, C. M., and D. R. Cornelius. 1957. Some effects of fire on a perennial range type. J. Range Manage. 19:39-41, illus. Ferguson, R. B., and J. V. Basile. 1966. Topping stiniu lates bitterbrush twig growth. J. Wildl. .Manage. .30(4): 8.39-841. Giunta, B. C, R. Steve.ns, K. R. Jorge.nsen, and A. P. Plummer. 1978. Antelope bitterbrush— an impor- tant wildland shrub. Utah State Div. Wildl. Re- sources Publ. 78-12. HosKiNS, L. W., AND p. D. Dalke. 1955. Winter browse on the Pocatello big game range in southeastern Idaho. J. Wildl. Manage. 19:21.5-225. JuLA.NDER, O. 1952. Forage habits of mule deer during the late fall as measured by stomach content analyses. U.S. For. Serv. Int. For. and Range Exp. Sta. Res. Note-2. 5 pp. KoMAREK, R. 1965. Fire and changing wildlife habitat. Proc. Tall Timbers Fire Ecol. Conf. 2:35-43. Leach, H. R. 1956. Food habits of the Great Basin deer herds of California. Calif. Fish and Game 43(4):243-308, illus. LoNGHURST, W. M., A. S. Leopold, and R. F. Das.ma.nn. 1952. A survey of California deer herds, their ranges and management problems. Calif. Dept. Fish and Game Bull. 6. 1.36 pp., illus. Mace, R. LI. 1957. Oregon's mule deer. Oregon State Game Comm. Wildlife Bull. No. 3. 25 pp. Miller, H. A. 1963. Use of fire in wildlife management. Proc. Tall Timbers Fire Ecol. Conf. 2:19-30. NoRD, E. C. 1965. Autecology of bitterbnish in Califor- nia. Ecol. Monogr. .35:307-.3.34. Pechanec, J. F., G. Stewart, and J. P. Blaisdell. 1954. Sagebrush burning— good and bad. U.S.D..\. Farmer's Bull. 1948. .34 pp. Reynolds, T. A., Jr. 1960. The mule deer— its history, life history, and management in Utah. Utah State Dept. Fish and Game Inform. Bull. 60-4. .32 pp. Smith, \. D., and R. Hubbard. 1954. Preference ratings for winter deer forages from northern Utah ranges based on browsing time and forage con- sumed. J. Range Jv-lanage. 7(6):262-265. SxuTii, J. G. 1952. Food habits of mule deer in Utah. J. Wildl. Manage. 16(2): 148-155, illus. TERRESTRIAL VERTEBRATE FAUNA OF THE KAIPAROWITS BASIN N. Diiane Atwood', Clyde L. Pritchctt', Richard D. Porter', and Benjamin W. Wood' .\bstr^ct.- This report inehides data collected during an investigation by Brighani Young University personnel from 1971 to 1976, as well as a literature review. The fauna of the Kaiparowits Basin is represented by 7 species of mnphihians (1 salamander, 5 toads, and 1 tree frog), 29 species of reptiles (1 turtle, 16 lizards, and 12 snakes), 183 species of birds (plus 2 hypothetical), and 74 species of mammals. Geographic distribution of the various species within the basin are discussed. Birds are categorized according to their population and seasonal status. .Avian habitat relationships are discussed, and extensions of range are reported for 5 species of birds. Three threatened or endan- gered avian species occur in the basin. Four avian species seem to have declined significantly in numbers in recent \ears. The early activities and exploration of trappers, missionaries, and government sur- vey workers provided little information to om- knowledge of fauna in the Kaiparowits Basin. Most of these early expeditions skirted around the basin on all sides or were con- fined to the depths of the Colorado River Canyon. The Domingues-Velez de Escalante party traveled along the north side of the Colorado River between Lee's Ferry and the well-known Crossing-of-the-Fathers. During this portion of their journey, it became neces- sary for the party to eat their horses (Auer- bach 1943). No mention is made in Father Escalante's journal of any fauna observed. Records of Powell's expeditions of 1869-1870 and 1871-1872 indicate observations of wild- life were limited to the larger game animals, probably those that could be used for food. In 1892 the American Museum of Natural History sent an expedition into the San Juan region just east of the Kaiparowits Basin (Al- len 1893). Wetherill, Flattum, and Sterns (1961) made a trip by boat up the Colorado River from Lee's Ferry to Rainbow Bridge. Both expeditions recorded the animals ob- served. The number of scientific investigations af- ter those of Powell into the Glen Canyon were few indeed (Crampton 1959). The Na- tional Park Service recognized this fact and, in measure, filled some of the gaps by send- ing out an expedition that descended by boat the San Juan and Colorado Rivers to Rock Creek. From this point, they spent about two weeks on the southwest end of the Kaiparo- wits Plateau. The results of this work were reported by Hall (1934). Others such as Gregory (1917, 1938, 1945, 1947, 1948) and Gregory and Moore (1931) have made sub- stantial contributions to our knowledge of the Colorado River drainage, particularly with respect to the geology. Observations and comments were also recorded regarding the biota. Members of the Department of Zoology at Brigham Young University' (BYU) initiated a .series of biological investigations of the up- per Colorado River Basin; of these, the fol- lowing were within the Kaiparowits Basin: In July 1927, a group visited Lee's Ferry and Bryce Canyon, and in 1936. a party of four biologists spent 810 man hours in the Esca- lante River drainage. In the fall of 1937, V. M. Tanner and C. L. Hayward studied in the Paria Valley. A party headed by D. E. Beck entered the region in the fall of 1938 and 1939, exploring the course of the Escalante River. Beck also .spent the last part of May and earlv June of 1940 in the area east of Willow Tank Spring down to the Escalante River. In 1946, 1952, and 1953, other BYU expeditions entered the area in both Kane and Garfield Counties. The Navajo Mountain and Wahweap Drainage were visited in 1955 and 1958 (Hayward et al. 1958). 'U.S. Forest Service, Provo, Utah 84601. ■Department of Zoology, Brigham Young I'niversity. Prove, Utah 84602. U.S. Fish and Wildlife'Service, Provo, Utah 8460r(.325 North .300 West. Mapleton, Utah 84663). 'Department of Botany and Range Science. Brigham Young University, Provo, Utah 84602. 303 304 Great Basin Naturalist Vol. 40, No. 4 11 ,; .A 30/ ^^J' V 7 Aa;^'^ ;|smok* NfV^j -i ,• , Ky'x \ ^^'vA 1/ S ,v ^-x^;^ ;, NIPPIE BENCH "^^^ 0' ...^ Fig. 1. Map of Brigham Young University Xavajo-Kaiparowits study sites, Kane and San Juan Counties, Utah, and Coconino Count\. Arizona. Benson (1935) published an important pa- per on the fauna and flora collected during an expedition led by Alexander, Kellogg, and Benson in the Navajo Mountain region in 1933. They had six stations as follows: one five miles .south of the mountain, 10-12 June; one at War God Spring on top of the moun- tain at 8400 feet, 13-20 June; one at Bridge Canyon, 21-24 June, two on the mesa south of the mountain; and the other near Navajo Mountain Trading Post, 25-26 June. In addi- tion, naturalists assigned to Bryce Canvon National Park have, since 1932, made contri- butions toward our knowledge of the fauna and flora of the region. Woodbury and Russell (1945), in their comprehensive report on the birds of the Navajo countrv, presented data on specimens collected and observations made in Glen Canvon, on Navajo Mountain, and on Kai- parowits Plateau. Behle and three associates made a boat trip down the Colorado River 13-17 April 1947. They made observations at various places between Hite and Lee's Ferry, December 1980 Atwood et al.: