é Futye ne ota reel “ ; fe eee ne #32 345 sane yi A 3 Oe a ekg eer SN bes eee Poe Rome St . ¢ . ade Pood x Aas fede CEOS? pe ea . . a ie : dir A hie at ‘ Se RATA? hire nynns : oT et . nas ay NERC OE, 5 4 . ae 5 ae aot eR UT : ore hes : tytn - 4 . ry eo) eee aay Cee 5 : le | rot Cerra iy . 2A fetes : Singis AS ARMED Ane Re, biter ia se EA at thot atts ; rh c ; Bay yes o han an we ‘ eB we akan spar tary ‘ Path , 2 yin . ans ae aos aoe ASN =n : . ta A ‘ bag > . i . sae Cee en 5 4 : “Oye tA eer ye bs 4 a Cpa tee date Note iy H Ais rere avasenyn yt Ne em ba Angi st : 4 ; P “eta < * m4 Held ee ee MA Nh Ban LL 8 <2 : hat . é fae vehi Asotin Rade ea Hada ME ; : arte Coded Aman See ents aan Bue roe Ema " ‘ . har ate . yen?) Es Deana ine me Oty Obe ria bear ae Ms Sirdar ir. W9e42A9o steed et needs - Srl alee sy deny tent When Leg) ad el ee vie dm int VERE De iMintae ada e ntti dns ae Ree Oe a reed on Mas shake. erik pren dey) 4 ‘aie a” Lie a oe eA S a i tad ey tay St RN fe tite, ae Wek, ekg pala dy Cpe atee tenes yedes a Faju a Fes yee bbe page ire AU age Cae Peds hfe, aoe Wphuadis ogee ads fag aoe Te Pree Lew earl ATK ages ato Paani y See eee Fi OY Rn ae ad Pane eee a Lb iss 4! ged. Tagish eth ats ee SEI IES) Ve geweed 4 Erbe ie oie ee eee ey ee eee ord id baer eel Caged fae der gtseay cag vee bere jeges TT EP ey ae ee CE oe Et ahaa iavets oF seer ae er are yet ta ie piiaige Ghd Sep Fear d lye Hea eat DEVE atin) BoD tae eh ade grad greeny VeGRpa bh pide hg eyeg pean eds ts fhe zed Le Ggipe eA oes Lae deg ayetisg pdt ata et: il nah gad pighe Pir S0z PTTEREMLV IU AL OTC U REL Te ies Cea eae aA NEP 4 EE] PFA ee ie EERE P AS OV A TIALE EDIE fe ee A ALF Lied ATi spades PUREE Ree ea) igs Te Paee eR EN Ee leo | seeped Sea AY “BF aid ag thar SY AGIL TAS P93 Eo + pha deib eae ahs Dh gadt ree pun ia eg Pree piney cag errr e er ae EVAN EI FL Ay TN LPL EE dP tet a MOgsIE 4 | BIL EE RL Thi a L1G VE Mab pip bisa crass apse A ie 2 Tg SVE 8a UE SEe pa ew EM Ed tT ALS Pees eer a Le ea a eae Aires belt 4 af 422) swiwie z: paeaf ii? A GPT Fh den 4854 fa bgigry! Me Pubtpoeraeaey e156 -d2 pte Lee Aa} prahirees ere are Pee tues oss Perey Ee Soyo op ea at be fea taay hin Varta 32 pouty} ee be chee ashe? BISLAN ET GS EBL T seg C255 ARE TE mena re ad LEA SALMO aA; eee CU ea ge fea te Vg nd te a tponent Z HARVARD UNIVERSITY e Library of the Museum of Comparative Zoology : : ty 7 s - = ie ) 7 4 ~ Ré rT s : i rey © ae 7 *; “ MH : * x ) “ Ss - J A 2 os oe ™ soe = 2 1 - . 2) ra 7 ss Eat t i i a) i 7 ie - - ; = wr =) e | = > S 7 . ¢ < i ves 1 ‘ 7 , wal : i t - { t . a y : . n The Great Basin Naturalist VOLUME 46, 1986 EDITOR: STEPHEN L. Woop PUBLISHED AT BRIGHAM YOUNG UNIVERSITY, BY BRIGHAM YOUNG UNIVERSITY TABLE OF CONTENTS Volume 46 31 January 1986 (Number 1) a4 a Dynamic landforms and plant communities in a pluvial lake basin. James A. Young, Raymond A. Evans, Bruce A. Roundy, and John’A® Brown) 24-4250 eee Vegetation and flora of Pine Butte Fen, Teton County, Montana. Peter Lesica. ....... Effects of suspended sediment on leaf processing by Hesperophylax occidentalis (Tri- choptera: Limnephilidae) and Pteronarcys californica (Plecoptera: Pteronarcidae). ©. Evan Homig and Merlyn A; Brusven. 9-..-25...-- 45. 40) cee Effects of watershed alteration on the brook trout population of a small Black Hills stream. Timothy Modde, Henry G. Drewes, and Mark A. Rumble.................... Floristic analysis of the southwestern United States. Steven P. McLaughlin. ......... Utah flora: Apiaceae (Umbelliferae). Sherel Goodrich.....................:..52555 Quercus (Fagaceae) in the Utah flora. Stanley L. Welsh. .......................... Seasonal phenology and possible migration of the Mourning Cloak butterfly, Nymphalis antiopa (Lepidoptera: Nymphalidae) in California. Arthur M. Shapiro. ......... Seasonal microhabitat relationships of blue grouse in southeastern Idaho. Dean F. Stauffer and Steven R: Peterson... . 4. .0.. ciememes oon: 0 eee Fall diet of blue grouse in Oregon. John A. Crawford, Walter Van Dyke, S. Mark Meyers, and Thomas F.. Haensly. .s..... Se: eee 0 SE. 12 ee Status and distribution of California Gull nesting colonies in Wyoming. Scott L. Findholt.. sac. 0-84 5 ces Reng oe SOS oe New genus and species of leafhopper in the tribe Tinobregmini (Homoptera: Cicadelli- dae: Coelidiinae). M. W. Nielson. .... 2... 2... 29... ene 15 Oe New oriental genus of leafhoppers in the tribe Coelidiini with descriptions of new species (Homoptera: Cicadellidae: Coelidiinae). M. W. Nielson. ......0.............. Ecological differences of C, and C, plant species from central Utah in habitats and mineral composition. C. Morden, Jack D. Brotherson, and Bruce N. Smith............. Response of winterfat (Ceratoides lanata) communities to release from grazing pressure. Lars L. Rasmussen and Jack, Brotherson.....:......5.. 5.000 CeeeEeeeee Isozymes of an autopolyploid shrub, Atriplex canescens (Chenopodiaceae). E. Durant McArthur, Stewart C. Sanderson, and D. Carl Freeman....................-. Winter nutritive content of black sagebrush (Artemisia nova) grown in a uniform garden. Barbara Behan‘and Bruceils, Welch. <:-...2.472 6 44.. 4400 oee eee Winter food habits of the pine marten in Colorado. Christine C. Gordon. ............ Foliage age as a factor in food utilization by the western spruce budworm, Choristoneura occidentalis. Elizabeth A. Blake and Michael R. Wagner. ..................-- Apple maggot (Rhagoletis pomonella) adaptation for cherries in Utah. Clive D. Jor- gensen, Darin B. Allred, and Richard LL. Westcott... ). 5...) sone New records for Monotropa hypopithys (Ericaceae) from Colorado. William Jennings, Loraine Yeatts, and Velma Richards.....)..-.- 594.4500 4-2 Tree densities on pinyon-juniper woodland sites in Nevada and California. Susan Komik isis as, aja evs 4 site oud esas Seegily 4e es 30 April 1986 (Number 2) Biology of Red-necked Phalaropes (Phalaropus lobatus) at the western edge of the Great Basin in fall'migration. Joseph R; Jehl} Jr. .245..5- 5-55 oe Some relationships of black-tailed prairie dogs to livestock grazing. Craig J. Knowles. . . ‘ul 2B) Andersen, Lauritz A. Jensen, H. Dennis McCurdy, and Craig R. Nichols. ...... Jam-raised fawns, an alternative to bottle feeding. Kathrin M. Olson-Rutz, Philip J. Une sswan cduleaunay Ama imMessucN Nn a nares een eas La ai ean peti OP ee bia as ubspecific identity of the Amargosa pupfish, Cyprinodon nevadensis, from Crystal Spring, Ash Meadows, Nevada. Jack E. Williams and James E. Deacon. ........ wo aberrant karyotypes in the sagebrush lizard (Sceloporus graciosus): triploidy and a “supernumerary oddity. Pamela Thompson and Jack W. Sites, Jr.............. ood habits of clouded salamanders (Aneides ferreus) in Curry County, Oregon (Am- phibia: Caudata: Plethodontidae). John O. Whitaker, Jr., Chris Maser, Robert M. y Stonmmyancyjosephyjarbeattyset ms dase vie ec aS 2 TRE ome 2 ANA \Vintering bats of the upper Snake River Plain: occurrence in lave-tube caves. David L. (| (GS INTEN BS at (Gnd aie ts een eg aac le ae tt a Tr ois OI Jan Rg W/o te srowth rates of mule deer fetuses under different winter conditions. Richard M. Bart- LUMENS a col'6 asc ire earls egae gee eee Up ya ag a UU RM, aT eo Denning habitat and diet of the swift fox in western South Dakota. Daniel W. Uresk and om@mshanoss saree ee er neti Miers ir re Nees ne ener eee aN * New taxa and combinations in the Utah flora. Stanley L. Welsh..................... New taxa in miscellaneous families from Utah. Stanley L. Welsh.................... "New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), Part } URES Cerone rye WOO Grae cies ey ore ute ae cea i ent emake s crate ertagn /Snergy and protein content of coyote prey in southeastern Idaho. James G. MacCracken AOeRIChanGeN les blanseMs chy fs ee oi ene, satiny soem sateen aan wage SE /istimates of site potential for Douglas-fir based on site index for several southwestern habitat types. Robert L. Mathiasen, Elizabeth A. Blake, and Carleton B. Edmin- | anne sia vee MeArthursandi Gaehma@ushing-0 Vengeance ee ye eae : hree-year surveillance for cestode infections in sheep dogs in central Utah. Ferron L. 3 . i ffect of excluding shredders on leaf litter decomposition in two streams. James R. | | /Wintering mule deer preference for 21 accessions of big sagebrush. Bruce L. Welch and | ID Unt CAN EMU gece ee cumin BUN heey FiOs Ue nia lame Gal celta ‘Coleoptera of the Idaho National Engineering Laboratory: an annotated checklist. ') Michael P. Stafford, William F. Barr, and James B. Johnson................... Microhabitat affinities of Gambel oak seedlings. Ronald P. Neilson and L. H. Wullstein. feeding habits of metamorphosed Ambystoma tigrinum melanostictum in ponds of high pre 9) brianveMillerand johnvH. Warsen, Jr0-.6...- 4-242 a0eo0e 49 aco sae Notes on the Swainson’s Hawk in central Utah: insectivory, premigratory aggregations, ancikleptopanasitisms NeiliD ss \Wotimdens me asc. 424 sache odes 16 oon ce Turkey vultures decline at a traditional roosting site. Daniel M. Taylor............... Barn owl diet includes mammal species new to the island fauna of the Great Salt Lake. an UID aN iartisane er pe een os rar he a aim) dalarae csevnue Weald cians 6 MRO Distributional study of the Zion Snail, Physa zionis , Zion National Park, Utah. David Ng TAG ATI SHED ATaIn Seep are Me seh) NOR cans ohce wali hu hal beard tenant uname acini 22% Blockage and recovery of nitrification in soils exposed to acetylene. Steven J. Burns and Rite seme A ed Gharstike ois cece bcos ne ma ete enor Smarr cee ates Bos eg eae er New thagriine leafhoppers from the Oriental Region, with a key to 30 species (Ho- moptera: Cicadellidae: Coelidiinae). M. W. Nielson. ..................00005: Genus Paralidia with descriptions of new species (Homoptera: Cicadellidae: Coelidi- TVA) MeV BNC Smee: Sree ie carb oer nia sein nis cdavgule ta are i ouita) seca tLe ee Montane insular butterfly biogeography: fauna of Ball Mountain, Siskiyou County, @aliformiagArthiumVs Shapinowe: sateen a ee ee oe tine it Comparative habitat and community relationships of Atriplex confertifolia and Sarcoba- tus vermiculatus in central Utah. Jack D. Brotherson, Lars L. Rasmussen, and Rich ance lackamee yard ete oe ters Aiea dit ani. yap Nemes ee ceodemeeet let 204 217 348 Trees used simultaneously and sequentially by breeding cavity-nesting birds. Kevin J. IN Gutzwiller and|Stanley He Anderson; 4240445 0 oboe eee 35) New species of Mentzelia (Loasaceae) from Grand County, Utah. Barry A. Prigge...... 36) N Utah flora: Juncaceae. Sherel Goodrich). 3). 3.4045-3; 25-0085 noe eee 36" Advertisement call variation in the Arizona tree frog, Hyla wrightorum Taylor, 1938. WN Brian K.. Sullivans: cect edn ees vk es sates Ee Oe STA 31 July 1986 (Number 3) g North American stoneflies (Plecoptera): systematics, distribution, and taxonomic refer- \0 ences. B. P: Stark,.S: W. Szezytko; and|R: W. Baumann...) -) 2 38, Three new records for diatoms from the Great Basin, USA. Samuel R. Rushforth, Lorin iG E. Squires, and Jeflrey R. Johansen). 0..0.....-). +42. 54+ dee eee 39 Movements by small mammals on a radioactive waste disposal area in southeastern Idaho. | Craig R. Groves and Barry L.Keller: =. -.<..0552...- 3) oe 40) | Roll of three rodents in forest nitrogen fixation in western Oregon: another aspect of mammal—mycorrhizal fungus—tree mutualism. C. Y. Li, Chris Maser, Zane Maser, i and Bruce A: Caldwell. o.002 0. Shoe gue beens bee bone cee ee 41 Canids from the late Pleistocene of Utah. Michael E. Nelson and James H. Madsen, Jr. a Note on food habits of the Screech Owl and the Burrowing Owl of southeastern Oregon. | Barbara A. Brown, John O. Whitaker, Thomas W. French, and Chris Maser. .... 42} | Diseases associated with Juniperus osteosperma and a model for predicting their occur- | rence with environmental site factors. E. D. Bunderson, D. J. Weber, and D. L. | Nelson: ¢ sicccc ee ce cone ss Re ns Ee eet Os Bi ec rr Status and distribution of the Fish Creek Springs tui chub. Thomas M. Baugh, John W. Pedretti, and James E..Deacon... 2.02. 0.205.505 bes oe Ponderosa pine conelet and cone mortality in central Arizona. J. M. Schmid, J. C. Mitchell, and'S\ A. Mata. ... 2. 0050 00. ake k oa a a ee Number and condition of seeds in ponderosa pine cones in central Arizona. J. M. Schmid, S. A.’ Mata, and J.C. Mitchell oc. ee a New species of Protocedroxylon from the Upper Jurassic of British Columbia, Canada. — David A. Medlyn and William D: Tidwell) <7.) -. 4a. c ae oe eee New taxa and nomenclatural changes in Utah Penstemon (Scrophulariaceae). Elizabeth GHNEESE: ook iis ee eld ates s toms ie UE ce eee Relict occurrence of three “American” Scolytidae (Coleoptera) in Asia. Stephen L. Wood and Hui-fen Yin... 0.3.0.0... 05.) s SEE eee New genus of Scolytidae (Coleoptera) from Asia. Stephen L. Wood and Fu-sheng AU aN oie ce ees een Se ee es ae eo oink ee 8 New Pseudoxylechinus (Coleoptera: Scolytidae) from India. Stephen L. Wood. ....... | Wildlife distribution and abundance on the Utah Oil Shale Tracts, 1975-1984. C. Val ) Grant e7 heh ese oe i Rs Fs Eee OEE OO 46! Comparison of vegetation patterns resulting from bulldozing and two-way chaining on a Utah pinyon-juniper big game range. J. Skousen, J. N. Davis, and Jack D. Brother- | SOM 3) 2) eo there Se ee SG al I ok I, Ce et 50% Vertebrate fauna of the Idaho National Environmental Research Park. Timothy D. ) Reynolds, John W. Connelly, Douglas K. Halford, and W. John Arthur. ........ ole Infection of young Douglas-firs and spruces by dwarf mistletoes in the Southwest. Robert } L.Mathiasen: ..f 0. O00 be OP 5284 Habitat relationships of saltcedar (Tamarix ramosissima) in central Utah. Jack D. Broth- | erson and Von Winkel. 2.0.04... 