Kaiparowits Vertebrates 305 including the mouth of the Escalante River, river mile 88; Hidden Passage, river mile 76; Aztec Creek, river mile 68.5; Crossing-of-the- Fathers; near Creek, river mile 40.5; and Lee's Ferry. Behle and associates also collect- ed birds at or near the confluence of Calf Creek and the Escalante River, 11-14 June 1953, and 7-9 May 1954 (Behle et al. 1958; Behle 1960). In 1958, Behle and Higgins (1959) made some observations at Hole-in- the-Rock (20 October), and the confluences of Kane Creek and the Escalante River with the Colorado River (19 October); birds were observed by Harold Higgins and Gerald Smith on a river trip extending from 1 Julv to 9 August 1958 from Hite to Lee's Ferry (Behle and Higgins 1959). We have not given the precise dates in the species accounts giv- en below of birds collected or seen bv Behle and his associates on June 1953 and Mav 1954 trips, or for Benson's (1935) trips in 1933, becau.se of the short duration of their investigations. If desired, such can be obtain- ed from the literature. The dates of observa- tions from our investigations, however, are reported herein. In 1957, the National Park Service in- itiated a salvage program for the Glen Can- \"on area preparatory to the constniction of Glen Canyon Dam. These data were pub- lished by the University of Utah and the Mu- seum of Northern Arizona in their respective journals. Earlier (as a result of a reconnais- sance trip, 8-14 August 1957, and as a part of tlie same project), Woodbury et al. (1959) had prepared an annotated checklist of the birds of the proposed Glen Canvon Reservoir area from various sources, including pub- li.shed and unpublished manuscript records. In addition, Behle and Higgins (1959) pub- lished a number of previouslv impubli.shed bird observations from Woodburv and Rus- sell's field notes from the Rainbow Bridge- Monument Valley expedition. In June 1971, Brigham Young University and Northern Arizona University initiated the Environmental Impact Studies for the Navajo and later in 1972 for the proposed Kaiparowits Generating Stations. Extensive collections and observations of the terrestrial vertebrate fauna and flora have been made by these groups. This report on the terrestrial vertebrate fauna is based on species reported in the lit- erature and collections or field observations made by BYU personnel from 1971 to the fall of 1976. The various taxa in the annotated lists are arranged phylogenetically following the order given in Schmidt (1953) for the am- phibians and reptiles; the .\merican Ornithologists Union (AOU) Check List, Fifth Edition (1957) and subsequent supplements for the bird.s; and Hall and Kelson (1959) for the mammals. Most of the studies dealing with the distri- bution and ecology of amphibians and rep- tiles within the Kaiparowits Basin have l^een restricted to areas along the Colorado River. The stretch of river generally known as Glen Canyon Gorge has been the area most com- monly studied. The most comprehensive list (28 taxa) of amphibians and reptiles for this area was by W. W. Tanner (1958a). Wood- bury et al. (1959) published a similar list, but it contained only 19 ta.xa. Both of these stud- ies were designed to obtain data prior to the constrution of Glen Canyon Dam and sub- sequent development of Lake Powell. .-Xs part of the Navajo-Kaiparowits environmental baseline studies. Toft (1972) prepared a field key ba.sed on the above literature and field observations and collections made during the summer of 1971 and 1972. .\dditional data have been added from subsecjuent studies made during the period of 1973-1975. These studies represent to date the most extensive field studies into the interior of Kaiparowits Basin. Amphibians Ambystomidae (Salamanders) Ambystoma trigrimim nehulosum Hallow- ell. Utah Tiger Salamander. V. NL Tanner (1930) and Weight ^1932) Bryce Canvon. W. W. Tanner (1975). Pelobatidae (Spadefoot Toads) Scaphiopus hammondi Baird. Hammonds Spadefoot Toad. V. M. Tanner (1930) and Weight (19.321 Bryce Canvon. Scaphiopus intermontanus Cope. Great Basin Spadefoot Toad. V. M. Tanner (1940a) 306 Great Basin Naturalist Vol. 40, No. 4 Tablf. 1. Vegetation types and locations of perniaiient sites for Biighaiii Young University Navajo-Kaiparowits study. Site No. Vegetation type Oryzopsis-Stipa-Ephedra Vanclevea Location At the base of Cedar Mt. on loo.se, deep sand, 3.2 mi W of GCC through the highway fence. T 4.3 S R 2E S\\' Part of the NW quarter of Sec. 21. Junipenis-Bouteloua On Cedar Mt. 4.1 mi on from Site 1. .\pprox. 20 vds N of fence on left of road. Exclosure 100 yd N of fence and W 30 yd R 44 S T 2E NE corner of Sec. 3. Hilaria-Ephedra Cedar Mt. .\pprox. 1.3 mi on from Site 2. Road is headed W and ca 50 vd before Pole Line on left. R 44 S T 2E Center of Sec. 10, ca 100 vd from state line. Kane Co., UT. Coleogyne Cedar Mt. 0.8 mi on from Site 3 after turning left under Pole Line. R 6E T 42.N SW part of the NE quarter of Sec. 5. .•\triplex corrugata- A triplex confertifolia .\pprox. 7.0 mi East GCC near U.S. W. B. and Dames and Moore weather station on Dakota formation on right, ca 75 yd from W. B. station. Pinyon-Juniper- Artemisia Smokij Mt. 14.2 mi from Last Chance Jet. Turn right .4 mi Site on right of road. On coal mine road turn right at top of Smoky (survey marker) 0.1 mi then left .4 mi T 41S R 4E NE part of the SE quarter of Sec. 8. 10. Gravia-Coleogvne-Hilaria Gravia-Hilaria Atriplcx-Kochia- Artemisia spinescens Orvzopsis Smoky Mt. 12.3 mi from Last Chance Jet. on top. E of road near dead juniper. T 41S R 8E NE part of the SE quarter of Sec. 21. Smokif Mt. 9.1 mi from Last Chance Jet. (on left of road) or 5.2 mi from coal mine road jet. on right side of road. T 41S R 9E SW part of the SE quarter of Sec. .34. Base of Sntokii Mt. 2 mi on from Ahlstrom Point Jet. on left. Turn off road in bottom of small wash to dead end at ca .2 mi Site 20 yd N. T 42S R 5E NW part of the NW quarter of Sec. .30. Alil.strotn Point Road 4.7 mi from Ahlstrom Point Jet. T 43S R 5E SE part of the NW quarter of Sec. 8. December 1980 Tal)Ie 1 tontiiuied. Atwood et al.: Kaiparowits V'ertebrates 307 Vegetation type Coleogvne Coleotrvne Grayia-Ephedra-Orvzopsis Boiiteloua-Hilaria Grayia-Coleogyne-Bouteloiia Popiilus-Tainaiix-L\ ciimi Tainari.\-Chr\'sothainn\is Spoiol) (Joleosivne Location Ahl.strom Point Road 5.0 mi or .3 mi from Site 10. T 4;3,S R 5E .\E part of the SE (|iiartL-r of Sec. 8. \il)))lc Bench In first well- developed Goleogyne ca .3.5 mi from jet. at top after leaving Tibbet Spring. On left of road, there i,s a .small ridge on right. T 42 S R .3E NE part of the S\\' qnarter of Sec. 9. \ipple Bench .\pprox. .6 mi on fiom Site 12. Tnrn np wash to the right ca 50 ft into Sec. 17 from marker, which is .1 mi off road up the woods. T 42 S R 3E \E part of the NE qnarter of Sec. 17. \ip})le Bench .Appro.x. 1.1 mi S of SE corner 17 on W of road. 2.1 mi S Site 13 T 42 S R 3E SE part of the SW ([iiarter of Sec. 21. Lmt ('liance Creek 31.8 mi from C.CXJ tnrn \ip creek bottom ca 1 mi T 415 R 5E SE part of the SE quarter of Sec. 4. .Salt Wash at Middle Branch Creek Across the creek and ca 40 vd downstream from washed out stock watering pond. T 41S R 4E SE part of the SE quarter of Sec. .3.5. Staleline 7.6 mi from (iC^C on Utah side of border turn left toward Lone Rock .Marina from U.S. 89. .\pprox. .2 mi on right of road. Statcline Same as above except on left of road 75 \cl and on a slight rise. Coleogyne Winihnill 9.3 mi S of Faue on 89 on left of highwav on pullout bv reflector post. T .39.\ R 8E SE part of the XW (luarter of Sec. 10. Bouteloua-Hilaria- Muhlenber<;ia Pinvon-Juniper Coleog)aie M mi post .5.35 ca 12 mi S of Page. 50 vd \ of post and 20 \d E of road on pullout. T .39N'R 8E SW part of the .\E Cjuarter of Sec. 20. Echo Cliffs 19.3 mi S of Page turn left and on ridge ca '4 mi. Xavajo Plant Site (undecided) 308 Great Basin Naturalist Vol. 40, No. 4 Table 1 continued. Site No. Vegetation type Location 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 3.3a. 33h. 33c. Coleogyne-Ephedra-Grayia Hilaria-Ephedra-Grayia Atriplex confertifolia Atriplex corrugata Pinyon-Juniper Artemisia tiidentata Pinyon-Jiiniper Grass-Ephedra Coleogvne Orvzopsis Moist hanging gardens with thin-leaved genera of trees and shrubs various genera of herbaceous plants. Cercis, Cladium, Rhamnus Ostrva, Cirsiium. Rubus Quercus. Gcltis. Populus, Baccharis, grass/forlj 1.2 mi N Tibbet Spring to jet. and thence 1.6 mi E on Cathy's Flat Road 50 yd N of road. T 41S R 3E NE part of the SE quarter of Sec. 33. 2.1 mi E Site 2.3. .\ppro.\. .2 mi beyond end of road. T 42S R 3E SE part of the NE quarter of Sec. 3. To be selected near Cath\"s Flat. 1(X) yd W of small twin flat-tops at Last Chance Summit on S of road ca 13 mi E of Warm Creek Jet. T 42S R .5E east central part of the SE quarter of Sec. 25. The Pine ,\pprox. 3 mi from cow camp at head of Wesses Canyon. T 40 S R 2E part of the SE quarter of Sec. 2. 1-1 mi W Drip Jet. 40 yd SW from Dead Juniper on S of road. T 40S R 3E NE part of the SW quarter of Sec. 7. Farthest distance out on Drip Point. T 40S R 3E SE part of the NE quarter of Sec. 21. T 41S R 1 W NE part of the SW quarter of Sec. 27. Brigham Plains Bench, ca 5 mi N of U.S. Highway 89, E of Paria River. Grand Bench ca 50 mi E of Glen Canyon City. T 42S R 6E. Grand Bench ca 54 mi E of Glen Canvon Citv. T 42S R 6E. In Driftwood Canvon on the .N side of Lake Powell ca'l mi NW of Rainbow Bridge Canyon. T 43S R 8E. Ribbon Canyon, between San Juan Drainage and Hole-in-the-Rock on the E side of Canyon. T 41S R lOE. Reflection Canyon (Cottonwood Gulch), between San Juan drainage and Hole-in-the-Rock on the W side of canvon. T 42S R 9E. December 1980 Atwood et al.: Kaiparowits Vertebrates 309 Table 1 continiifd. Site No. Vegetation type Location .33d. Quercus, Cercis, Cirsiuin 1 pii N of confluence of Forh/grass Colorado/San Jnan Rivers on E side of canyon. T 42S R 9E ,5.5 mi E of Glen Canyon Citv in Last Chance Creek. T 41S R 6E. .34. Taniarix-dcsert siiiuh K 42S H9E.55ini Euftilen Canyon C^itv in Last C^hance Creek. T 4IS R 6E. Willow Tank Spring and confluence Calf Creek/Escalante River; Hayvvard et al. (1958) Navajo Mountain; Russell and Thomp- son (1964) Bryce Canyon; BYU (1972) Grosvenor Arch. Bufonidae (Toads) Bufo cognatus Sav. Great Plains Toad. BYU (1971) Site 15. Biifo punctattis Baird & Girard. Red- spotted Toad. V. M. Tanner (1940a) Willow Tank Spring; Woodbury et al. (1959) Bridge Canyon, Rock Creek, Hidden Pa.ssage, Aztec Canvon, Rainbow Bridge, and Warm Creek; BYU (1971-1973) Warm Creek, Escalante drainage, Wahweap Creek, Driftwood Can- yon, Reflection Canyon, and Three Garden. Bufo woodhoiisei Girard. Woodhouses Toad. V. M. Tanner (1940a) Tropic, Esca- lante, and Escalante River; Hay ward et al. (1958) Navajo Mountain; Woodbury et al. (1959) Rock Creek and Kane Creek; Russell and Thompson (1964) Bryce Canvon; BYU (1972-1973) Cottonwood 'Wash Spring, Re- flection Canyon, and Tibbet Spring. Hylidae (Treefrogs) Hyla arenicolor Cope. Canyon Tree Frog. V. M. Tanner (1940a) confluence of the Esca- lante /Colorado River; Hayward et al. (1958) Paria Valley; Woodbury et al. (1959) Bridge Canyon and Rock Creek; BYU (1972-1973) Driftwood Canyon and Three Garden. Ranidae (True Frogs) Rana pipens brachycephala Cope. West- em Leopard Frog. V. M. Tanner (1940a) con- fluence of the Escalante/Colorado Rivers; Woodbury et al. (1959) Bridge Canyon, Hole- in-the-Rock, Hidden Pas.sage, Rock Creek, West Canyon, Padre Creek, and Warm Creek; Russell and Thompson (1964) Br\ce Canyon; BYU (1972) Escalante River, Reflec- tion Canyon, and Ribbon Canvon. Reptiles Emydidae (Water and Bo.x Turtles) Chrysemys picta belli Gray. Western Painted Turtle. Woodbury et al. (1959) Rock Creek, Labvrinth Canvon, and three miles above Face Canyon; Miller (1966) Hole-in- the-Rock. Iguanidae (Iguanid Lizards) Crotaphytus collaris subssp. Western C^ol- lard Lizard. Records of C. c. haileyi Stejnegar are: V. M. Tanner (1940a) Paria River and Henrieville; Woodbury et al. (1959) Rainbow Bridge and Rock Creek; W. W. Tanner (1958a) collected specimens between Lee's Ferry and Tuba City and indicated, "inter- gradations of .subspecies C. c. haileyi and C. c. auriceps occur in the region south of the San Juan River and west of its confluence with Colorado River." BYU (1971-1974) Sites 4 and 11, .south ba.se of Navajo Mountain. Glen Canyon, and Nipple Bench. C. r. bicin- tores Smith & W. W. Tanner was named from specimens collected at Crossing-of-lhe- Fathers. An additional collection of this ani- mal was made by BYU (1971) east of Smoky Mountain. Crotaphytus wislizenii punctatus Baird & Girard. Long-nosed Leopard Lizard. \'. .M. Tanner (1940a) Willow Tank Spring; Hay- ward et al. (1958) Navajo Mountain; Wood- 310 Great Basin Naturalist Vol. 40, No. 4 bury et al. (1959) Last Chance and Lee's Fer- ry; W. W. Tanner and Banta (1963) Hole-in- the-Rock, Lone Rock, Willow Tank Spring, Catstairs Canyon, Crossing-of-the-Fathers, Navajo Mountain Trading Post, and Lower Wahweap Creek; BYU (1971-1973) Sites 1, 14, and 27, and Cottonwood Wash. Tanner and Banta (1977). W. W. Tanner (1980, pers. comm.). Sauromalus obesus multiforaminatus Tan- ner & Avery. Upper Colorado River Chuck- walla. V. M. Tanner (1940a) Warm Creek; Hayward et al. (1958) Paria Valley; Wood- bury et al. (1959) Rainbow Bridge, Rock Creek, and Last Chance drainage; W. W. Tanner and Avery (1964) Crossing-of-the-Fa- thers, Hole-in-the-Rock, Warm Creek, and one mile upstream from Glen Canyon Dam; BYU (1971-1972) Navajo Creek, Warm Creek Bay, Glen Canyon Dam, Grand Bench, and Navajo Creek. Holbrookia maculata approximans Baird. Lesser Earless Lizard. BYU (1972) Site 20, Tietso Spring, and 13 miles south of Page along U.S. Highway 89. Sceloporus magister cephaloflacus Tanner. Orange-headed Desert Spiny-Lizard. V. M. Tanner (1940a) Willow Tank Spring and Wahweap Creek; W. W. Tanner (1954a) Kai- parowits Plateau, Lone Rock, Catstairs Can- yon, Escalante River, and 15 miles northwest of Hole-in-the-Rock; Hayward et al. (1958) Navajo Mountain; Woodbury et al. (1959) Bridge Canvon, Rock Creek, and Kane Creek; BYU (1971-1974) Sites 1, 2, 8, 12, 14, 16, 17, and 23, 5 miles south of Page, Tibbet Canyon, Tibbet Spring, Wahweap Bay area, Cottonwood Wash/U.S. 89, and Three Gar- den. Sceloporus undulatus elongatus Stejnegar. Northern Plateau Lizard. V. M. Tanner (1940a) Cannonville, Escalante, and Calf Creek/Escalante River; W. W. Tanner (1954d) Paria River drainage north of U.S. Highway 89; Hayward et al. (1958) Navajo Mountain; Russell and Thompson (1964) Bryce