4) l65. DO ee 53h Characteristics of mule deer beds. H. Duane Smith, Mark C. Oveson, and Clyde L. L Pritchett siaincd wd oe eG oe eR ee a ee ee 54o | Trumpeter Swan (Cygnus buccinator) from the Pleistocene of Utah. Alan Feduccia and Ff Charles G. Oviatti.i... cs go8 se Oh PSG hay one Vg oR eee 347 | ew species and a new combination of Mentzelia section Bartonia (Loasaceae) from the Colorado Plateauyle @hompsonand)B. Brigge.. 5554. .045. eee : ‘ew variety of Mentzelia multicaulis (Loasaceae) from the Book Cliffs of Utah. Kaye H. iiironmmerand trankseoimithams site ci te Coad ee ee ea Ma ‘ [ew variety of Mentzelia pumila (Loasaceae) from Utah. Kaye H. Thorne. ........... gathoxylon lemonii sp. nov., from the Dakota Formation, Utah. William D. Tidwell and Creconseehayn eras Vas Here Meh ems ee mi la ek BAN yl jsrazing and passerine breeding birds in a Great Basin low-shrub desert. Dean E. | IMIG GLFg: 5 S18 Sie kare ee cos al AU My toler 8 ane Ree en ocean ee oR hloroplast ultrastructure in the desert shrub Chrysothamnus nauseosus ssp. Albicaulis . Cracsh Colemanjand WalliamsRyAndersenyay- ana) a... »;enetic variation of woodrats (Neotoma cinerea) and deer mice (Peromyscus manicula- ; tus) on montane habitat islands in the Great Basin. William T. Mewaldt and Stephemibea|emkinse pore arcs Ninen eet sist cyte sae Ny (sreeding records for Clark’s Grebe in Colorado and Nevada. Richard L. Bunn 4 31 October 1986 (Number 4) | ! Jistory of fish hatchery development in the Great Basin states of Utah and Nevada. J. W. SigdeimancaWep bee SiGle rrr Niel rtns Meuertae eee lle nn ele tan, aes } Composition and abundance of periphyton and aquatic insects in a Sierra Nevada, California, stream. Harry V. Leland, Steven V. Fend, James L. Carter, and Albert TD), INU Oraye lot ya): as ita uc tet cae ere RE reise ice TR Gt agian eRe ) Diatom flora of Cowboy Hot Spring, Mono County, California. Laura Ekins and Samuel i ROBINS nbort ntipee err sete coi loro otialniintaiaila sie ulule nears ON eM RAS RC es Ne) ertea | nventory of Utah crayfish with notes on current distribution. James E. Johnson....... cryptogamic soil crusts: recovery from grazing near Camp Floyd State Park, Utah, USA. i Jetineyvakjohansemandélearny Myst Claim cre ns ae a ese New species and new records of North American Pityophthorus (Coleoptera: Scolytidae), : ant Agsbneseautus croup. Wonaldsb. Brightsa-c > be sae snes initial survey of acetylene reduction and selected microorganisms in the feces of 19 species of mammals. C. Y. Li, Chris Maser, and Harlan Fay................... ‘iotes on the birds of Cold Spring Mountain, northwestern Colorado. Peter O. Dunn and Vora ARM Le Terre eine ae ee ie te me te ey CsI ES AOk pe Di Life, strategies in the evolution of the Colorado squawfish (Ptychocheilus lucius). Harold Eee US treet uate Naa st ale GOL So ntate wlatu ec lala Gok heey eM ato toamt east: Parasites of the woundfin minnow, Plagopterus argentissimus , and other endemic fishes from the Virgin River, Utah. Richard A. Heckmann, James E.. Deacon, and Paul D. New Sclerocactus (Cactaceae) from Nevada. Ken Heil and Stanley L. Welsh.......... New species and new records of North American Pityophthorus (Coleoptera: Scolytidae) arta lle Ormal ue BTiS ty eave ney. aera a dere arc eis cmPaud Cale glurelnye Effects of dwarf mistletoe on spruce in the White Mountains, Arizona. Robert L. ~Mathiasen, Frank G. Hawksworth, and Carleton B. Edminster................ Hydrology of Bear Lake Basin and its impact on the trophic state of Bear Lake, Utah- Idaho. Vincent Lamarra, Chuck Liff, and John Carter. ....................5.- Understory seed rain in harvested pinyon-juniper woodlands. Richard L. Everett. .... Species diversity and habitat complexity: does vegetation organize vertebrate communi- ties in the Great Basin? David J. Germano and David N. Lawhead. ............ Comparison of insects from burned and unburned areas after a range fire. James D. ID IBYNKE) TY a -n.0 ab) 6:5 o.acove cueticabccle ato Recent oan lice By cae mars a rn tac ara eRe merry errr ear meroa Size, structure, and habitat characteristics of populations of Braya humilis var. humilis (Brassicaceae): an alpine disjunct from Colorado. Elizabeth E. Neely and Alan T. Wareniter wee ee eS Sg cats) ae BONS aster) Nyanee ROR A GS Soaks 549 599 507 509 567 573 O77 581 583 595 612 625 632 641 646 651 656 662 677 679 685 690 706 711 721 728 Biogeographic aspects of leeches, mollusks, and amphibians in the Intermountain Re- gion. Peter Hovingh: .. 0.3.2 sc.d $005. 05 Jes eed Hatching chronology of Blue Grouse in northeastern Oregon. John A. Caneen Walter Van Dyke, Victor Cogeins, and Martin St. Louis. .-..--922. 4. seep eee New South American leafhoppers in the genus Docalidia, with a key to 37 species (Cicadellidae: Coelidiinae, Teruliini). M. W. Nielson. ....................... Fossil birds of the Oreana local fauna (Blancan), Owyhee County, Idaho. Jonathan J. Becker... eos: Cee eto Oe Ea a ee 7 IEGREAT BASIN NAIURALIoT ie 46 No. 1 31 January 1986 Brigham Young University MCZ LIBRARY 8 hip ANY Z CRT if : ali (Uy D> Bi> |. WS SS Was 7 pal oS a) — WAC NS KP AY >. A - SS = { = GREAT BASIN NATURALIST Editor. Stephen L. Wood, Department of Zoology, 290 Life Science Museum, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Chairman, Botany; James R. Barnes, Zoology; Hal L. Black, Zoology; Stanley L. Welsh, Botany; Clayton M. White, Zoology. All are at Brigham Young University, Provo, Utah 84602. Ex Officio Editorial Board Members. Bruce N. Smith, Dean, College of Biological and Agricultural Sciences; Norman A. Darais, University Editor, University Publications. Subject Area Associate Editors. Dr. Noel H. Holmgren, New York Botanical Garden, Bronx, New York 10458 (Plant Taxonomy). Dr. James A. MacMahon, Utah State University, Department of Biology, UMC 53, Logan, Utah 84322 (Vertebrate Zoology). Dr. G. Wayne Minshall, Department of Biology, Idaho State University, Pocatello, Idaho 83201 (Aquatic Biology). Dr. Ned K. Johnson, Museum of Vertebrate Zoology and Department of Zoology, University of California, Berkeley, California 94720 (Ornithology). Dr. E. Philip Pister, Associate Fishery Biologist, California Department of Fish and Game, 407 West Line Street, Bishop, California 93514 (Fish Biology). Dr. Wayne N. Mathis, Chairman, Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 (Entomology). Dr. Theodore W. Weaver III, Department of Botany, Montana State University, Boze- man, Montana 59715 (Plant Ecology). The Great Basin Naturalist was founded in 1939 and has been published from one to four times a year since then by Brigham Young University. Previously unpublished manuscripts in English of fewer than 100 printed pages in length and pertaining to the biological natural history of western North America are accepted. Western North America is considered to be west of the Mississippi River from Alaska to Panama. The Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natural history longer than can be accom- modated 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 for private individuals is $16; for institutions, $24 (outside the United States, $18 and $26); and for student subscrip- tions, $10. The price of single issues is $6 each. All back issues are in print and are available for sale. All matters pertaining to subscriptions, back issues, or other business should be directed to Brigham Young University, Great Basin Naturalist , 290 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. See Notice to Contributors on the inside back cover. 6-86 650 23497 ISSN 017-3614 The Great Basin Naturalist PUBLISHED AT PROVO, UTAH, BY BRIGHAM YOUNG UNIVERSITY ISSN 0017-3614 VOLUME 46 31 January 1986 No. 1 DYNAMIC LANDFORMS AND PLANT COMMUNITIES IN A PLUVIAL LAKE BASIN James A. Young’, Raymond A. Evans', Bruce A. Roundy’, and John A. Brown? ABSTRACT. — Mapping units were developed based on landform, soils, and plant communities for the 40,000 ha of salt desert vegetation located below the maximum level of pluvial Lake Gilbert in Grass Valley, Nevada. Lake plain, beach, and bar features of the pluvial lake provide the dominant landforms. Fine textured lake sediments have produced salt- and Na-affected soils. The distribution and structure of plant communities are related to depth of the groundwater table, fluctuations in this depth, and the salt content of the groundwater. Wind and water erosion combine to continually evolve new environments for colonization by plants in this ecosystem. The structure and association of plant com- munities occupying basins in western North America that were formerly inundated by plu- vial lakes has long been of interest to ecolo- gists (Shantz and Piemeisel 1940, see West 1983 for description of ecosystems). The pro- gressively finer textured soils from the edge of the basins and increasing concentration of sol- uble salts that formed as the lakes dried of- fered the potential of relating plant communi- ties to soils, and especially to the salt content of soils (Kearney et al. 1914, Clements 1920, Billings 1945). Such salt-plant community ori- entations are complicated by the extreme aridity of many pluvial lake basins. In the Great Basin of the western United States, the entire province is located in the rain shadow of the Sierra Nevada and the Cascade Moun- tains. In addition, the mountains that rim many of the subbasins range from 2,150 to more than 3,000 m in elevation, casting their own rain shadows across the arid bottoms of the valleys (Houghton et al. 1975). The result is that the natural vegetation of the valley bottoms may reflect lack of atmospheric pre- cipitation as much as the reduced osmotic potential of soil water solutions (Billings 1949). The concept of pluvial basins containing concentric rings of soils with increasing salt content helps explain the distribution of many salt desert plant communities (Flowers 1934). The occurrence of other plant communities such as those dominated by Ceratoides lanata or Atriplex confertifolia is not associated with specific soil features (Gates et al. 1956). The distribution of Atriplex nuttallii on soils from low to high salinity is a good example of eco- typic differences within a species in relation to salt tolerance (Goodman 1973). Genetic dif- ferences must be considered when relating plant distribution to edaphic factors in the salt desert. Stutz (1978) has presented evidence that the new habitats provided by the drying of pluvial lake basins provide space for the explosive evolution of perennial species of Atriplex. Some of the landscape occupied by species of Atriplex exist in complex polyploid 1U_S. Department of Agriculture, Agricultural Research Service, 920 Valley Road, Reno, Nevada 89512. Formerly with Agric. Res. Ser., currently at University of Arizona, Tucson. “Formerly research hydrologist, University of Nevada, Reno. Deceased. Plant names based on Cronquist et al. (1972) and Munz and Keck (1968). Plant specimens-on file USDA/ARS Herbarium, Reno, Nevada. 2 GREAT BASIN NATURALIST series. These desert landscapes dominated by woody chenopods apparently have limited re- generation and appear, superficially, to be composed of nearly identical individuals. Ac- cording to Stutz’s hypothesis, the endless sameness of woody chenopods is a mirage con- cealing dynamic evolutionary processes. Distinct patterns of vegetation on pluvial lake sediments were postulated by Miller et al. (1982) to be due to differences in soil-wa- ter-plant relations. Differences were caused either by the depth to groundwater or from differences in water-retention capacities of soils deriving water only from precipitation. In a recent review, West (1982) logically refuted the stereotype concept that salt desert shrubs exist in spatial harmony in equilibrium with the very limited environmental potential of their environment. In fact, chenopod shrubs tend to be grouped in competitive clumps where litter-fall on coppice mounds has enhanced the potential of seedbeds to support germination and seedling establish- ment (Charley and West 1975). The chance recruitment of seedlings into these communi- ties may be conditioned by erratically occur- ring episodic climatic events (West 1979). Due to the lack of atmospheric precipita- tion and subsequent runoff water, many of the lower pluvial lake basin environments appear remarkably stable once the soil surface be- comes stabilized by vegetation and desert pavement formed by wind erosion. We pro- pose that much of this apparent stability is as much a mirage as the genetic stability of the shrub populations. The pluvial lakes shaped the basin bottoms with currents and waves. Sedimentations in the deep water areas were usually very fine textured. Lowering the wa- ter level in the lakes lowered the base level of the attendant streams. Landforms and hydro- logic processes in and surrounding pluvial lake basins are striving toward stability in a radically different environment. Our purpose was to characterize the major plant communi- ties of a pluvial lake basin in relation to land- forms and soils. METHODS The study was conducted in Grass Valley, Nevada, 39°52’ latitude, 116°37’ longitude. Grass Valley is aclosed basin with a watershed Vol. 46, No. 1 basin of 1,500 km*. During the Pleistocene, the basin contained pluvial Lake Gilbert, which had a surface area of 400 km?