Canyon; BYU (1973) Tibbet Canyon. Sceloporus graciosus graciosus Baird & Gi- rard. Great Basin Sagebrush Lizard. V. M. Tanner (1930), Weight (1932), and Presnall (1935) Bryce Canyon; V. M. Tanner (1940a) Calf Creek/Escalante River, Tropic, and Es- calante/Colorado Rivers; Havward et al. (1958) Navajo Mountain; BYU (1971-1974) south base of Navajo Mountain and at Sites 1, 10, 13, 14, 21,23, and 27. Ufa stansburiana uniformis Pack & Tan- ner. Upper Colorado Basin Side-blotched Lizard. V. M. Tanner (1940) Escalante, Wil- low Tank Spring, and Calf Creek/Escalante River; W. W. Tanner (1954d) Paria River drainage north of U.S. Highway 89; Hayward et al. (1958) Navajo Mountain; Woodbury et al. (1959) Rainbow Bridge Trail and lower Bridge Canyon; Russell and Thompson (1964) Brvce Canvon; BYU (1971-1974) Sites 1, 2, 4, 6, 7, 8, 9, 10,1 3, 14, 17, 18, 19, 20, 21, 22, 23, 27, 28, and 30, Glen Canyon City, Four Mile Bench, Three Garden, and Tibbet Can- yon. Urosaurus ornata wrighti Schmidt. Colo- rado Tree Lizard. W. W. Tanner (1954d) Paria River drainage north of U.S. Highway 89; Hayward et al. (1958) Navajo Mountain; Woodbury et al. (1959) Bridge Canyon, Rock Creek, and Last Chance; BYU (1971) three miles west of Site 19. Phrynosoma douglasii hernandesi Girard. Short-horned Lizard. V. M. Tanner (1930), Weight (1932), and Presnall (1935) Bryce Canyon; BYU (1972) Sites 14 and 23, and Warm Creek Bay. W. W. Tanner (1975). Phrynosotna platyrhinos calidairum Cope. Sonoran Desert Horned Lizard. W. W. Tan- ner (1954d) Paria River drainage north of U.S. 89; Woodbury et al. (1959) Rock Creek and Lee's Ferry; BYU (1971) Utah-Arizona state line north of Page along U.S. Highway 89. Xantusidae (Night Lizards) Xantusia vigilis utahensis Tanner. Utah Night Lizard. W. W. Tanner (1957) records topotvpes taken from Trachyte Creek, Gar- field Co., Utah; W. W. Tanner (1958b) in- dicates this species possibly is found within the area defined as the Kaiparowits Basin. Teidae (Whiptail Lizards) Cnemidophorus tigris Baird & Girard. Western Whiptail. Two subspecies occur in the Kaiparowits Basin, viz. C. t. tigris (Baird & Girard) reported by V. M. Tanner (1930), Weight (1932), and' Presnall (1935) Bryce December 1980 Atwood et al.: Kaiparowits Vertebrates 311 Canyon and V. M. Tanner (1940a) Calf Creek /Escalante and Paria River drainage; and C. t. septentrionalis Burger reported by Woodbury et al. (1959) from Rainbow Bridge Trail, Beaver Creek, and lower Bridge Can- yon, BYU (1971-1973) Sites 1, 2, 6,^^13, 16, 17, 19, 20, 22, 23, 27, and 30, and the south base of Navajo Mountain. Cnemidophorus velox Springer. Plateau Whiptail. Woodbury et al. (1959) Rock Creek; Schmidt (1953) lists C. sacki innotatus from Kanab, Kane Countv. Cnemidophorus sexlineatus perplexus Baird & Girard, Six Lined Racerunner. V. M. Tanner (1940a) Canonville and Escalante. Scincidae (Skinks) Eumeces skiltonianus utahensis Tanner. Great Basin Skink. Russell and Thompson (1964) Bryce Canyon beneath logs and stones. Colubridae Thamnophis cyrtopsis cyrtopsis Kennicott. Western Black-necked Garter Snake. Schmidt (1953) lists this species for Utah, Ari- zona, southern Colorado, New Mexico, and southward. Thamnophis elegans vagrans Baird & Gi- rard. Wandering Garter Snake. V. M. Tanner (1930), Weight (1932), and Presnall (1935) Bryce Canyon; V. M. Tanner (1940a) Tropic and confluence Calf Creek /Escalante River; Woodbury et al. (1959) Rock Creek. Masticophis taeniatus taeniatus Hallow- ell. Desert Striped Whipsnake. W. W'. Tan- ner (1954d) Paria drainage north of U.S. Highway 89; Woodbury et al. (1959) Beaver Creek, Kaiparowits Plateau, and Hole-in-the- Rock; Russell and Thompson (1964) Bryce Canyon. Salvadora hexalepis mojavensis Bogert. Mojave Patch-nosed Snake. W. W. Tanner (1953) near the old town site of Adairville, Kane County, Utah; W. W. Tanner (1954c) Wahweap Creek, one mile southeast of Lone Rock; Hayward et al. (1958) Paria Valley; Woodbury et al. (1959) Rock Creek; BYU (1971) Paria River three miles south of U.S. Highway 89 and at Glen Canyon City. Arizona elegans philipi Klauber. Painted Desert Glossy Snake. W. W. Tanner (1964) 14 miles south of Page, 9 miles west of Page, and 2 and 5 miles west of the Paria River all along U.S. Highway 89; BYU (1972) Sites 1 and 3, Arizona-Utah border between Page and Glen Canyon City, and 5 miles northwest of Page. Pituophis meUinoleucus deserticola Stejne- gar. Great Basin Gopher Snake. Presnall (1935) Bryce Canyon; V. M. Tanner (1940a) Tropic, Escalante, Wahweap Creek, and W'illow Tank Spring; W. W. Tanner (1954d) Paria River drainage north of U.S. Highway 89; Hayward et al. (1958) Navajo Moimtain; BYU (1971-1973) Sites 1, 2, 14, 17, and 20. Lampropeltis getulus californiac Blain- ville. California King Snake. \'. M. Tanner (1940a) 30 miles south of Escalante; W. W. Tanner (1958b) undoubtedly extending well into the upper Colorado Basin; BYU (1971-1973) one observed dead on U.S. High- way 89 by Glen Canyon City and another collected west of Cockscomb Ridge near U.S. Highway 89. Rhinocheilus lecontei lecontei Baird & Gi- rard. W'estern Long-nosed Snake. W. W. Tanner (1964) Wahweap road at junction with U.S. 89 and Buck Tank Draw. Sonora semiannulata isozona Cope. West- ern Ground Snake. BYU (1972) seven miles southeast of Glen Canyon City on U.S. High- way 89. The specimens collected by BYU ex- tend the range of this subspecies into Kane County and east almost to the Colorado River. Hypsiglena torquata deserticola Tanner. Desert Night Snake. W. W. Tanner (1954b) northeastern Kane County, Utah, in the area south and east of the Vermillion Cliffs; Rus- sell and Thompson (1964) Bryce Canyon. Hypsiglena torquata loreala Tanner. Plateau Spotted Night Snake. Woodbury et al. (1959) Labyrinth Canyon; BYU (1973) Re- flection Canyon. Tantilla planiceps utahensis Blanchard. Utah Black-headed Snake. W. W. Tanner (1954c) Paria River drainage 38 miles east of Kanab in Catstairs Canvon. Crotalidae (Rattlesnakes) Crotalus viridis lutosus Klauber. Great Basin Rattlesnake. V. .\1. Tanner (1930), Weight (1932), and Presnall (1935) Bryce 312 Great Basin Naturalist Vol. 40, No. 4 Canyon; W. W. Tanner (1958a) indicates that this subspecies ranges east at least to the Paria River. Pritchett (1962) extended the range to the plateau east of the Paria. Crotalus viridis nuntius Klauber. Hopi Rattlesnake. W. W. Tanner (1958a) southeast portion of Navajo Mountain near the Utah- Arizona border and at the confluence of Es- calante River, Willow Tank Spring, and Wahweap Creek; W. W. Tanner (1958a) north Escalante/ Colorado River junction; BYU (1972-1974) Sites 2, 3, 10, 14, 22, and 28, Tibbet Canyon, two miles southeast of Glen Canyon, Tibbet Spring, and Paria, Utah. Crotalus viridis concolor Woodbury. Midget-faded Rattlesnake. V. M. Tanner (1940a) Tropic, Escalante River, Willow Tank Spring, and Wahweap Creek; W. W. Tanner (1958a) north of Escalante/Colorado River junction; BYU (1972-1974) Sites 2, 3, 10, 14, 22, and 28, Tibbet Canyon, two miles southeast of Glen Canyon City, Tibbet Spring, and Paria, Utah. Birds Present avian classification systems have divided Fringillidae into two families and have separated them in their positions in the phylogenetic order and have removed other species such as the Bushtits {Psaltripanis) from tlie families where they have been well established. Additionally, new positions in the phylogenetic order of a number of Passerine families have been proposed. Hay- ward et al. (1976) have discussed these di- verse taxonomic changes in considerable de- tail and have deviated from the Fifth Edition of the AOU Check-list (1957) to conform with some of the proposed changes. In view of the present diversity of opinion regarding avian classification and because the AOU Check-list committee has not yet made a de- cision on the proposed changes, we have fol- lowed the Fifth Edition of the AOU Check- list and its supplements. Our ornithological data were collected from July 1971 through Febniary 1974; the number of individual birds seen on a monthly or seasonal basis and the number seen in vari- ous vegetational associations on a seasonal basis are biased, since only two years data were collected for the months of March through June. Also no data were collected in December 1972. The data are biased further by the fact that not every observer deter- mined numbers, nor did they always report the vegetational associations in which they saw the species. Furthermore, the vegeta- tional associations were not sampled equally for bird species composition and numbers; some were sampled more intensively than others. Therefore, our data pertaining to avian seasonal population trends and their preferences for certain vegetational associ- ation are only suggestive. Additionally, since birds usually show decided preferences for the architectural structure of the vegetation rather than the plant species composition of specific associations, many of the plant asso- ciations have been lumped into Pi7ius-Juni- perus or Juniper us associations, often referred to as woodland, riparian {Populus fremontii, Salix, Tamarix) associations, and desert shrubs. Additional data are available on habi- tat relationships of the vertebrate animals discussed here by comparing the sites (Fig. 1, Table 1) where the animal was seen or col- lected with the vegetational associations found at that site. All river miles listed for the Colorado River are reckoned from Lee's Ferry. In the following species accounts, numbers followed by localities and dates refer to the number of specimens taken. Podicipedidae (Grebes) Podiceps nigricollis C. L. Brehm. Eared Grebe. BYU (1971) 50 seen at Warm Creek Bay, 8 Nov.; BYU (1972) 12 seen at river mile 56, 7 on Lake Powell near mouth of San Juan River, and 17 at Three Garden on Lake Pow- ell, 2 Nov. Behle et al. (1958) regarded the species as a regular spring migrant through the Kanab area. Late fall resident on Lake Powell. Aechmophorus occidentalis (Lawrence). Western Grebe. BYU (1971) Lake Powell 6 seen on Warm Creek Bay, 8 Nov. and one seen on Wahweap Bay, 1 Dec. A total of 80 Western Grebes were counted at the follow- ing localities of Lake Powell 2 Nov. 1972: river miles 12, 14, 17, 18 and 20, Dangling Rope Canyon, Driftwood Canyon, and December 1980 Atwood et al.: Kaiparowits Vertebrates 313 Navajo Creek. A 29 June 1973 observation on Lake Powell suggests either a straggler or possible nesting. Late fall transient south- eastern Utah (Behle 1960). Podilymbus podiceps (Linnaeus). Pied- billed Grebe. BYU (1971) on Lake Powell, one seen at Warm Creek Bay, 8 Nov. and an- other at Wahweap Bay, 1 Dec; BYU (1972) 36 were recorded at the following localities on Lake Powell: Gregory Butte near Rock Creek Bay, Dangling Rope Bay, and river miles 12 and 15, 2 Nov. Late fall resident on Lake Powell. Pelecanidae (Pelicans) Pelecanus erythrorhynchos Gmelin, White Pelican. Presnall (1937) Bryce Canyon; Woodbury and Russell (1945) three miles be- low Rock Creek on Colorado River, 28 July 1937. Rare transient. Ardeidae (Herons and Bitterns) Ardea herodias Linnaeus. Great Blue He- ron. Behle (1948) Aztec Creek, Rock Creek, Last Chance, and river miles 3, 6, and 21; Woodbury and Russell (1945) one at Lee's Ferry, Aug. 1909 as recorded by the Nelsen and Birdseye trip; Behle and Higgins (1959) from previously unpublished observations of Woodbury and Russell on the Rainbow Bridge-Monument Valley expedition, 23 herons seen from 1 to 11 Aug. 1938; they were seen from 2V2 miles below Lee's Ferry to river mile 69, 3100-3200 ft; Behle (1960) common from Hite to Lee's Ferry, May through Oct.; BYU one at Warm Creek/Lake Powell, 8 Nov. 1971 and 3 May 1972; BYU (1973) one at Three Garden alcove, 21 Mar. The presence of a nesting colony at river mile 117 (Woodbury et al. 1959) indicates that the species also nests in the area. Not un- common resident between 21 Mar. and 8 Nov. Egretta thiila (Molina). Snowy Egret. Woodbury and Russell recorded in their un- published field notes observing this species at Forbidding Canyon along the Colorado River 18 July 1937 and seeing tracks on shore of Colorado River between river miles 63-69 (Behle and Higgins 1959); Behle (1960) Hig- gins saw 17 at Wahweap Creek and river mile 17 and one at the dam site, 7 .\ug. 1958; BYU (1972) one seen in Populus at Navajo Creek, 27 Apr.; BYU (1973) one seen by Rob- ert Whitmore at junction of Paria Riv- er/Colorado River, 6 June (riparian vegeta- tion), and 2 more were seen bv Whitmore feeding from a sandbar in the Colorado River one mile north of Lee's Ferrv, 15 June. Spring and summer resident, possiblv breeds. Nycticorax nycticorax (Linnaeus). Black- crowned Night Heron. \\'oodbur\- and Rus- sell (1945) specimen: river mile 64, 4 Aug. 1938 and six were seen in 65 miles, i.e., be- tween river mile 63 and 2'/2 miles below Lee's Ferry, between 4 and 11 .\ug. 1938; Behle (1948) two were seen 16 .\pr. 1947 at Crossing-of-the-Fathers near Kane Creek and another pair seen at Wahweap Creek. None were seen during the present investigation. Summer and spring records suggest breeding. Threskiornithidae (Ibises and Spoonbills) Plegadis chihi (Vieillot). White-faced Ibis. Behle and Higgins (1959) reported that ood- bury saw two at river mile 13, 13 Sept. 1957. Uncommon transient along the Colorado Riv- er. Anatidae (Swans, Geese, and Ducks) Branta canadensis (Linnaeus). Canada Goose. Grater (1947) Bryce Canyon; Behle and Higgins (1959) noted that Woodbury saw two at river mile 50, 12 Sept. 1957; Behle and Higgins (1959) observed a pair of adults with six young that were unable to fly at riv- er mile 106, 16 July 1958; they considered the species a common sinnmer resident and transient in Glen Canyon. Sununer resident, possiblv nesting near Lake Powell. Anas platyrhynchos Linnaeus. .Mallard. Behle (1948) six were seen near Hole-in-the- Rock; BYU (1971) three were seen on a small islet at Warm Creek/Lake Powell, 8 Nov. Uncommon transient along the Colorado Riv- er and Lake Powell. Anas strepera Linnaeus. Gadwall. Behle (1948) saw a pair at Hole-in-the-Rock, five at the mouth of San Juan River, and two at Wahweap Creek; BYU (1972) five were seen on a pond at Grosvenor's Arch, 2 Mar. and two at Lake Powell /Cottonwood Wash. 22 314 Great Basin Naturalist Vol. 40, No. 4 July (Ephedra-gr ass); BYU (1973) three on a pond near Coyote Creek, 5 June. Uncommon spring and summer resident. Anas acuta (Linnaeus). Pintail. Behle (1948) six near Hole-in-the-Rock; BYU (1973) two on a pond at Wiregrass Spring, 5 Nov. Spring and fall transient. Anas crecca (Linnaeus). Green-winged Teal. Woodbury