* (Mifflin and Wheat 1979). The maximum level of Lake Gilbert was 1,766 m. The bottom of the basin is now 1,728 m. Some authorities (e.g., Hubbs and Miller 1945) consider that Lake Gilbert overflowed at its maximum level and formed part of the Lake Lahontan systems. Mifflin and Wheat (1979) point out that the maximum beach ridge of Lake Gilbert is 39 m below the supposed outlet pass. Color infrared aerial photographs, scale 1:15840, were used for identifying landforms and drainage patterns; plant communities were mapped on 1:62500 U.S. Geological Survey maps. Shrub vegetation in each com- munity was sampled with 10 plots, each 10 m? in area. The plots were randomly located along paced transects from a starting point selected on the aerial photographs. The pro- jected crown cover, height, and density of shrubs were recorded by species. The herba- ceous vegetation and cryptogamic soil crust (Anderson 1978) were sampled for frequency and cover with four replications of 100 step points using the procedures described by Evans and Love (1957). Soil prefiles were excavated in each major vegetation assemblage. Profiles were de- scribed according to the Soil Survey Manual (Anonymous 1951). Soils were classified ac- cording to the U.S. Soil Conservation Service system (Anonymous 1975). Soil samples were analyzed for percent gravel by straining through a 2-mm screen; percent sand, silt, and clay were determined using the Bouyocus (1962) method. Electrical conductivity of the saturated soil paste extract was determined by a conductivity meter (Black 1965). In 1981 a network of shallow wells was in- stalled in three transects from the alluvial fans across the lake plain to the central playa. The wells were distributed along the transects on the basis of plant communities and landforms. The wells were drilled to 4.6 m, with a 10-cm rotary drill and cased with perforated plastic pipe 3.75 cm in diameter, and gravel was packed around the casing. The depth of the water table was measured periodically, and samples were collected and analyzed for elec- trical conductivity and temperature. A por- table signal-enhanced seismograph was used | \ | i i i January 1986 YOUNG ET AL.: LAKE BASIN COMMUNITIES 3 TABLE 1. Landforms, vegetation, soil texture, percentage of total area, and number of stands sampled for the land area below the maximum level of pluvial Lake Gilbert, Grass Valley, Nevada. M ap units refer to Figure 1. Landform Map Vegetation Soil Area Percentage Number Major subunit unit Dominant species texture ha oftotal of stands PLAYA Playa 1 None Clay 6040 15.0 64 Wet depression la None Clay 10 eas) 3 Hummocks lb Allenrolfoa occidentalis Silty clay loam 10 — 3 Mobile dunes 2 Sarcobatus vermiculatus Clay 120 0.3 12 Haystack dunes 2a Sarcobatus vermiculatus Clay 10 _ 6 Recent Holocene beach 2b Kochia americana Clay 10 as 2 6200 15.4 LAKE PLAIN Older Holocene beach 3 Sarcobatus vermiculatus Silty clav loam 2660 7.0 3 Stable dunes 4 Sarcobatus vermiculatus Silt loam 160 0.4 3 Clay dunes 5 Sarcobatus vermiculatus Clay 380 0.9 ll Dune basins 6 Distichlis/Sporobolus Clay 50 1.4 2 Lake plain 7 Chrysothamnus-Sarcobatus- Silty clay loam 2130 5.3 7 Artemisia/Elymus 8 Sarcobatus/Distichlis Silty clay loam 90 0.2 5 9 Sarcobatus-Atriplex Silty clay loam 2160 5.4 il confertifolia 10 Chrysothamnus albidus/ Silty clay 10 — 3 Puccinella 11 Atriplex nuttalii Silty clay loam 30 0.1 3 8070 =. 20.2 Thermal springs 12 Distichlis/Juncas Clay 460 Mel 2 Barrier beach 13 Sarcobatus vermiculatus Clay 820 2.0 2 14 Sarcobatus/Kochia Clay loam 190 0.5 2 15 Atriplex confertifolia Silty clay loam 116 0.3 2 1120 2.8 Lagoon 16 Allenrolfea occidentalis Clay 1470 Oat 4 17 Chrysothamnus nauseosus/ Silty clay loam 990 2) 3 Distichlis eima 2460 6.2 Sand dunes 18 Tetradymia/Grayia Sand 130 0.3 9} Alluvial fans 19 Artemisia tridentata Loam to gravelly 7350 18.3 12 sandy loam 20 Atriplex confertifolia Silty clayloam 13760 34.3 21 to gravelly sandy loam 21110 52.6 Bars 21 Atriplex confertifolia Gravelly sandy 370 0.9 3 Lagoons (upland) 22 Sarcobatus vermiculatus Silt loam 130 0.3 3 23 Atriplex confertifolia Silt loam 60 0.2 3 190 0.5 in the summer of 1981 to determine depth to the water table at 35 locations in the valley. The seismographic data were calibrated against wells with known water tables. RESULTS The area surveyed was approximately 40,110 ha (Table 1). We established a total of 23 map units (Fig. 1, Table 1). Most of these landform-vegetation-soil map units represent ranges in more or less continuous variation caused by several interacting variables.Some divisions are obvious, but others are arbitrary because of the limits of our current knowledge of these environments. Plant Communities of the Playa The surface of the Grass Valley playa is usually free of vegetation (Fig. 2). Obviously the salt content of the surface soils of the playa are limiting to plant growth, but occasional prolonged periods of flooding may be the lim- Vol. 46, No. 1 GREAT BASIN NATURALIST Fig. 1. Distribution of map units for pluvial Lake Gilbert below the maximum lake level. Units based on landform, soils, and vegetation. Area designation (based on dominant plant species) and area of map units given in Table 1. - : Fig. 2. Grass Valley playas, viewed from a central location looking north. Hot Springs Point (2,300 m), Sawtooth Mountains (2,230 m), Cortez Pass (1,805 m), and Mount Tenabo (2,820 m) on skyline in background from left to right. Note typical polygonal cracking of playa surface. Range pole 1 m divided into dm. spread of rhizomes of the grass Distichlis spi- iting factor for the most extreme halophytes. cata var. stricta.’ Rhizomes, several meters in Flooding is probably the factor limiting the January 1986 YOUNG ETAL.: LAKE BASIN COMMUNITIES 3) se iy Fig. 3. Wet depressions on margin of playa with effervescence of salt crystals. Middle distance shows group of puff dunes. Holocene beach with scattered plants of Atriplex nuttalii. Range pole 1 m divided in dim. length, can be observed extending down eroded slopes to the playa surface. Very occasionally, a sparse cover of sum- mer-annual herbaceous vegetation is found on the margin of the playa. Suaeda occidentalis, Atriplex truncata, and Halogeton glomeratus form these communities. The depth and electrical conductivity of groundwater on the playa varied greatly, de- pending on the sampling location and season. In the fall, when the playa was dry, the water table in observation wells ranged from 1.5 to 1.8 m below the playa surface. Electrical con- ductivity of water from the wells ranged from 34 to 105 dS m '. In the spring, when the groundwater reached the surface of the playa, electrical conductivity of the water dropped to 3to 10dS m * Portions of the playa were surveyed and brass plaques were mounted on 3.75—cm— diameter galvanized steel pipe to mark sec- tion corners in 1917-1918. By 1979 the pipe supports were faint discolorations in the playa soil profiles. Wooden fence posts planted dur- ing the same time period now have bases swollen by salt crystallization to three times their current aerial diameters. Hummocks.—The wet depressions on the margins are free of vegetation (Fig. 3), but the margins of these depressions are partially ringed with elongated hummocks scarcely 25 cm above the playa surface. The mounds are salt encrusted, and the unwary playa hiker who steps on the mounds for footing in the slimy-slick soils of the wet depression is greeted by a puff of salts that instantly can be tasted. Despite the salinity of the mounds, they are densely vegetated with clumps of Allenrolfea occidentallis (Fig. 4). The electri- cal conductivity of extract from the surface soils of the wet depressions reached values of 400 dS m ' and averaged 34 dS m . The texture of the soils of the wet depressions consists of at least two-thirds clay-sized parti- cles, whereas the texture of the mounds is at least two-thirds silt-sized particles. These mounds may form during drying cy- cles on the playa when the crystallization of salts around the wet depressions causes miniature salt domes to form. The domes trap 6 GREAT BASIN NATURALIST Fig. 4. silts in their irregular surface topographies and eventually rise high enough above the playa surface to permit the establishment of Allenrolfea plants. The silt particles are highly permeable compared to the clays of the playa and therefore may be readily leached of solu- ble salts. Mobile Dunes.—Mobile dunes are found on the western margin of the playa, especially where ephemeral streams dump bed loads onto the playa (Fig. 2). These dunes are from 1.0 to 2.0 m in height and are egg shaped in outline (Fig. 5). The pointed end of the dunes is oriented toward the southwest. The dunes are vegetated with clumps of Sarcobatus vermiculatus, a highly variable species. Juvenile plants have a distinct fleshy leaf morphology. Plants growing on upland sites where the groundwater table is not close to the surface are small, stunted, and gray in color. This form is classified as S. vermiculatus var. baileyi, or it is sometimes raised to the species level (S. baileyi). On the mobile dunes the Sarcobatus plants are highly polymorphic. The west-southwest portions of the dunes are usually bare. During Vol. 46, No. 1 Small puff dune with Allenrolfea occidentalis. Range pole 1 m divided in dm. dry periods electrical conductivity of the bare soils may reach 250 dS m '. The tops of the dunes are covered with dense patches of dwarfed Sarcobatus stems scarcely 2 dm tall. Electrical conductivities of the soils under the dwarf shrubs may reach from 60 to 85 dS m’. The east slopes of the dunes support relatively sparse stands of Sarcobatus plants, which reach a meter or more in height. Maximum electrical conductivity of soils on the east slope seldom exceeds from 25 to 40 dS m'. The Sarcobatus plants eventually die and the largely clay textured dunes melt away from the skeletons of crowns and roots. Each dune is apparently vegetated with a single clone of Sarcobatus. Haystack Dunes.—Haystack dunes are sparingly distributed along the southeastern margin of the playa, opposite the wave cut escarpment. These dunes range to 7.5 m in height, the tallest being higher than the adja- cent lake plain. The east slopes of these large dunes have sparse stands of Sarcobatus plants. Recent Holocene Beach .—This landform is only preserved in discontinuous stretches January 1986 YOUNG ET AL.: LAKE BASIN COMMUNITIES 7 a Fig. 5. Mobile dunes vegetated with Sarcobatus vermiculatus plants. West side of dunes are largely bare with dwarf plant on top and 1-m tall plants on the east side. If the clones of Sarcobatus die, the largely clay-texutred dunes melt away (rilled dune in middle distance) and erode across the playa. Range pole 1 m divided in dm. around the margins of the playa. It apparently represents some recent rise of the lake of suffi- cient duration to erode a definite beach into the escarpment of the surrounding lake plain. The discontinuous nature of this beach pro- vides evidence of the highly erodible nature of the interface between playa and lake plain. Vegetating these fragments of Holocene beach is a relatively dense overstory of shrubs (density 0.9 plants per per m*, cover 8%) con- sisting of S. vermiculatus, Atriplex confertifo- lia, A. nuttallii, and Kochia americana. Virtu- ally, the only herbaceous species in these communities is Thelypodium flexuosum. Plant Communities of the Lake Plain Older Holocene Beach. —This is the larg- est mapping unit of the lake plain associated landforms, constituting 7% of the area of the total pluvial lake basin (Table 1). This land- form is located about 2 m higher than the recent Holocene beach that incompletely sur- rounds the current playa. The general topog- raphy is nearly flat from a prominent beach ridge located at 1,730 m elevation to the es- carpment at the playa edge (Fig. 6). On the southeast margin of the playa, stable dunes cover the older Holocene beach, and on the northeast side of the playa, clay dunes inter- rupt drainage to the playa, forming dune basins on the older Holocene beach surface. Occasional drainage channels have eroded channels into the lake plain, creating minia- ture badland topographies. The microtopog- raphy consists of coppice dunes occurring around shrubs and averaging from 20 to 30 cm in height. The coppice dunes support thin, ragged-edge microphytic crusts. The inter- space between shrubs is usually bare of vege- tation and microphytes. Soils of the lake plain largely belong to the order Inceptisols (Anonymous 1975). A typi- cal soil for the older Holocene beach would be classified as a fine, montmorillonitic (cal- careous), mesic Aeric Halaquept. A typical profile consists of a dark, grayish brown salt- and sodium-affected silty clay surface horizon, over a thick (2 m) light gray lacustrine clay 8 GREAT BASIN NATURALIST Vol. 46, No. 1 Fig. 6. Sarcobatus vermiculatus -dominated plant community located on older Holocene beach on lake plain. horizon. Below this layer iron mottles are common. Sodium absorption ratios of these soils are 45 or greater, with average electrical conductivity of the saturation extract of 8 to 12 dS m’. In the upper 30 cm, the predominant salts are sodium chloride and sodium sulfate (Roundy 1984). Salt concentrations of the sur- face soil decrease due to leaching by winter and spring precipitation and increase with the lack of precipitation in the summer (Roundy et al. 1983). The depth of the water table on the older Holocene beaches averages 3.2 m during dry periods and 2.1 m in the spring. However, the amount of variation measured was highly de- pendent on location. Apparently the nature of the watershed and alluvial fan at the mouth of the watershed influences the groundwater on the lake plain adjacent to the alluvial fan. Snowmelt in surrounding mountains causes ephemeral streams to flow on the mountain escarpment in May and early June. As the water leaves the pediment surface, it disap- pears on the relatively coarse alluvial fans. The water moving down through the fans ap- parently strikes buried, fine-textured lacus- trine deposits from previous high rises of the lake. Lateral movement of groundwater oc- curs with discharge in springs or seeps where past wave action has truncated alluvial fans. There is a delay of a month to six weeks be- tween the water disappearing at the mouths of canyons along the mountain escarpment and rising groundwater levels on the lake plain. Below some alluvial fans, even if they were deposited at the mouths of large watersheds, the groundwater level on the lake plain does not vary. Similar relationships were apparent for sol- uble salt content of the groundwater. In areas where the water table varied, electrical con- ductivity of water from the wells dropped as the water table rose and increased as the wa- ter table dropped. Where the water table re- mained stationary, electrical conductivity of the water was very constant. The plant communities of the older Holocene beach are dominated by Sarcobatus vermiculatus shrubs with varying amounts of Atriplex confertifolia (Fig. 3). Cover of both species of shrubs was only 12% (Table 2). Only six herbaceous species were recorded in the January 1986 understory (Table 3). A Cruciferae, Thely- podium flexuosum, was the most frequent herbaceous species encountered. Atriplex confertifolia tended to occur in rel- atively pure stands within a matrix of S. ver- miculatus. The A. confertifolia-dominated areas had deeper water tables. Sarcobatus vermiculatus is a pronounced phreatophyte with reported rooting to water tables at depths of 7.5 m (White 1932). However, the very low permeability of the clay soils, cou- pled with the scant atmospheric precipitation associated with the central basin locations, may never result in soil wetting to the water table to permit contact of S. vermiculatus roots with the groundwater fringe. Stable Dunes .