and Russell (1945) one, Col- orado River between Rock Creek and Lee's Ferry, 17 Aug. 1937; Behle and Higgins (1959) Woodbury saw seven at river mile 82, 11 Sept. 1957; BYU (1972) three at Wahweap Creek Bay on a rain pond, 30 Oct. Transient Lake Powell and the Colorado River. Anas discors Linnaeus. Blue-winged Teal. Behle (1948) near Klondike Bar; Behle and Higgins (1959) Higgins saw six near mouth of Rock Creek at river mile 58, 28 July 1958. Uncommon spring and summer transient Col- orado River and Lake Powell. Anas cyanoptera Vieillot. Cinnamon Teal. Woodbury and Russell (1945) one, Paria /Colorado Rivers, 11 Aug. 1938; Behle (1948) mouth of Wahweap Creek; Behle et al. (1958) found one dead on highway south of Escalante, 8 May 1954; Behle and Higgins (1959) reported that in 1958 Woodbury saw one between river mile 63 and 50, 5 Aug. and one at V2 mile below Lee's Ferry, 11 Aug. Spring and summer transient. Anas americana Gmelin. American Wid- geon. BYU (1972) two at Rainbow Landing on Lake Powell, 2 Nov. Uncommon transient on Lake Powell. Aythyinae (Diving Ducks) Aythya valisineria (Wilson). Canvasback. Behle (1948) near mouth of Last Chance Creek. Uncommon transient. Bucephala clangula (Linnaeus). Common Goldeneye. Behle (1948) saw 12 at Crossing- of-the-Fathers, 12 at river mile 36, and 16 at river mile 31; BYU (1972) one at Coyote Creek Pond, 8 Dec. Spring and winter tran- sient. Bucephala albeola (Linnaeus). Bufflehead. BYU (1972) five on Lake Powell at Warm Creek, 7 Feb.; three on gravel pit pond at Wahweap Bay, 27 Mar.; and a female on Colorado River at Last Chance Bay, 5 July. Uncommon spring and summer transient. Cathartidae (American Vultures) Cathartes aura teter Friedmann. Turkey Vulture. Presnall (1934) Bryce Canyon; Behle and Higgins (1959) reported that Woodbury saw two along the Colorado River between miles 13 and zero, 10 Aug. 1938; Woodbury and Russell (1945) one, Navajo Mountain Trading Post on 26 July 1936 and another on 8 Aug. 1936 near Lee's Ferry, and observed on Kaiparowits Plateau; BYU (1971) two seen 12 miles south Page, Ariz., 24 July; BYU (1972) Site 2 (flying), 23 July; Site 12, 16 Apr. 1972 (Coleogyne); Site 20; Driftwood Can- yon, 24 June; three seen at Dry Rock Creek, 3 May; and one the confluence of the San Juan River/Lake Powell, 17 July. Woodbury and Russell (1945) reported the Turkey Vul- ture as a common summer resident almost everywhere in the Navajo country area in the 1930s. They observed it nearly every day from 1 June through 15 Sept. over a period of years. Yet, present observations covering a three-year span yielded fewer than a dozen sightings, suggesting a drastic decline in num- bers. This is particularly significant when one considers, as noted by Woodbury and Russell (1945), "that this conspicuous bird is seldom missed in observations in both time and space," a factor that tends to overemphasize its occurrence by comparison with less con- spicuous birds (Woodbury and Russell 1945). Uncommon summer resident from 28 Mar. to 1 Oct. (in the general area). Accipitridae (Eagles, Hawks, and Harriers) Accipiter gentilis atricapillus (Wilson). Goshawk. Woodbury and Russell (1945) Nav- ajo Mountain, i.e.. War God Spring, 23 July 1936; Russell and Thompson (1964) Bryce Canyon. Uncommon summer resident. Accipiter striatus velox (Wilson). Sharp- shinned Hawk. Presnall (1934) Bryce Point; Woodbury and Russell (1945) Navajo Moun- tain, 13 July 1933, and Kaiparowits Plateau, 30 July 1937, and young birds out of nest being fed by parents on Kaiparowits Plateau, 5-6 Aug. 1937; Benson (1935) base of Navajo Mountain, mid- June 1933; BYU (1971) Site 8, 1 Aug. and one Navajo Mountain, 13 Oct. Fairly common summer resident and spring and fall transient. December 1980 Atwood et al.: Kaiparowits Vertebrates 315 Accipiter cooperii (Bonaparte). Cooper's Hawk. Presnall (1934) and Grater (1947) Bryce Canyon; Woodbury and Russell (1945) specimens: Navajo Mountain, 9 Aug. 1935 and 25 July 1936 and two seen nesting on Kaiparowits Plateau, 5 Aug. 1937; Behle and Higgins (1959) Rock Creek river mile 58, 28 July 1958; BYU (1971) two near Site 1 in Jiinipenis, 14 and 18 July; BYU (1973) one at Wahweap Creek, 30 Apr. {Tamarix in Salt- wash). Summer resident, spring and fall tran- sient. Buteo jamaicensis calurus (Cassin). Red- tailed Hawk. Presnall (1934) Bryce Canyon; Woodbury and Russell (1945) two at Navajo Mountain, 7 July 1936 at 3250 ft and seen at Beaver Creek, War God Spring, Soldier Seep (Navajo Mountain), and on Kaiparowits Plateau; Behle and Higgins (1959) reported that Woodbury found them common along Colorado River, 4-23 July 1936, and one nest with young hawks was also noted; Behle and Higgins (1959) abundant in Glen Canyon, 1 July to 9 Aug. 1958. BYU (1971-1973") Sites 1, 2, 6, 12, 13, 15, 18, 20, 22, 23, 27, and 30, and Cottonwood Wash, Tibbet Canyon, Ce- dar Mountain, Grosvenor Arch, Wahweap Creek, and specimen: Cockscomb /Highway 89, 14 Jan. 1972. Recorded every month; April-July (21 seen) and November-Januarv (17 seen). The months January-June are un- fairly represented since no records were ob- tained for these months in 1971. During the winter months most observations were in Juniperus and associated species (17), and the remainder were in grass (2), grass-shrub (2), perched on a ledge (2), and in Tamarix (1). During the spring and summer nesting season they were seen flying over desert shrubs (9) consisting of Atriplex, Vanclevea, Chryso- thamnus, Artemisia, and shrub-grass. Four of the nine were in Coleogijne; three others were in Juniperus and/or woodland, and one each in grass and washbottom situations. Per- manent resident. Buteo swainsoni Bonaparte. Swainson's Hawk. Presnall (1934) reported this species to be common at Bryce Canyon during the summer. Buteo lagopus (Pontoppidan). Rough- legged Hawk. In 1935 Long (1937) observed one in Bryce Canyon. Sparse winter resident. Buteo regalis (Gray). Ferruginous Hawk. Long (1937) and Russell and Thompson (1964) Bryce; BYU (1973) one perched on power lines south of Glen Canyon near Warm Creek, 27 Apr. Uncommon transient. Aquila chrysaetos canadimsis (Linnaeus). Golden Eagle. Presnall (1934) Rainbow Point; Grater (1947) Bryce Canyon; Wood- bury and Russell (1945) Navajo Mountain and Glen Canyon; Behle and Higgins (1959) sighted adults and immatures near mouth of Aztec Creek, 26 July 1958; BYU (1971-1974) Sites 1, 6, 12, and 23, Glen Canyon City. Church Wells, Cockscomb, Grosvenor Arch, Last Chance Creek, Tibbet Canyon, Wah- weap Creek, and Warm Creek. Recorded 42 times at 26 locations during every month of the year except October. Fewest birds were seen October to November and .March to April, and most May- August followed bv De- cember-Febniary. Only one eagle was seen during our study, in January. The vegeta- tional type over which they were flving or in which they were perched during Janu- ary-June consisted of desert shrubs (6), cliffs or rock ledges (4), grasslands (2), woodland (1), and washbottom (1). In August, Septem- ber, and December they were recorded in mixed shrubs and cliffs (4). Common per- manent resident. Haliaeetus leucocephalus (Linnaeus). Bald Eagle. Behle and Higgins (1959) report that Woodbury observed one flying on the east side of the Kaiparowits Plateau below the cliffs north of Glen Canyon, 4 Aug. 1938; BYU (1972) one seen near Rock Creek Bay and another at confluence of San Juan Riv- er/Lake Powell, 2 Nov.; BYU (1975) Last Chance Wash, Dec. Sparse fall transient; the 4 Aug. bird seen by Woodbury may haNC been a postnesting eagle from the small nest- ing populations in central Arizona. Circus cyaneus hudsonius (Linnaeus). Marsh Hawk. Woodbury and Russell il945^ one, Kaiparowits Plateau, 9 Aug. 1937; BYU (1971) two at Site 17, 27 Aug. (gras.slands); two at Site 20, 6 Nov. (grassland): BYU (1972) specimen (male): Wahweap Greek. 