—Floristically and physiog- nomically, the plant communities of the stable dunes are very similar to those of the older Holocene beach. The soil of the stable dunes is classified as a coarse-silty, mixed (cal- careous), mesic, Aquic Durorthidic Tor- riorthent of the order Entisol. The soils are very deep and moderately well drained be- cause of the relatively coarse texture. Typi- cally the surface layer is light gray silt. The underlying material to a depth of 2 m is light brownish gray and light gray stratified and very fine sandy loam and silt loam with discon- tinuous weak silicon concentration in the up- per part. Despite the quite different soil and a generally greater depth to the groundwater table, only the shrub Grayia spinosa was added to the plant communities of the stable dunes compared to the older Holocene beaches of the lake plain (Table 2). Clay Dunes.—With only 8% plant cover, there is little vegetation on the clay dunes (Table 2 and 3). The microdrainages and val- leys among the dunes are occupied by flows of salt crystals that are apparent after a rain. Vegetation is generally restricted to the top and east side of the dunes. Dune Basins. —The dunes, either stabi- lized or clay, interrupt drainage from the lake plain to the playa. In wet years ephemeral lakes form in the basins (Fig. 7). The typical soils of these basins are classified as fine silty, mixed (calcareous) mesic Aeric Fluvaquents. These soils range from slightly to strongly sa- line. The slightly saline soils are in the lowest depressions that are subject to spring pond- ing. The soils are very deep and poorly YOUNG ET AL.: LAKE BASIN COMMUNITIES 9 drained. Typically the surface layer is light gray, strongly salt- and sodium-affected clay loam. ; Sampling the observation wells indicated that the water table depth under this landform ranges from 0.5 to 1.0 m below the soil sur- face. The electrical conductivity of the water ranged from 0.8 to 1.0 dS m '. Repeated flooding limited shrubs in the dune basin community to occasional plants of Chrysothamnus nauseosus ssp. consimilis, but a total of 27 herbaceous species was rec- orded in the basins (Tables 2 and 3). Distichlis spicata var. stricta was the most frequent herbaceous species. Lake Plain. —The lake plain itself accounts for more than 10% of the basin below the maximum level of the pluvial lake basin. Gen- erally the lake plain is physically positioned above the offshore bar that forms the upper boundary of the older Holocene beach (1,730 m elevation) and a prominent beach that trun- cates alluvial fans at 1,740 m elevation. The elevations are from the top of one bar to the top of the other bar. Once the plunge pit behind the bottom bar is passed, the lake plain is nearly flat until the 1,740 m bar is reached. The lake plain is not distributed symmetrically in the basin (Fig. 1). Most of the lake plain is located between the south- eastern margin of the playa and the wave- truncated alluvial fans at the base of the moun- tain escarpment. On the northwest margin of the playa, the playa extends to the alluvial fans with no lake plain. The genesis of the soils of the lake plain apparently combines: (a) periods of relatively deep water flooding with deposition of fine- textured lacustrine sediments; (b) recurring periods of lake desiccation with aeolian ero- sion and deposition, coupled with alluvial fan encroachment onto the plain; and (c) recur- rent flooding with truncation of the alluvial fans and subsequent incorporation of alluvial material into the lacustrine sediment by wave and current action. Such complex soil formation factors pro- duce a range of soil morphologies depending on the volume of material, the physical and chemical nature of the inputs from alluvial fans, and the physical position in the pluvial lake basin. A typical soil for the bulk of the lake plain above the older Holocene beach 10 GREAT BASIN NATURALIST Vol. 46, No. 1 TABLE 2. Density per m* and projected cover of shrub species in plant communities (mapping unit number) of the lake plain and associated landforms. Landform, plant community, density, and cover Older Holocene beach (3) Stable dunes (4) Parma dunes (5) Dune basins (6) Sarcobatus Sarcobatus Sarcobatus Distichlis- vermiculatus vermiculatus vermiculatus Sporobolus Density Cover Density Cover Density Cover Density Cover per m” % per m? % per m? % per m? % CHENOPODIACEAE Atriplex confertifolia 0.2 2.0 Grayia spinosa wl 0.2 Sarcobatus vermiculatus 0.8 10.0 1.1 14.0 0.6 4.0 COMPOSITAE Artemisia tridentata ssp. tridentata Chrysothamnus nauseosis 0.3 0.2 ssp. consimilis Chrysothamnus albidus Totals 1.0 12.0 We 14.2 0.6 4.0 0.3 2.0 TABLE 3. Frequency (%) and total cover of herbaceous species in plant communities (mapping unit numbers) of the lake plain and associated landforms. * Older Holocene beach (3) Stable dunes (4) Parna dunes (5) Dune basins (6) Sarcobatus Sarcobatus Sarcobatus Distichlis - vermiculatus vermiculatus vermiculatus Sporebolus Frequency Cover Frequency Cover Frequency Cover Frequency Cover % % % % % % % % CAPRIFOLIACEAE -Cleonella plocusperma 1 CHENOPODIACEAE Atriplex truncata Halogeton glomeratus Monolepis nuttalliana Nitrophilia occidentalis Salsola iberica 4 COMPOSITAE Aster occidentalis . Crepis runcinata Haplopappus lanceolatus Haplopappus uniflorus Hymenoxys lemmonii Solidago spectabilis Taraxacum officinale CRUCIFERAE Capsella bursa-pastoris Descurinia sophia Lepidium perforiatum 4 2 Thelypodium sagittatum Thelypodium flexuosum 86 92 54 GRAMINAEAE Distichlis spicata var. stricta 2 4 46 Elymus cinereus Elymus triticoides Hordeum pusillum Muhlenbergia rohardsonis Puccinellia airoides Puccinellia lemmonii Sitanion hystrix Spartina gracilis Sporobolus airoides JUNCAGINACEAE Triglochin maritina PLANTAGINACEAE Plantago eripoda ROSACEAE Potentilla anserina Potentilla pectinisecta Ivesia kingii twp to Ao AARP rPHAN yA | Ou wo oN Re we le e i aor January 1986 YOUNG ETAL.: LAKE BASIN COMMUNITIES 1] Lake plain Chrysothamnus-sarcobatus/ Sarcobatus/ Sarcobatus/Atriples (9) Chrysothamnus /(10) Distichlis (11) Artemisia-Elymus (7) Distichlis (8) Puccinella Density Cover Density Cover Density Cover Density Cover Density Cover per m? % per m? % per m? % per m2 % per m2 % Hall 9.2 0.2 2.0 1.0 16.0 6 3.0 1.0 15.1 4 4.0 0.2 3.0 0.3 3.0 2 ee wei Es 0.4 3.0 1.6 21.1 1.2 19.0 ea NPA) 0.4 + oO 0.3 3.0 Lake plain Chrysothamnus-Sarcobatus-(7) Sarcobatus/ (8) Sarcobatus -(9) Chrysothamnus /(10) Distichlis (11) Artemisia/Elymus Distichlis Atriplex Puccinella Frequency Cover Frequency Cover Frequency Cover Frequency Cover Frequency Cover % % % % % T T 2 T 1 2 1 a as 2 3 T 4 1 1 1 1 3 8 85 8 77 72 2 1 2 36 48 12 16 Ie 1 2 — eel ie _ ee ereues all 16 49 11 74 12 GREAT BASIN NATURALIST Be ales Vol. 46,.No. 1 Fig. 7. Dune basin with fairly dense ground cover of Distichlis spicata var. stricta. In background stabilized dunes with Sarcobatus vermiculatus shrubs. would be classified as a mixed (calcareous), mesic family of Aquic Durorthidic Tor- riorthent. Typically the surface horizon is pale brown and sodium-aftected (sodium absorp- tion ratio >40) silt loam about 10 cm thick. The upper 50 cm of the underlying material is pale brown silt loam that is weakly silicon-con- centrated in the lower part. This horizon rests upon at least 2 m of light gray to yellow clay. The above soil is associated with a complex plant community that combines as dominant species Artemisia tridentata ssp. tridentata, Chrysothamnus nauseosus ssp. consimilis, Sarcobatus vermiculatus/Elymus cinereus (Ta- bles 2 and 3, Fig. 8). This is a plant community that should not exist because the autecology of the woody dominants is in apparent opposi- tion to each other. Artemesia tridentata ssp. tridentata is probably the most highly evolved taxon of the A. tridentata polyploid complex that characterizes the landscape above the maximum level of pluvial lakes in the Great Basin (McArthur and Plummer 1978). It is not considered to be a salt-tolerant species. Chrysothamnus nauseosus ssp. consimilis is a morphologically distinct subspecies in a complex group of root-sprouting shrubs (McArthur et al. 1978). The subspecies consimilis is the only Chrysothamnus of the nauseosus group that is found in abundance in saline/alkaline habitats (Roundy et al. 1981). Sarcobatus vermiculatus is not competitive in upland nonsaline/alkaline sit- uations, and the soluble salt residue from S. ver- miculatus litter has been demonstrated to in- crease the salinity of the surface to the point of excluding reproduction of A. tridentata (Rickard 1965). The distribution of the dominant shrubs is highly variable, with occasional patches contain- ing an equal representation of these three spe- cies. Essentially, $. vermiculatus and A. triden- tata represent saline and _ nonsaline soil extremes, respectively, with C. nauseosus ssp. consimilis being more or less able to compete in both extremes as long as the water table is rela- tively shallow. The swirling, apparently mean- ingless patterns of distribution of the three dom- inant shrub species in this environment appar- ently reflect equally complex evolution of salina- tion-desalination of the older lake plain soils. January 1986 YOUNG ET AL.: LAKE BASIN COMMUNITIES 13 Fig. 8. Most abundant plant community on the older lake plain landforms. Overstory dominants Artemisia tridentata ssp. tridentata, Chrysothamnus nauseosus ssp. consimilis, Sarcobatus vermiculatus. Understory dominant, which was the mammoth, tufted perennial grass Elymus cinereus, is now much depleted by grazing of domestic livestock. The herbaceous dominant, Elymus cinereus, has been greatly depleted by past excessive grazing of domestic livestock (Les- perance et al. 1978). Under pristine condi- tions, in particularly favorable locations for soil moisture on the lake plain, the annual biomass production of this grass probably reached 2000 kg/ha. The Sarcobatus-Atriplex- dominated plant communities of the older lake plain are similar in appearance and structure to those found on the older Holocene beach (Tables 2 and 3). These xeric communities are associated with areas with deeper water tables that do not fluctuate seasonally. On the mesic end of the spectrum, several communities exist where the groundwater reaches the soil surface at least sometime dur- ing the growing season. In Chrysothamnus albidus/Puccinella species, the perennial grasses occur on miniature pedicels. The Dis- tichlis meadows are wet to the soil surface relatively late in the season, when water per- colating through the alluvial fans comes to the surface on the lake plain. This coincides with the phenology of Distichlis for summer growth. In the high water table portion of the lake plain, the soluble salts in the soil profile are largely concentrated on the soil surface by capillary discharge of groundwater. Located on the far southern extremes of the lake plain are several islands of vegeta- tion completely dominated by Atriplex nut- tallii. Only afew plants of Kochia americana share the communities. The heavily grazed A. nuttalli plants are under 10 cm high, so the communities stand out sharply from the 1-m tall mixed shrubs of the surrounding lake plain communities. Other than the very regular lower contact boundary of the soil horizons beneath the A. nuttallii com- munities, there are no obvious soil differ- ences between the A. nuttallii islands and the surrounding vegetation. 