17 Jan.; three seen at Wahweap Creek near Glen Canyon City, 7 Feb., 18 Apr., and 3 July (washbottom). Uncommon permanent resi- dent; more common spring and fall transient. 316 Great Basin Naturalist Vol. 40, No. 4 Falconidae (Falcons) Falco mexicanus Schlegel. Prairie Falcon. Presnall (1934) and Russell and Thompson (1964) Bryce Point; Woodbury and Russell (1945) Navajo Mountain; one seen in Glen Canyon in 1938 between river miles 41 and 25, 8 Aug.; and three on cliffs between miles 13 to zero, 10 Aug.; Behle and Higgins (1959) report that in 1938 the species was seen by Woodbury in two places between river miles 78 and zero, 4 July; Behle (1948) pairs (prob- ably nesting) near mouth of Escalante River and junction of Bridge and Aztec Canyons; BYU (1971) five sightings; Site 3, 16 Aug. (grasslands); Site 15, no date; Glen Canyon City/Wahweap Creek, 1 June 1972 (grass- lands); Church Wells, 14 Nov. 1971 (grass- lands); and Nipple Creek (chasing dove), 23 May 1973 (washbottom). Common summer resident, less common in spring and fall; probably a permanent resident. Falco peregrinus anatum (Bonaparte). Peregrine Falcon. Russell and Thompson (1964) Bryce Canyon; Woodbury and Russell (1945) saw peregrines at Navajo Mountain at three different sites, 4-10 July 1936, and at Beaver Creek, Navajo Mountain, 2-16 Aug. 1936; Behle (1960) Glen Canyon near Wah- weap Creek, river mile 17, 6 Aug. 1958; BYU (1971) two observations believed to be per- egrines (both uncertain) at Site 15, 8 Oct. Un- doubtedly the species formerly nested in Glen Canyon, along the Colorado River, and its side canyons as well as on Navajo Moun- tain. With the construction of Lake Powell, the area may now be more suited for pere- grines than formerly (Porter and White 1973). Since early investigators did not exam- ine the area specifically for peregrines, they probably underestimated the size of the population. The entire area now needs to be thoroughly surveyed, especially Lake Powell. Falco columbarius bendirei Swann. Mer- lin. Grater (1947) Bryce Canyon. Falco sparverius sparverius Linnaeus. American Kestrel. Presnall (1934) Bryce Can- yon; Benson (1935) Navajo Mountain; Wood- bury and Russell (1945) two 4 Aug. 1935 and 11 July 1936 (9,500-10,000 ft, Picea-Ahies and Pinus flexilis), and seen on Kaiparowits Plateau and at Lee's Ferry; Behle et al. (1958) one seen 10 miles south of Escalante in stand of Juniperus, 6 May 1954; Hayward et al. (1958) Escalante drainage along stream- sides; BYU (1971-1974) Sites 1, 2, 3, 7, 10, 12, 14, 16, 19, and 26; one mile south of Glen Canyon City, two specimens: 26 Aug. and 13 Sept. 1971; Brigham Plains, Cottonwood Wash, Grosvenor Arch, Smoky Mountain, Driftwood Canyon, Tibbet Canyon, Church Wells, Crosby Canyon (nesting on cliff face, 28 Apr. 1973), Lee's Ferry, and Wahweap Creek. Very common (over 75 birds seen at 31 sites); earliest spring sightings, 20 March 1972 (1 Apr. 1973) and latest fall sightings, 7 Nov. 1971 (3 Aug. 1972 and 9 Aug. 1973). The greatest numbers were seen in April (16), June (17), and July (22). Most observations were in desert shrubs in March-July (29). They were also seen in Tamarix (2), Juniperus (1), saltwash (1), Juniperus in March and April (4), grassland (4), and Tamarix in May- July. The shrubs represented among the desert shrubs were Vanclevea, Atriplex, Graij- ia, Chrifsothamnus, Artemisia tridentata, Co- leogyne, and Epiiedra. Summer resident, a few may winter. Tetraonidae (Grouse) Dendragapus obscurus (Say). Blue Grouse. Presnall (1934), Grater (1947), and Russell and Thompson (1964) Bryce Canyon. Per- manent resident. Centrocercus urophasianus urophasianus (Bonaparte). Sage Grouse. Russell and Thompson (1964) Bryce Canyon. Permanent resident. Phasianidae (Chukars, Pheasants, and Quail) Lophortyx garnbelii gambelii Gambel. Gambel's Quail. Hayward et al. (1958) Calf Creek area; BYU (1971) one at Site 13, 15 Oct. {Coleogyne) and five birds in Cotton- wood Wash; also specimen, 16 Dec. Uncom- mon permanent resident. Phasianus colchicus Linnaeus. Ring- necked Pheasant. Behle et al. (1958) observed and heard in the fields around Escalante, 7 and 8 May 1954. Uncommon permanent resi- dent near areas of cultivation. Alectoris chukar (Gray). Chukar. BYU (1972) Cottonwood Wash. Sparse permanent resident. December 1980 Atwood et al.: Kaiparowits Vertebrates 317 Rallidae (Rails, Gallinules, and Coots) Fulica americana Gmelin. American Coot. Woodbury and Russell (1945) below mouth of Bridge Canyon, 15 July 1936; Behle and Higgins (1959) river mile 47, flushed from S«- hx, 30 July 1958; BYU (1971) on Wahweap Creek, three miles east of Glen Canyon Citv, 6 Nov. {Tamarix around pond); BYU (1973) Site 14, 9 May. Uncommon summer, spring, and fall resident. Charadriidae (Plovers) Charadrius vociferus vociferus Linnaeus. Killdeer. Woodbury and Russell (1945) one, Kaiparowits Plateau, 11 Aug. 1937 (pond, 7000 ft); Behle et al. (1958) confluence Calf Creek/Escalante River; BYU (1971) two, north of Church Wells, 28 Sept.; BYU (1972) two near Glen Canyon City, 14 July and two at Wahweap Creek, 13 June (Tamarix); BYU (1973) one at Coyote Creek, 5 June. Uncom- mon summer resident and spring and fall transient. Scolopacidae (Sandpipers and Willet) Actitis macularia (Linnaeus). Spotted Sandpiper. Woodbury and Russell (1945) one. Rock Creek /Colorado River, 20 July 1937 (3300 ft), and as reported from their impub- lished field notes (Behle and Higgins 1959) they observed 17 individuals from river mile 44 to Lee's Ferry, 7-10 Aug. 1938; Behle and Higgins (1959) common along Colorado throughout Glen Canyon during summer of 1958 and nesting along many of the smaller streams; Behle et al. (1958) confluence Calf Creek/Escalante River; BYU (1973) one at Coyote Creek Pond, 5 June and 2 at junction Paria/ Colorado Rivers, 6 June. Formerly a common summer resident, this species seem- ingly has declined in numbers since the con- struction of the Glen Canyon Dam and the filling of the reservoir. Catoptrophorus semipalmatus inornatus (Brewster). Willet. Presnall (1937) Bryce Canyon; BYU (1972-1973) Four Mile Bench, no date. Sparse transient. Calidris mauri (Cabanis). Western Sand- piper. Woodbury and Russell (1945) near Lee's Ferry, 1 1 Aug. 1938. Sparse transient. Phalaropodidae (Phalaropes) Steganopus tricolor \'ieilIot. Wilson's Phalarope. Woodbury and lUissell (1945) specimen from flock on Kaiparowits Plateau. 