14 GREAT BASIN NATURALIST Vol. 46, No. 1 Fig. 9. Little Hot Springs, Grass Valley, Nevada. Springs support a few square meters of saline/alkaline meadow surrounded by Sarcobatus vermiculatus plants located on a peninsula in the center of the playa. The Little Hot Springs is the only source of water for a large area. Mounds were caused by cattle trampling in mud trying to drink. Poles have been placed in caldera of hot springs to discourage cattle from entering. THERMAL SPRINGS There are three groups of thermal springs below the maximum level of pluvial Lake Gilbert (Fig. 1, map unit 12). The largest group of springs, the Walti Hot Springs, are on the east central edge of the valley near the maximum level of the lake. The flora around these hot springs has been highly disturbed by agricultural activities, but the occurrence of species such as Spartina gracilis, Juncus balticus, and the central Nevada endemic J. longistylis suggests what the vegetation com- position of the shoreline may have been around pluvial Lake Gilbert. The Little Hot Springs, located in the cen- ter of the playa, support only a few square meters of Juncus-Distichlis meadow (Fig. 9). Hot Springs Point on the west side of the playa has several thermal springs on large mounds of tufa. Runoff from these springs supports Distichlis meadows. Initially we did not take the temperature of groundwater in the wells when water samples were obtained. In the second year of sam- pling, when temperatures were taken, it was determined that on the same day the temper- ature of the surface of the water table varied by more than 20 C among wells. We consid- ered none of the wells to be located in the thermal springs areas, but slightly geothermal groundwater is widespread on the lake plain. The almost complete lack of tufa deposits in the basins of pluvial Lake Gilbert is notewor- thy considering the thick mantles of tufa de- posited in Lake Lahontan (Morrison 1969). The hot springs should have built tufa domes if they were active during the pluvial lake period (Papke 1976). Plant Communities of Barrier Bar and Lagoon In the central part of the Valley, south of the playa, extensive areas of Sarcobatus vermicu- latus communities were delineated. Some of these communities supported cryptogamic crusts in the interspaces between shrubs and stands of Kochia americana, which is highly January 1986 YOUNG ETAL.: LAKE BASIN COMMUNITIES 15 : 2 . . TABLE 4. Density per m’ and projected cover of shrub species in plant communities on the barrier bar and lagoon. Barrier beach Lagoon Sarcobatus Atriplex Sarcobatus- Allenrolfea Chrysothamnus nauseosus vermiculatus confertifooia Kochiaamericana _ occidentalis ssp. consimilis/Distichlis Total Total Total Total Total Density cover Density cover Density cover Density cover Density cover m? % m? % m? % m? % m2 % CHENOPODIACEAE Allenrolfoa occidentalis 0.4 5.2 Atriplex confertifolia 2 3.0 13 8.0 Kochia americana 1.8 2.0 Sarcobatus vermiculatus a) 11.0 8 4.3 TBD) 16.0 0.2 31 0.2 0.4 COMPOSITAE Artemisia tridentata ssp. T tridentata Chrysothamnus nauseosus T 0.8 8.4 ssp. consimilis Totals ll 14.0 PRL NB} 3.0 18.0 0.6 8.3 1.0 8.8 TABLE 5. Frequency (%) and total cover of herbaceous species on the barrier bar and lagoon landforms. Barrier beach Lagoon Sarcobatus Atriplex Sarcobatus- Allenrolfea Chrysothamnus nauseosus vermiculatus confertifolia Kochia americana _ occidentalis ssp. consimilis/Distichlis Total Total Total Total Total Frequency cover Frequency cover Frequency cover Frequency cover Frequency cover % % % % % CAPRIFOLIACEAE Cleomella plocasperma 2 T 6 T 2 T CHENOPODIACEAE Atriplex truncata 6 1 CRUCIFERAE Thelypodium flexuosum 92 2 94 2 88 3 GRAMINAEAE Distichlis spicata 6 T 4 T 100 8 100 38 var. stricta Elymus cinereus 2 2 3 8 38 preferred by domestic livestock. We at first assumed the communities represented a higher level of range condition than was previ- ously observed on similar soils on the older Holocene beach. Distance from water would have limited grazing in these central valley communities (Stewart et al. 1940). A map pro- duced from satellite data showed this area as a separate unit from the remaining lake plain (unpublished data, P. T. Tueller, Division of Range, Wildlife, and Forestry, University of Nevada, Reno). Further investigations re- vealed a large depression to the south (up watershed). This depression contained red- dish colored dunes composed of clay and salt particles. The vegetation on these dunes was almost entirely Allenrolfea occidentalis, an extremely salt-tolerant chenopod. We delineated a large rectangular island about 3.3 by 5 km stretching across the valley from the older lake plain on the east to a 2.5-km-long gravel bar on the west side of the valley. On the north and south sides of the island, escarpments of about 4.5-m height ex- isted. Callaghan Creek was incised into the island with a multiterrace cut about 4.5 m deep. Cowboy Rest Creek incised a channel along the west boundary of the island before continuing on to the playa. We theorized that the island was a result of isostatic rebound following the evaporation of the waters of plu- vial Lake Gilbert. Isostatic rebound has been noted for the deeper parts of Lake Lahontan (Mifflin and Wheat 1979). Closer examination suggested that waves driven by northerly winds on the long axis of the valley built a 16 GREAT BASIN NATURALIST Vol. 46, No. 1 Fig. 10. Chrysothamnus nauseosus ssp. consimilis/Distichlis spicata var. stricta plant communities on the south maring of the Allenrolfea dunes in the eroded lagoon. Range poles 1 m divided in dm. barrier bar across the south end of the basin. A similar bar, on a much smaller scale, is being built across the northern end of the present playa. A third barrier bar may have been built upstream on Callaghan Creek ex- tending out from the most southerly gravel bar (Fig. 1). The only inconsistency with the barrier bar hypothesis for the origin of the central island landform is the lack of a gravel veneer on the bar. The lack of gravel may have been the result of the deep water location of the bar, or the gravel veneer may be buried by subsequent subaerial deposition on the area. Whatever the origin of the landform, what we identified as the barrier bar served two functions in the evolution of soils and vegeta- tion assemblages in the central valley area. First, the island is elevated above the sur- rounding landforms, so no overland flow is received from adjacent landforms. The lack of microdrainage patterns and the well-devel- oped cryptogamic crust on the soil surface indicate a very stable surface landscape. Sec- ondly, the island structure apparently blocked drainage from Callaghan Creek to the central playa, creating a large lagoon. Since the central lake level dropped to the present level of the playa, Callaghan and Cowboy Rest creeks have breached the barrier, allowing erosion of the sediments trapped in the lagoon and the drainage pattern of the entire south end of the valley to erode toward a new base level. The sediments in the lagoon contain more sol- uble salts than the surface soils of the present playa. Some soils from the A. occidentalis field were 50% soluble salts. Microscopic examina- tion of these soils revealed aggregations of salt crystals that were worn by saltation until well rounded. The plant communities of the barrier bar and lagoon are characterized by a poverty of species (Tables 4 and 5). South of the A. occidentalis dunes in the lagoon area, extensive areas of C. nauseosus ssp. consimilis/D. spicata var. stricta plant communities occur (Fig. 10). Plant Communities of Sand Dunes Sand-textured soils are very rare in the January 1986 landscape is covered with sand dunes (Table 1). The well-stabilized dunes are located on an older lake plain surface on the west side of the valley. The sands are not salt affected and support a diverse shrub and herbaceous plant community. Besides the shrubs A. tridentata and S. vermicu- latus, the dunes support Grayia spinosa and Tetradymia comosa. Several herbaceous species such as the grasses Oryzopsis hymenoides and Sitanion hystrix, which are characteristic mem- bers of plant communities found on alluvial fans, were found on the sand dunes. Plant Communities of the Alluvial Fans Over half the area below the maximum level of pluvial Lake Gilbert is occupied by alluvial fans spreading out from the mountain escarpment. Most of this area is covered by various plant communities that are dominated by Atriplex confertifolia. In the southern end of the basin, the pluvial lake was very shallow (less than 7.5 m). The lake sediments are mixed with alluvial material. The Atriplex confertifolia plants that dominate this area are less than 0.3 m tall, with total projected crown cover of the shrub around 10% (Fig. 11). Mixed with the A. confertifolia are Sarcobatus baileyi plants. Where the alluvium has mixed with shallow lake sediments, gravel has sorted to the soil surface to form desert pavement in the interspaces among shrubs. The shrubs are grow- ing on small mounds, from 10 to 35 cm above the interspace surfaces. Several soils are found in the area dominated by A. confertifolia, most of which are Aridisols. The oldest landforms support Haplargids soils. These soils have an argillic horizon. Commonly a calcium horizon has been developed below the argillic horizon. Many of the soils on the pluvial lake sediments are Orthids. These soils com- monly have horizons of accumulations of soluble salts and carbonates. The soils do not have argillic horizons. On the west side of the valley, there is an area of A. confertifolia/Oryzopsis hymenoides in ap- parent high seral status, as a result of protection from grazing by distance from water (Table 6). The density of shrubs here is about the same as in the grazed areas, but the shrub interspaces sup- port a good stand of the perennial grass O. hy- menoides , with the perennial grass Sitanion hys- trix found under shrub canopies. YOUNG ETAL.: LAKE BASIN COMMUNITIES 117 The relationship between Atriplex and Artemisia communities on the alluvial fans is very complex. The most abundant A. conferti- folia community in the basin consists of a mo- saic of Artemisia tridentata ssp. wyomingensis in the microdrainageways, with the bulk of the intervening residual soil occupied by A. confertifolia. The alluvial fan consists of a series of fans of differing age. On the southeasterly margin of the valley, A. conferifolia extends up the allu- vial fans to the mountain escarpment without the intervening Artemisia-dominated com- munities. Along the east central margin of the basin, A. tridentata ssp. wyomingensis com- munities extend to the lake plain without in- tervening A. confertifolia communities. This distribution may be due to orographic influ- ences on precipitation, both mountains to the east that accumulate precipitation and moun- tains to the west that cast rain shadows. Where A. tridentata and A. confertifolia communities abut each other laterally along the alluvial fans, A. confertifolia appears to occupy the older alluvial fan. The plant communities of the alluvial fans dominated by A. tridentata ssp. wyomingen- sis have been described in detail by Cluffet al. (1983). The herbaceous vegetation in the un- derstory of the Artemisia community is domi- nated by the alien annual grasss Bromus tecto- rum. Apparently this weed can not tolerate the salt content of the lake plain soils. Ruderal and disturbance weeds in this environment are Salsola iberica and Halogeton glomera- tus. Plant Communities of Bars Ranging from the playa to well upon the older lake plain, there are several current bars largely composed of well-sorted gravels. Some of these are 10 km long and rise from the lake plain like railroad embankments (Fig. 12). The typical soils on these bars are Xerollic Camborthids. The most abundant plant com- munities are dominated by A. confertifolia/ Artemisia spinescens . A second form of bars are offshore bars formed between alluvial fans along the mar- gins of the valley. The soils of these bars are similar to those noted for the larger current bars. Wave plunge pits were formed behind these offshore bars. Most of the plunge pits 18 GREAT BASIN NATURALIST Vol. 46, No. 1 Fig. 11. Atriplex confertifolia-dominated communities at the south end of Grass Valley, Nevada. Range pole 1 m divided in dm. TABLE 6. Density per m? and projected cover of shrub and frequency and cover of herbaceous species in a high condition Atriplex confertifolia/Oryzopsis hymenoides community. Shrub Herbaceous Total Density Cover Frequency cover per m> % % SHRUBS Chenopodiaceae Atriplex confertifolia 8 8 Sarcobatus baileyii 59) 2 Compositae Artemisia spinescens 1 l Total 1.1 ll | HERBACEOUS Cruciferae Thelypodium flexuosum 9 Graminaeae Sitanion hystrix 27 Oryzopsis hymenoides 64 are now filled with silt-textured sediments material was reeroded to the natural basins of that are thought to be the result of wind ero- _ the plunge pits. We called the filled-in plunge sion off the playa and subsequent subaerial _ pits lagoons. i despoition on the alluvial fans (Young and The soils of the lagoons are Durorthidic Evans 1984). The fine-textured deposition Torriorthents. Some of the lagoons support | January 1986 YOUNG ET AL.: LAKE BASIN COMMUNITIES 19 Fig. 12. Atriplex confertifolia-Artemisia spinescens located on the north slope of a large gravel bar on the west side of Grass Valley, Nevada. stands of S. vermiculatus, others A. conferti- folia. Both species are out of place when the surrounding alluvial fans have A. tridentata spp. wyomingensis plant communities. The structure of the surface horizon of the lagoon soils, especially in the interspaces be- tween shrubs, greatly limits moisture pene- tration into the soil profile. In the spring, after winters with below-average precipitation, there is some moisture available for plant growth in soil profiles of Artemisia communi- ties. At the same time soils of the lagoons would be completely dry below the surface. SYNTHESIS OF DYNAMIC LANDFORMS The basin of pluvial Lake Gilbert is an elon- gated bow filled with stairsteplike terraces of predominately clay-sized particles. Atmo- spheric drought and reduced osmotic poten- tials caused by soluble salts combine to limit vegetation cover and subsequent protection from erosion. Water as a mechanism for erosion is re- stricted in the basin because of lack of precipi- tation and the porous nature of the surround- ing alluvial fans, which largely absorb runoff from the surrounding mountains and limit surface flow. The fine texture of the lake sedi- ments and the limited vegetation cover pro- mote wind erosion. In a previous study in this lake basin, we documented the erosion and subaerial deposition of fine-textured sedi- ments (Young and Evans 1985). Although we characterized the role of water erosion as restricted, it must not be over- looked or underestimated. Without periodic moisture events that produce stream flow and/or overland flow to eroded rills, gullies, and washes, the development of crypotgamic crust as now exists on the barrier bar would probably stabilize the interspaces among shrubs and protect them against wind erosion. When sufficient water is available on the lake plain to flow, small streams are faced with extremely flat gradients. In addition, the flat gradients are often interrupted by dunes or offshore bars. Streams tend to meander on the flat lake plains, with loads of sediment being deposited 20 in dune basins and plunge pits. Once the bar- riers to flow are broken by breaching a barrier bar or dune dam, large amounts of sediment are suddenly available for deposition on the next lower level of the playa itself. Two factors complicate this deposition pattern: (1) the sediments that are moving down the levels in the pluvial basins are loaded with soluble slats, and (2) the redeposited sediment often develops. vesicular crust, which limits seedling establishment. These erosion pro- cesses, which in a humid climate would have proceeded to a new base level milleniums ago, are retarded to an almost imperceptible pace by the current aridity of the basin. The erosion and deposition processes are proceeding at a microscale in virtually every location in the basin. On the other end of the scale is the 10 km of braided channels above the dune desert, the erosion of which was instigated when the barrier bar was breached. The basin of pluvial Lake Gilbert, below the maximum lake level, is composed of dy- namic landforms evolved toward a new equi- librium. The landform-soil dynamics provide a fertile template to be colonized and domi- nated by evolving plant species. LITERATURE CITED ANDERSON, D. C. 1978. Cryptogamic soil crusts: factors influencing their development in Utah deserts and their recovery from grazing on Utah winter ranges. Unpublished dissertation, Brigham Young University, Provo, Utah. 81 pp. ANONYMOUS. 1951. Soil survey manual. Soil Conservation Service, U.S. Department of Agriculture, Wash- ington, D.C. 504 pp. 1975. Soil taxonomy. U.S. Department of Agri- culture, Soil Conservation Service. Agric. Hand- book 436. U.S. Government Printing Office. Washington, D.C. 754 pp. BILLINGS, W. D.1945. The plant associations of the Carson Desert region, western Nevada. Butler Univer- sity, Botany Studies 7:89—123. ____. 1949. The shadscale vegetation zone of Nevada and eastern California in relation to climate and soils. American Midland Naturalist 42:87—109. BLACK, C. A., ED. 1965. Methods of soil analysis. Part I. Physical and mineralogical properties, including statistics of measuring and sampling. Agronomy Publ. 9. American Society of Agronomy, Madison, Wisconsin. 770 pp. Bouyoucos, G. J. 1962. Hydrometer method improved for making particle size analysis of soil. Agronomy J. 54:464—465. GREAT BASIN NATURALIST Vol. 46, No. 1 CHARLEY, J. L., AND N. E. WEsT. 1975. Plant-induced soil chemical patterns in some shrub-dominated semi- desert ecosystems in Utah. Journal Ecology 63:945—-963. CLEMENTS, F. E. 1920. Plant indicators. Carnegie Insti- tute, Washington, D.C. Publ. 353:1-388. CLuFF, G. J., J. A. YOUNG, AND R. A. Evans. 1983. Edaphic factors influencing the control of Wyoming big sagebrush and seedling establishment of crested wheatgrass. J. Range Manage. 36:786—797. CRONQUIST, A., A. H. HOLMGREN, N. H. HOLMGREN AND J. L. REVEAL. 1977. Intermountain flora. New York Botanical Garden, New York. Vols. I and VI. Evans, R. A., AND R. M. Love.1957. The step-point method of sampling—practical tool in range re- search. J. Range Manage. 10:208—212. FLOWERS, S. 1934. Vegetation of the Great Salt Lake region. Botanical Gazette 95:353—418. Gates, D. H., L. H. STODDART, AND C. W. Cook. 1956. Soil as a factor influencing plant distribution on salt deserts of Utah. Ecol. Monogr. 26:155-175. GoopMaAN, P. J. 1973. Physiological and ecotypic adapta- tions of plants to salt desert conditions in Utah. J. Ecology 61:473—494. HoucGutTon, J. G., C. M. SAKAMOTO, AND R. O. GIFFORD 1975. Nevada's weather and climate. Nevada Bu- reau of Mines and Geology Special Publ. 2. Mackay School of Mines, University of Nevada, Reno. 78 pp. Huss, C. L., AND R. R. MILLER.1945. The zoological evi- dence. Bulletin of the University of Utah. 78(20): 17-166. KEARNEY, T. H., L. J. Briccs, H. L. SHANTZ, J. W. MCLANE, AND R. L. PIEMEISEL. 1914. Indicator significance of vegeta- tion in Tooele Valley, Utah. Agric. Res. 1:365—417. LESPERANCE, A. L., J. A. YouNG, R. E. ECKERT, JR., AND R. A. Evans.1978. Great Basin Wildrye. Rangeman’s J. 5(4):125—127. MCARTHUR, E. D., D. L. Hanks, A. P. PLUMMER, AND A. C. BLAUER.1978. Contributions to the taxonomy of Chrysothamnus species using paper chromatogra- phy. J. Range Manage. 31:216-223. MCARTHUR, E. D., AND A. P. PLUMMER. 1978. Biogeography and management of native western shrubs: a case study, section Tridentatae of Artemisia. Great Basin Nat. Mem. 2:229-243. MIFFLIN, M. D., AND M. M. Wueat. 1979. Pluvial lakes and estimated pluvial climates of Nevada. Bulletin 94. Nevada Bureau of Mines and Geology, Mackay School of Mines and Geology, University of Nevada, Reno. 57 pp. MILLER, R. F., F. A. BRANSON, I. S. MCQUEEN, AND C. T. SNYDER.1982. Water relations in soils as related to plant communities in Ruby Valley, Nevada. J. of Range Manage. 35:462—468. Morrison, R. B. 1969. Lake Lahontan: Geology of southern Carson Desert, Nevada. U.S. Geological Survey Prof. Paper 401. 156 pp. Munz, P. A., AND D. D. Keck. 1968. A California flora with supplement. University of California Press, Berkeley, California. 1905 pp. PapkE, K. G. 1976. Evaporites and brines in Nevada playas. Bulletin 87. Nevada Bureau of Mines and Geology, Mackay School of Mines, University of Nevada, Reno. 39 pp. January 1986 RICKARD, W. H. 1965. The influence of greasewood on soil moisture and soil chemistry. Northwest Sci. 39:36-—42. ROUNDY, B. A., J. A. YOUNG, G. J. CLUFF, AND R. A. EVANS. 1983. Measurement of soil water on rangelands. Agric. Res. Results ARR-W31. Agricultural Re- search Service, U.S. Department of Agriculture, Oakland, California. 27 pp. Rounpy, B. A., J. A. YOUNG, AND R. A. Evans. 1981. Phe- nology of salt rabbitbrush (Chrysothamnus nau- seosus ssp. consimilis) and greasewood (Sarcoba- tus vermiculatus ). Weed Sci. 29:448—454. ROuNDY, B. A. 1984. Estimation of water potential compo- nents of saline soils of Great Basin rangelands. Soil Science Society of America Journal Vol. 48, #3, May-June 1984. ROUNDY, B. A., J. A. YOUNG, AND R. A. Evans. 1983. Sur- face soil and seedbed ecology in salt desert plant communities. In The biology of Atriplex and re- lated chenopods. U. S. Department of Agricul- ture, Forest Service, Intermountain Forest and Range Experiment Station, General Technical Re- port, Ogden, Utah. YOUNG ETAL.: LAKE BASIN COMMUNITIES 21 SHANTZ, H. L., AND R. L. PIEMEISAL. 1940. Types of vegetation in Escalante Valley, Utah, as indicators of soil condi- tions. U.S. Department of Agriculture Technical Bul- letin 713. 46 pp. STEWaRT, G., W. P. CorraM, AND S.S. Hutcuincs. 1940. Influ- ence of unrestricted grazing in northern salt desert plant associations in western Utah. J. Agric. Res. 60:289-316. Stutz, H. C. 1978. Explosive evolution of perennial Atriplex in western America. Great Basin Nat. Mem. 2:161-168. West, N. E. 1979. Survival patterns of major perennials in salt desert shrub communities of southwest Utah. Range Manage. 31:43—45. —_—. 1982. Dynamics of plant communities dominated by chenopod shrubs. International Journal of Ecology and Environmental Sciences 8:73-84. West, N. E.1983. Intermountain salt desert shrublands. Pages 375-397 in N. E. West, ed., Temperate deserts and semi-deserts. Ecosystems of the world. Elsevier Scientific Publ. Co., Amsterdam. Vol 5. Wuirte, W.N. 1932. A method of estimating groundwater sup- plies based on discharge by plants and evaporation from soil. U.S. Geological Survey, Water Supply Pa- per. 659(A):1—105. VEGETATION AND FLORA OF PINE BUTTE FEN, TETON COUNTY, MONTANA : 1 Peter Lesica ABSTRACT.—The Pine Butte Fen, situated east of the Rocky Mountains in north central Montana, is a boreal, patterned peatland occurring in a relatively dry climatic region. It is one of the southernmost mires of its kind in North America. The vegetation communities present in the fen are described, and possible causes of vegetation patterning are discussed. The Pine Butte Fen is a minerotrophic fen with 93 species of vascular plants represented in an area of approximately 450 ha. Floristic similarities between the Pine Butte Fen and 11 other peatlands in North America reported in the literature are low. Similarity of this fen to other peatlands tends to decrease with increasing distance between the sites and decreasing pH of surface water at other sites. Possible causes for these trends and the floristic uniqueness of the Pine Butte Fen are discussed. Peatlands (mires) are a common feature in the boreal zones of the earth. Bogs and fens are abundant in northern and central Alberta and Saskatchewan and often cover hundreds of square kilometers in the Hudson Bay Lowlands of Ontario and the Glacial Lake Agassiz region of northern Minnesota (Sjors 1959, Heinselman 1963, Glaser 1983). West of the Continental Di- vide in Montana, small mires are common in forested areas at low to midelevations where cli- mate is relatively moist and there has been a history of glaciation. East of the Divide mires are generally small and restricted to montane areas. There are no previously published vegetation studies of Montana peatlands. This study reports the vegetation and flora of the Pine Butte Fen, a large patterned fen in north central Montana situated at the interface of the Northern Great Plains and the east front of the main range of the Rocky Mountains. The Pine Butte Fen is part of the Pine Butte Pre- serve, a sanctuary established by The Nature Conservancy to protect the last area in the conti- nental United States where large numbers of grizzly bears, Ursus horribilis, still migrate onto the plains to feed. Inaccessible and inhospitable terrain has allowed the large wetland complex surrounding Pine Butte to remain the last low- elevation stronghold of the grizzly in the lower 48 states. This report is part of a larger study providing a classification system for and descrip- tions of the wetland and riparian vegetational communities found in this unique area (Lesica 1982). Peatlands have been studied extensively in Scandanavia (Sjors 1950, 1980), England (Pearsall and Lind 1941, Pearsall 1955), Canada (Moss 1953, Sjors 1959, Vitt et al. 1975, Slack et al. 1980), and the Great Lakes region of the United States (Heinselman 1963, 1965, 1970, Schwintzer 1978, Schwint- zer and Tomberlin 1982, Glaser et al. 1981, Glaser 1983). Good general reviews of the literature concerning peatland vegetation and ecology are provided by Gorham (1957) and Moore and Bellamy (1974). There is a large body of literature dealing with relationships between mire vegetation and water and soil chemistry (Sjors 1950, Jeglum 1971, Waugh- man 1980, Schwintzer and Tomberlin 1982, Glaser et al. 1981, and Karlin and Bliss 1984). The surface patterning typical of boreal bogs and fens has been discussed by Sjors (1961), Heinselman (1963, 1970), and Glaser (1983). STUDY AREA The Pine Butte Fen is on the west side of Pine Butte, approximately 45 km west of Cho- teau in Teton Co., Montana (47°50’'N, 32°30'W, Fig. 1). The fen covers approxi- mately 450 ha on a gentle southeast-trending slope. The Pine Butte area is underlain by glacial outwash derived from calcareous shales and limestones from the main range of the Rocky Mountains that rise abruptly 9 km to the west. Water flowing south from the | Teton River through this permeable till rises _ 'The Nature Conservancy, Big Sky Field Office, Box 258, Helena, Montana 59624. 22 January 1986 Pelelonen Fen Carr Dwarf Carr Aspen Study Site ec. Montana LESICA: PINE BUTTE FEN 23 Fig. 1. Vegetation community types in the Pine Butte Fen. Numbers refer to sample stands listed in Table 1. to the surface in the Pine Butte Fen, provid- ing a nearly constant supply of cold, nutrient- enriched water (Nimick et al. 1983). Fen veg- etation occurs on organic soils (peat) 0.5—3.0 m thick. Mean annual precipitation is esti- mated to be 430 mm and mean annual tem- perature is 6.0 C (USDA 1980). The precipita- tion/evaporation ratio along the east front of the Rockies is appreciably lowered by the presence of frequent, strong westerly winds. Upland vegetation surrounding the fen is pre- dominately mixed grass and foothills prairie dominated by grasses such as Agropyron spi- catum, Bouteloua gracilis, and Stipa comata. Localized uplifts support open forests domi- nated by Pinus flexilis.. METHODS I collected plant specimens and made obser- vations during six trips to the study area from May through August 1982. Nomenclature fol- lows Hitchcock and Cronquist (1973) for vascular plants and Crum et al. (1973) for mosses. Speci- mens were deposited in the herbarium at the University of Montana, Missoula (MONTU). 