2-3 Aug. 1938. Sparse fall transient. Laridae (Gulls and Terns) Larus californicus Lawrence. California Gull. BYU (1971) Warm Ocek Bav. 8 .Nov.; BYU (1973) Lone Rock Ba>, 8 Aug. Sparse fall transient. Lams delawaremis Ord. Ring-bilkd (iiili. Behle (1948) three at Last Chance Creek and again at the mouth of Kane Creek, on the Colorado River. Sparse spring transient. Columbidae (Doves and Pigeons) Columba fasciata fasciata Say. Band- tailed Pigeon. Benson (1935) Navajo Moun- tain; Behle et al. (1958) Bryce Canvon. Sum- mer resident. Zenaida macroura (Linnaeus), .Mourning Dove. Presnall (1934) Br\ce Canvon; Benson (1935) Navajo Mountain; Tanner (1940a) Es- calante River drainage; Woodburv and Rus- sell (1945) Rock Creek, 24-26 July; Kaiparo- wits Plateau, 3 Aug. (1937); and War (iod Springs at Navajo Mountain, 16 .Vug. 1935. common breeder up to 9000 ft elevation on Navajo Mountain; Behle et al. (1958) con- fluence of Calf Oeek/Escalante River; Behle (1960) abundant Coyote Gulch (Whitney) north side of Kaiparowits Plateau. Davis Gulch, and along Escalante River 55 miles southeast of Escalante, Aug. 1957; Behle and Higgins (1959) abundant in Glen (.'anyon. summer 1958; BYU (1971-1973) specinK-n; one mile west Site 2 on 23 July 1971, Sites 1. 6, 10, 13, 14, 19, 28, 30. and 34. Glen Canyon City, Cannonville, Cottonwood Wash. Grosvenor Arch, Tibbet Sprinu, Lee's Ferry. Nipple Creek, Wahweap Creek. Warm Creek, Driftwood Canyon, and Hiree (harden one mile above confluence of .San Juan with Lake Powell. During the .\pril and .May mi- gration period most Mourning Doves were encountered in washbottoins 102'. 5 were seen in grasslands, 14 in desert shrubs, and 4 were distributed more evenly among varioas vegetational types; in contrast, during fall 318 Great Basin Naturalist Vol. 40, No. 4 migration fewest were seen in washbottoms. For example, 24 were seen in grasslands, 22 on desert shrubs, 21 in Popidus, 12 in Tama- rix, 15 in woodland, 6 in hanging gardens, 3 in open fields, 2 in rocks, and 2 in washbot- toms. In AugiLst-November grasslands were most used where 39 individuals were seen, followed by desert shrubs (27) and woodland (2). This species was sighted 155 times with 1335 individuals recorded. The earliest spring record was 15 Apr. (1972) and the latest fall record was 1 Nov. (1973). The peak of spring and fall, passage through the area was in May and in August, respectively. The monthly to- tals for 1972 and 1973 combined are as fol- lows: Apr. (29), May (121), June (72), July (68), Aug. (463), Sept. (109), Oct. (197), and Nov. (2). Abundant summer resident. Strigidae (Typical Owls) Otus asio (Linnaeus). Screech Owl. Russell and Thompson (1964) Bryce Canyon. Per- manent resident. Otus flammeolus (Kaup). Flam mula ted Owl. Woodbury (1939) indicates that during July 1936 H. N. Russell, Jr., collected a speci- men in Salix near War God Spring on Navajo Mountain, 6 July 1936. Others were seen or heard calling from 6, 12, 20, and 21 July; they made additional observations at Beaver Creek, eight miles north of Navajo Mountain, 8 and 9 Aug. Resident, breeding in Piniis ponderosa and pygmy conifers. Bubo virginianus pallescens Stone. Great Homed Owl. Presnall (1934) and Behle et al. (1958) Bryce Canyon; Woodbury and Russell (1945) Navajo Mountain at 9000 ft; BYU (1971-1973) specimen: Site 1, near Glen Canyon City, 29 Aug. 1971; Site 1, 16 Feb. 1972 (grasslands); Tibbet Spring, 15 Apr. 1972 {Populus fremontii, washbottom); BYU (1973) Four Mile Bench, 13 May. Permanent resident. Glaucidium gnoma californicum Sclater. Pygmy Owl. Russell and Thompson (1964) Bryce Canyon. Permanent resident. Athene cunicularia hypugaea (Bonaparte). Burrowing Owl. Phillips et al. (1964) south- east portion of the Kaiparowits Basin; BYU (1972) Cottonwood Wash, 9 Aug. (grassland); five miles west of Glen Canyon City, 4 June and 21 July; four miles west of Glen Canyon City {Artemisia fUifoIio) sitting in burrow on side of road; U.S. 89/Cottonwood Wash Road, 22 and 23 July; BYU (1973) three to four miles west of Glen Canyon City, 5 June (open grassland) nest under construction nearby; BYU (1974) Utah/Arizona state line north of Page on U.S. 89. Uncommon sum- mer resident. Strix occidentalis lucida (Nelson). Spotted Owl. Woodbury (1939) reported that this species was collected 13 Aug. 1936 in Navajo Canyon by H. N. Russell, Jr.; Behle (1960) Davis Gulch, a tributary of the Escalante River, 55 miles southeast of Escalante, Aug. 1957; two were seen several times in a small side canyon of Glen Canyon near river mile 101, 17 July, and another at the mouth of the Escalante River, 19 July 1958. Uncommon permanent resident. Asio otus (Linnaeus). Long-eared Owl. BYU (1973) Site 30, 9 June (grassland). Per- manent resident. Caprimulgidae (Goatsuckers) Phalaenoptilus nuttallii nuttallii (Audu- bon). Poorwill. Woodbury and Russell (1945) Navajo Mountain Trading Post, 22 July 1936, and Kaiparowits Plateau, 29 July and 3 Aug. 1937 (all in pygmy conifers); common in Aug. On Navajo Mountain, nests were found containing two downy young 23 July 1936 in pygmy conifers, a family of three poorly feathered juveniles in pygmy forest, Kaiparo- wits Plateau, 7000 ft, 29 July 1937, and sim- ilar family on 3 Aug. not far away in Arte- uiisio-Quercus; Russell and Thompson (1964) Bryce Canyon; BYU (1971) Paria Plateau, 29 Sept.; BYU (no dates) Site 3 and Church Wells. A recent decline in numbers is evi- denced by the paucity of sightings in the present study as compared to the relative abundance of the species in earlier studies. Uncommon summer resident. Chordeiles minor henryi Cassin. Common Nighthawk. Presnall (1934) Bryce Point; Ben- son (1935) Navajo Mountain; Tanner (1940a) Escalante drainage; Woodbury and Russell (1945) top Navajo Mountain, 14 July 1936; BYU (1971-1973) Sites 2, 3, 4, and 28; Glen Canyon Dam; one mile west, six miles east, and 2.5 miles southwest of Grosvenor Arch; and Four Mile Bench. Nighthawks were seen December 1980 Atwood et al.: Kaiparowits Vertebrates 319 IS early as 19 May (1972) and as late as 5 3ct. (1972). Monthly distribution of observa- :ions during our study is one in May, 8 in [une, 26 in July, 37 in Aug., none in Sept., ind one in Oct. During June and Julv night- lawks were recorded in the following vege- :ation types and in the following niuubers: 3oleogyne (2), mixed shrub (1), Graijia (1), 'iiniperus (6), and grassland (7); from August ^n they were distributed as follows: Coleog- /tie (1), grass (35), Juniperus (9), and Arte- nisia (1). Common summer resident. Apodidae (Swifts) Aeronatites saxatalis saxatalis (Wood- louse). White-throated Swift. Presnall (1934) 3ryce Canyon; Benson (1935) Navajo Moun- :ain; Tanner (1940a) Kaiparowits Basin; ^oodburv and Russell (1945) one, Kaiparo- wits Plateau, 27 Julv 1937 (7000 ft); Behle et il. (1958) Escalante River/Calf Creek; Hay- A'ard et al. (1958) Escalante drainage; Behle 1960) Coyote Gulch and Kaiparowits Plateau, Aug. 1957; Behle and Higgins (1959) ^ive numerous observations for lower Glen Hanyon from the field notes of Woodbury ind Russell. In these notes a colony of swifts was observed in a crevice of a cliff at the iiouth of Rock Creek, 20-26 July 1937; six at iiile 2 on the San Juan River, 1 Aug. 1938, md others were seen between miles 41 and 15, 8 Aug., and at mile 25, 9 Aug. BYU 1971-1973) Sites 1, 2, 3, 4, 10, and 23, Cot- :onwood Wash, Hackberry Canyon, Lee's Ferry, Kelly Grade, Tibbet Spring, Wiregrass spring, Reflection Canyon, Ribbon Canyon, md Three Garden one mile above San Ju^n •onfluence with Lake Powell. Although most ;wifts were seen near cliffs, those migrating were seen away from cliffs and over various