24 To characterize the vegetation, I conducted quantitative sampling during the last week in August, essentially following the techniques of Daubenmire (1959). I subjectively placed 10 transects in distinct and homogenous stands of vegetation. For each stand I laid out a 50-m baseline parallel to the slope and placed 20 plots 20 < 50 m at regular intervals along this line. I estimated canopy cover for each species in each plot by assigning it to one of seven classes: T=0-1%, 1=2-5%, 2=6-25%, 3=26-50%, 4=51-75%, 5= 76-95%, and 6=96—100%. For each transect, average cover for each species is the mean of the 20 midpoints of the assigned cover classes. I estimated average shrub height to the nearest dm. I determined pH and conduc- tance values of surface water from natural de- pressions using portable meters. I mapped vegetation using a 1:24000 infrared pho- tograph supplemented by on-site inspection. Prominence values (PV) were obtained us- ing the formula (PV=CVF) where C=% canopy cover and F=absolute frequency (Beals 1960). Sorenson's Index of Similarity (S,) was computed using the formula S,=2w/ a+b where w=number of the species com- mon to both areas, and a and b are the num- bers of species in areas A and B, respectively (Mueller-Dombois and Ellenberg 1974). RESULTS AND DISCUSSION Surface Patterning The Pine Butte Fen displays patterning similar to that of boreal mires and is among the southernmost patterned peatlands in North America, occurring in a region with a rela- tively low precipitation/evaporation ratio. The recurring pattern found throughout the Pine Butte Fen is one of parallel low ridges (strings) approximately 0.5 m high and 0.5-1.0 m wide alternating with shallow wa- ter-filled depressions (flarks) 0.5—2.0 m wide. Strings and flarks lie transverse to the slope, perpendicular to the direction of water move- ment (Fig. 2). Similar patterned mires have been described by Gorham (1957), Heinsel- man (1963, 1965), Sjors (1959, 1963), and Glaser et al. (1981). A number of theories attempt to explain the origin of these patterns. Since strings and GREAT BASIN NATURALIST Vol. 46, No. 1 flarks are generally found on a slope and are always aligned perpendicularly to it, the most plausible explanation involves gravity. Down- hill slippage of peat may result in a series of ridges separated by splits in the surface that fill with water (Pearsall 1955). These rudimen- tary strings and flarks may then be further differentiated by the relatively higher produc- tivity of the more aerobic string environment. The ponds located at the north end of the Pine Butte Fen, although not extremely elongate, are aligned across slope and may also be the result of downhill slippage (Erman 1976, Moore and Bellamy 1974). Sjors (1961, 1980) feels that string and flark patterns are not caused but merely oriented by the sloping condition. According to his theory, flarks are due to excessive waterlogging, and strings are caused by lateral pressure of ice during freeze-thaw cycles. The regeneration com- plex theory of Sernander and Von Post and modified by Kulcynski attempts to explain hummock-hollow microtopography in terms of differential growth rates of mosses, princi- pally Sphagnum (Moore and Bellamy 1974). A completely satisfactory and all-encompassing explanation for string and flark patterning has yet to be worked out. Vegetation Results of the vegetation and water chem- istry analysis are presented in Table 1. Based on differences in species composition, devel- opment of the shrub layer, and physiognomy of the habitat, the vegetation of the Pine Butte Fen may be divided into three community types (c.t.’s): open fen, dwarf-carr, and carr. The open fen c.t. is further divided into typi- cal and Scirpus phases. In some areas of the fen these community types appear distinct; elsewhere they form a continuum. Water chemistry analyses were too superti- cial for determining significant correlations; however, pH was consistently lower in the shrub-dominated carr and dwarf-carr c.t.’s. I was unable to detect any correlation between ionic concentration as measured by specific conductance and vegetation. In a study of Swedish mires, Sjors (1950) also found pH to be superior to conductivity as a predictor of vegetation type. Open fen community type .—Open fen veg- etation is dominated by graminoids and 25 PINE BUTTE FEN LESICA January 1986 inkles. Fig. 2. Aerial photograph of Pine Butte Fen. Note string-flark patterning that appears as parallel wr 26 GREAT BASIN NATURALIST Vol. 46, No. 1 TABLE Ll. Species association table for 10 stands in the Pine Butte Fen. Values indicated are prominence values (see Methods). Conductivity and pH measurements were taken from samples of standing water. Stands are grouped by community type. Open fen Open fen Scirpus phase Dwarf carr Carr Stand number 1 4 8 2 5 9 3 W 6 10 Water (%cover) 42, 31 45 38 40 54 0 0 5 45 Bare ground 0 0 8 0 0 0 0 0 (0) 0 Shrub height (dm) 4 4 4 4 4 9 13 18 12 18 Conductivity (umho/cm) 445 520 600 440 490 430 560 395 520 510 pH TW Tell Cot U2 Poul 7.1 6.7 6.8 6.6 6.9 MOSSES Drepanocladus revolvens 174 80 17 146 59 0 25 0 3 0 Scorpidium scorpioides 60 25 0 51 29 92 0 0 0 0 Campylium stellatum 178 320 145 206 204 132 158 71 207 1 Rhynchostegiella compacta 0 0 2 0 0 1 29 54 89 1 Calliergon giganteum 0 0 0 0 0 0 0 46 0 0 GRAMINOIDS Carex livida ily/ 9 19) 5 6 15 0 0 0 0 Carex limosa 1 52 0 34 113 7 0 0 0 0 Eleocharis pauciflora 2 53 25 68 13 1 3 0 0 0 Muhlenbergia glomerata 25 13 if 3 5 15 0 0 1 0 Carex simulata 250 250 210 38 76 27 0) 2 233 0 Scirpus acutus 0 0 0 268 268 256 0 0 0 0 Carex buxbaumii 0 1 0 0 21 0 0 0 1 0 Carex aquatilis 63 1 40 22 0 1 0 165 0 0 Juncus balticus 10 102 46 1 103 37 239 342 197 0 Carex rostrata/aquatilis — i —— — 1 — 170 — — 311 Carex lasiocarpa 0 0 0 0 0 0 32 17 0 0 FORBS Menyanthes trifoliata 279 29 0 136 1 0 0 0 0 0 Utricularia vulgaris 0 0 67 9 11 65 MV . @ 0 0 Utricularia minor 49 4 1 7 2 ll 0 0 0 0 Aster junciformis 7 4 4 18 ae 1 2 0 0 0 Galium boreale 42 1 0 64 DT 1 23 6 15 0 Triglochin maritima 20 155 151 10 1 Ci 9 6 23 0 Equisetum laevigatum 0 0 0 0 0 0 1 38 1 47 Equisetum arvense 0 0 0 0 0 0 0 32 0 0 SHRUBS Potentilla fruticosa 65 18 21 117 14 20 118 1 22 0 Betula glandulosa 26 26 1 26 33 34 137 4] 76 190 Cornus stolonifera 0 0 0 0 0 0 0 18 24 12 Salix candida 2 1 2 2} 2 1 1 1 74 2 Salix phylicifolia 0 0 0 0 0 0 1 ll 1 75 bryophytes. Shrubs are less than 1.0 m tall with cover values rarely exceeding 30%. This type is associated with poorly drained fibrous peat of the Dougcliff Series (USDA 1980). The surface of the peat displays string and flark patterning, with the flarks containing stand- ing water during all or most of the growing season. In many areas the peat is so unconsoli- ated and water saturated that the entire sur- face seems to be floating and “quakes” when stepped on. The strings are dominated by Carex simu- lata, C. aquatilis, Juncus balticus, Muhlen- bergia glomeata, Betula glandulosa, and Po- tentilla fruticosa. Common forbs are Triglochin maritima, Galium boreale, Aster junciformis, Viola nephrophylla, and Dode- catheon pulcherrimum. The mosses, Camp- ylium stellatum and Drepanocladus revol- vens, form an almost continuous ground layer. Vegetation of the flarks is dominated by the aquatic dicots Utricularia vulgaris, U. mi- January 1986 nor, and Menyanthes trifoliata and the mosses Scorpidium scorpioides and Drepan- ocladus revolvens. Carex simulata, C. livida, and Eleocharis pauciflora are common graminoids. Throughout the open fen are 1-5 ha patches of vegetation dominated by the bul- rush Scirpus acutus. Scirpus is abundant on both strings and flarks, partially replacing Carex simulata. Potentilla fruticosa and Be- tula glandulosa have greater cover and often attain greater height in these patches than in typical open fen vegetation. All the dominant species of typical open fen vegetation are also associated with the Scirpus. Since the small patches of Scirpus-dominated vegetation can- not easily be mapped from aerial photo- graphs, this vegetation is best referred to as the Scirpus phase of the open fen c.t. Carr community type. —This community type is dominated by shrubs ranging in height from 1.0 to 3.0 m and attains total cover of greater than 50%. It is associated with mucky peat of the Winginaw Series (USDA 1980). The surface is often of ahummock-hollow mi- crotopography, but distinct strings and flarks do not occur. Standing water, as much as 0.5 m deep, is often present throughout the growing season. In the Pine Butte Fen, carr vegetation occurs along the margins and occasionally on isolated areas of higher ground. The hummocks are dominated by Betula glandulosa, Salix monticola, S. phylicifolia var. planifolia, S. serrissima, and Cornus stolonifera. Depressions around the shrubs are dominated by the coarse sedges Carex rostrata and C. aquatilis and by Equisetum laevigatum. Forbs are uncommon, and mosses are present only at the base of shrubs. Dwarf-carr community type. —Dwarf-carr vegetation is intermediate in appearance and composition to the carr and open fen c.t.’s, and, in some instances, a continuum of all three types occurs. Shrub development is no- ticeably greater than in the open fen but less than in the carr. Shrubs commonly attain heights of 0.5—2.0 m and cover of greater than 30%. Hummock-hollow microtopography is present, but distinct strings and flarks are generally not apparent. Standing water is present early in the growing season. The dwarf-carr c.t. is most common along the LESICA: PINE BUTTE FEN ik edges of the Pine Butte Fen, but it is also found around ponds and in patches through- out the open fen c.t. Dominant shrubs are Betula glandulosa, Cornus stolonifera, Salix candida, and Poten- tilla fruticosa. Important herbaceous species are Carex aquatilis, C. simulata, C. lasio- carpa, Juncus balticus, Triglochin maritima, Equisetum laevigatum, E. arvense, and Gal- ium boreale. Shrubs and most forbs occur only on hummocks, whereas graminoids and Equi- setum spp. occupy both hummocks and hol- lows. The masses Campylium stellatum and Rhyncostegiella compacta are abundant on hummocks beneath litter. Narrow bands of vegetation dominated by Carex rostrata, C. sartwellii, C. lanuginosa, and C. aquatilis occur occasionally through- out the fen. These bands of coarse sedges run parallel to the slope, and surface water move- ment is often apparent. Heinselman (1963) refers to similar communities as water tracks. I did not sample this vegetation as it occupies an insignificant portion of the study area. Aspen (Populus tremuloides) dominated vegetation occurs on islands of mineral soil along the east side of the Pine Butte Fen (Fig. 1). These communities were not considered in this study. Vegetation patterns in the Pine Butte Fen are complex (Fig. 1). Throughout the fen, wa- ter percolates up through the underlying min- eral substrate (Nimick et al. 1983). Percola- tion may not be uniform through space. Water appears stagnant in most flarks, but water flow is apparent in small drainages along the mar- gins as well as around ponds and in water tracks. Different rates of water flow may be responsible for much of the vegetation pat- terning. Jeglum (1974) states that the two most im- portant environmental gradients affecting the vegetation, floristics, and productivity of peatlands are the moisture-aeration and pH- nutrient regimes (see also Sjors 1950, Gorham 1957, and Heinselman 1970). In saturated or- ganic soils, oxygen is often a limiting factor. Even in mires fed by nutrient-enriched wa- ter, phosphorus and nitrogen may be limiting (Slack et al. 1980, Schwintzer and Tomberlin 1982). Areas with increased water flow will have access to greater amounts of oxygen and minerals. It has also been suggested that re- 98 GREAT BASIN NATURALIST ducing conditions in peat may promote the accumulation of toxic compounds around sub- terranean plant organs, a situation that is am- meliorated by increased water flow (Moore and Bellamy 1974). Heinselman (1963) found that better tree growth in Minnesota peat- lands was correlated with increased water movement, and Ingram (1967) believed that in some cases water movement alone may be responsible for the presence of more eu- trophic vegetation. The open fen community type had the greatest number of species common to bogs and fens throughout boreal North America. Plants such as Utricularia spp., Menyanthes trifoliata, Muhlenbergia glomerata, Carex livida, C. limosa, and Eriophorum spp. are adapted to the poor aeration and nutrient regimes of waterlogged soils (Sjors 1961, Heinselman 1970). Areas occupied by open fen probably have little water movement. The presence of the Scirpus phase may indicate somewhat greater water movement, perhaps resulting from increased subsurface up- welling. Lewis and Dowding (1926) and Slack et al. (1980) report similar Scirpus communi- ties in Alberta fens occurring along drainage ways, ponds, and other areas of increased wa- ter flow. Carr vegetation is probably associated with soils having relatively better oxygen and nu- trient relations. Water movement is often vis- ible in the carr c.t., and beaver activity, gen- erally associated with moving water, is greatest in this vegetation. Greater move- ment of the water along the margin of the fen is expected in light of the low hydric conduc- tivity of peat (Boelter and Verry 1977). The peat mass acts as a dam, and water, unable to pass over or through it quickly enough, flows around the margin. Moore and Bellamy (1974) attribute the existence of carr vegetation along the margin of a mire in England to the presence of drainage water circumscribing the main peat mass. In a study of peatlands in central Saskatchewan, Jeglum (1971) corre- lated cover attained by individual species with depth to water table. Water tables ranged from 80 cm below to 60 cm above the soil surface. Betula glandulosa, Cornus stolon- ifera, and five of seven species of Salix at- tained maximum cover values at depth to wa- ter table exceeding 20 cm, indicating a Vol. 46, No. 1 preference for the increased aeration of the better drained soils. In the Pine Butte Fen, Betula, Cornus, and Salix attained maximum height and cover values in the carr c.t., proba- bly responding to the more favorable condi- tions provided by greater water movement or better aerated soil. The dwarf-carr c.t., inter- mediate in shrub development, is probably also intermediate in its requirements for nu- trients and oxygen in the rooting zone. An alternative explanation for the distribu- tion of vegetation in the Pine Butte Fen fol- lows the lines of classic hydrarch succession in which the direction of change is from an aquatic to a terrestrial environment (Oosting 1948). Aquatic plants are replaced by herba- ceous emergents, which are in turn replaced by woody plants that increase transpiration and lower the water table (Dansereau and Segadas Vianna 1952). Since the peripheral carr vegetation frequently has deeper surface water than the center of the mire, a simple explanation based on hydrarch succession seems untenable. Although the major vegeta- tion patterns displayed in the Pine Butte Fen can be accounted for by a theory based on the movement of water, confirmation of this the- ory requires thorough analysis of water chem- istry throughout the fen. Floristics I observed 93 species of vascular plants and nine species of mosses in the Pine Butte Fen (Appendix A). I believe the vascular plant list to be nearly complete, whereas the list of bryophytes is certainly incomplete and proba- bly includes only the most common moss spe- cies. The majority of vascular species oc- curred on hummocks, often in association with shrubs. Many, such as Fragaria virgini- ana, Galium boreale, Senecio pauperculus, Smilacina stellata, and Vicia americana, al- though common in the fen, were at least as common in adjacent upland communities. A much smaller number of species was found in the standing water of depressions. Moore and Bellamy (1974) state that low oxygen tensions associated with waterlogged soils is one of the major problems faced by peatland vegetation. Possibly the better aeration of the hummock soils provides a habitat suitable to many spe- cies not specifically adapted to mires. Eleven species of vascular plants found in the Pine January 1986 LESICA: PINE BUTTE FEN 29 TABLE 2. Floristic similarities between the Pine Butte Fen and other peatlands in North America. S, = Sorenson’s Index of Similarity (see Methods for explanation). Ericads = number of woody ericaceous species present. Location and reference Se Distance pH Tae (km) Slack et al. (1980) 28 650 6.8-7.9 4 Alberta. Rich fens. Jeglum (1971) 27 850 6.0-—7.9 4 Saskatchewan. Eutrophic peatlands. Vitt et al. (1975) 8 900 5.0 8 Alberta. Poor fens. Baker (1972) California. a 1350 5.2—7.0 1 Fen. Heinselman (1963) 18 1400 5.3-6.4 3 Minnesota. Patterned fens. Glaser et al. (1981) 17 1400 4.3-6.9 6 Minnesota. Rich fens. Sjors (1963) Ontario. 24 2000 5.8-7.1 5 Rich fens. Schwintzer (1978) Michigan. Fens. 10 2250 5.7-7.0 3 Vitt and Slack (1975) 9 2200 4.9-6.7 11 Michigan. Bogs and fens excluding forest. Montgomery and 1 3150 — 10 Fairbrothers (1963) New Jersey. Bogs. Drury (1956) Alaska. 19 4000 — 8 Bogs. Butte Fen are restricted to bogs and fens throughout their range (Appendix A). A major environmental factor influencing mire vegeta- tion is the mineral concentration and coinci- dent hydrogen ion concentration (pH) of the water (Waughman 1980). A gradient of om- brotrophic (mineral-poor, low pH) to minero- trophic (mineral-enriched, high pH) water determines the continuum of bog to rich fen (Sjors 1950). Ombrotrophic bogs are charac- terized by “acid-loving” species-poor floras, whereas minerotrophic rich fens typically have a greater number of vascular species, some of which are calciphiles (Gorham 1957). Karlin and Bliss (1984) feel that the greater species diversity of strongly minerotrophic mires is a reflection of the complex interac- tions of microrelief and substrate chemistry gradients. The Duhr Fen displays a high spe- cies diversity common to rich fens, and many of the common mosses, such as Scorpidium scorpioides, Drepanocladus revolvens, and Campylium stellatum, are considered rich fen indicators (Sjors 1950, Slack et al. 1980). Many bogs and fens throughout North America have been studied floristically. Al- though vascular plant species lists reported for other mire systems may be incomplete, comparisons using Sorenson's Index of Simi- larity (S,) indicate degree of relationship and point to factors controlling floristic differences (Table 2). In general, floristic similarity to the Pine Butte Fen decreases with decreasing pH and increasing distance of separation. All other mire systems reported in the literature that I surveyed have at least one species of ericaceous shrub, and, in those cases where bryophytes were also included, all have at least one species of Sphagnum. Rich fens in Alberta and Saskatchewan show the greatest similarity to the Pine Butte Fen; however, none of the bogs and fens surveyed have a similarity to the Pine Butte Fen greater than 30%. In contrast, Wheeler et al. (1982) found much greater similarity between their study area in northern Minnesota and peatlands in Ontario, Saskatchewan, and Michigan (S, > 40%). A major factor responsible for the floristic uniqueness of the Pine Butte Fen is climate. Bogs and fens generally occur in regions with acool, moist climate (Dansereau and Segadas- 30 GREAT BASIN NATURALIST Vianna 1952, Sjors 1961). Walther and Leith (1960) have prepared world climatic zone maps based, in part, on length of growing season and precipitation/evaporation ratios. These maps place the Pine Butte Fen in the arid continental zone, whereas all other pat- terned fens of significant size reported in the literature for North America occur in more mesic zones (e.g., typical temperate, warm temperate, boreal, and arctic). The aridity of the Pine Butte area may account for the dis- tinctness of its peatland flora in two ways. First, Sphagnum spp. are important compo- nents of most northern mire vegetation, and, although Sphagnum can establish in cal- careous regions (Gorham 1957, Heinselman 1963), it grows only in moist areas (Moss 1953). The absence of Sphagnum and the lo- calized acidity associated with it may be re- sponsible for the absence of ericaceous shrubs, Drosera spp., and other “acid-loving” species from the Pine Butte Fen. Secondly, a large number of species found in the Pine Butte Fen are derived from adjacent upland communities and are adapted to the regional aridity. Many of these species are not avail- able to mire systems occurring in different climatic regions. The presence of nutrient-en- riched groundwater, the microtopographic heterogeneity due to the various surface pat- terns, the large size of the mire, and the rela- tively xeric climate of the region are undoubt- edly all factors contributing to the large and unique flora of the Pine Butte Fen. ACKNOWLEDGMENTS I am grateful to Gerald Moore and David McAllister for assistance in the field. Robert Dorn identified the willows, and Bruce Mc- Cune determined a number of the mosses. Anne Bradley and Robert Antibus gave help- ful criticisms on the manuscript. Joanne An- tibus provided the illustrations, and Susan Molina typed the manuscript. Funding was provided by The Nature Conservancy. APPENDIX A List of vascular plants and bryophytes of the Pine Butte Fen. The * indicates species that are restricted to bogs and fens throughout their range. Vol. 46, No.1 APIACEAE Cicuta douglasii (DC.) Coult. & Rose ASTERACEAE Antennaria pulcherrima (Hook.) Greene Antennaria microphylla Rydb. Aster junciformis Rydb. Aster occidentalis (Nutt.) Torr. & Gray Crepis runcinata (James) Torr. & Gray ssp. hispidulosa (Howell) Babc. & Stebb. Helianthus nuttallii Torr. & Gray Senecio pauperculus Michx. Solidago canadensis L. var. salebrosa (Piper) Jones Solidago nemoralis Ait. var. longipetiolata (Mack. & Bush) Palmer & Steyerm. Taraxacum laevigatum (Willd.) DC. Taraxacum officinale Weber BETULACEAE *Betula glandulosa Michx. var. glandulosa CAMPANULACEAE Campanula rotundifolia L. Lobelia kalmii L. CORNACEAE Cornus stolonifera Michx. var. stolonifera CYPERACEAE Carex atherodes Spreng. Carex aquatilis Wahl. Carex aurea Nutt. Carex buxbaumii Wahl. Carex capillaris L. Carex diandra Schrank Carex dioica L. var. gynocrates (Wormsk.) Ostenf. Carex disperma Dewey Carex interior Bailey Carex lanuginosa Michx. Carex lasiocarpa Ehrh. var. americana Fern. *Carex limosa L. *Carex livida (Wahl.) Willd. Carex nebrascensis Dewey Carex oederi Retz. var. viridula (Michx.) Kuek. Carex rostrata Stokes Carex sartwellii Dewey Carex scirpoidea Michx. var. scirpoidea Carex simulata Mack. Eleocharis palustris (L.) R. &S. Eleocharis pauciflora (Lightf.) Link *Eriophorum polystachion L. *Eriophorum viridicarinatum (Engelm.) Fern. Scirpus acutus Muhl. January 1986 EQUISETACEAE Equisetum laevigatum A. Br. Equisetum variegatum Schleich. ERICACEAE Pyrola asarifolia Michx. var. asarifolia FABACEAE Vicia americana Mubhl. var. truncata (Nutt.) Brew. GENTIANACEAE Gentiana amarella L. Gentiana dentosa Rottb. IRIDACEAE Iris missouriensis Nutt. JUNCACEAE Juncus alpinus Vill. Juncus balticus Willd. var. montanus Engelm. JUNCAGINACEAE Triglochin maritimum L. Triglochin palustre L. LENTIBULARIACEAE *Utricularia minor L. Utricularia vulgaris L. LILIACEAE Allium schoenoprasum L. Lilium philadelphicum L. Smilacina stellata (L.) Desf. MENYANTHACEAE *Menyanthes trifoliata L. ONAGRACEAE Epilobium palustre L. ORCHIDACEAE Corallorhiza trifida Chat. Cypripedium calceolus L. var. parviflorum (Salisb.) Fern. Habenaria hyperborea (L.) R.Br. Spiranthes romanzoffiana Cham. var. romanzoffiana POACEAE Agropyron caninum (L.) Beauv. var. unilaterale (Vasey) Hitche. Agrostis alba L. var. alba Bromus ciliatus L. Calamagrostis inexpansa Gray var. inexpansa Deschampsia cespitosa (L.) Beauv. var. cespitosa *Muhlenbergia glomerata (Willd.) Trin. Muhlenbergia richardsonis (Trin.) Rydb. POLEMONIACEAE Phlox kelseyi Britt. var. kelseyi LESICA: PINE BUTTE FEN PRIMULACEAE Dodecatheon pulchellum (Raf.) Merrill var. pulchellum Lysimachia thrysiflora L. RANUNCULACEAE Anemone parviflora Michx. Thalictrum venulosum Trel. ROSACEAE Fragaria virginiana Duchesne var. glauca Wats. Potentilla fruticosa L. Potentilla gracilis Doug]. var. elmeri (Rydb.) Jeps. RUBIACEAE Galium boreale L. SALICACEAE Salix bebbiana Sarg. var. perrostrata (Rydb.) Schneid. * Salix candida Fluegge Salix drummondiana Barratt Salix monticola Bebb. Salix myrtifolia Anderss. Salix phylicifolia L. var. planifolia (Pursh) Hiit. Salix rigida Muhl. var. watsonii (Bebb.) Cronq. *Salix serrissima (Bailey) Fern. SAXIFRAGACEAE Parnassia palustris L. Scrophulariaceae Castilleja miniata Dougl. var. miniata Castilleja sulphurea Rydb. Pedicularis groenlandica Retz. VALERIANACEAE: Valeriana edulis Nutt. var. edulis VIOLACEAE Viola adunca Sm. var. adunca Viola nephrophylla Greene var. cognata (Greene) Hitche. MOSSES Bryum pallescens Schleich. Calliergon giganteum (Schimp.) Kindb. Campylium stellatum (Hedw. ) C.Jens. Drepanocladus exannulatus (S.S.G.) Warnst. Drepanocladus revolvens (Sw.) Warnst. Mnium rugicum Laur. Platydictya jungermanioides (Brid.) Crum Rychostegiella compacta (C. Muell.) Loeske Scorpidium scorpioides (Hedw.) Limpr. 32 GREAT BASIN NATURALIST LITERATURE CITED BAKER, H. G. 1972. A fen on the northern California coast. Madrono 21:405-—416. BEALS, E. 1960. Forest bird communities of the Apostle Islands. Wilson Bull. 72:156—181. 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WALTER, H., AND H. LEITH. 1960. Klimadiagramm weltat- las. Fischer, Japan. WaAUGHMAN, G. J. 1980. Chemical aspects of the ecology of some German peatlands. J. Ecol. 68:1025- 1046. WHEELER, G. A., P. H. GLASER, E. GORHAM, C. M. WET- MORE, F. D. BOWERS, AND J. A. JANSSENS. 1983. Contributions to the flora of the Red Lake Peat- land, northern Minnesota, with special attention to Carex. Amer. Midl. Nat. 110:62—96. Ce EFFECTS OF SUSPENDED SEDIMENT ON LEAF PROCESSING BY HESPEROPHYLAX OCCIDENTALIS (TRICHOPTERA: LIMNEPHILIDAE) AND PTERONARCYS CALIFORNICA (PLECOPTERA: PTERONARCIDAE)! C. Evan Hornig’ and Merlyn A. Brusven” ABSTRACT. —The effects of suspended sediments on stream invertebrate detrital processing were investigated under replicated conditions in light and temperature-controlled chambers in the laboratory. The leaf-shredding insects Pteronarcys californica and Hesperophylax occidentalis were studied. Mean daily ingestion rates were lower among insects subjected to suspended sediments (1.5 and 3.0 g/l) than insects held in suspended sediment-free environments for seven of the eight trials. In five of the eight trials, mean ingestion rates were suppressed by >41% when compared to insects held in suspended sediment-free environments. Feeding inhibition was typically greater at the end of the feeding trials (14 days) than at the beginning (0-4 days). The effects of suspended sediments on ingestion were apparently related to the feeding status of the insects at the time of a trial. Insects in an active feeding mode were less influenced by suspended sediment than those in an inactive feeding mode. We conclude that, depending on the season and the duration of impact, suspended sediment can suppress processing of coarse particulate organic matter and thus adversely influence important nutrient and energy pathways in low-order streams. Inorganic sediment introduction is a com- mon and ubiquitous cause of water quality deterioration. Certain practices used in agri- culture, forestry, road construction, mining, and urban development may contribute to increased sediment loads of streams. The ef- fects of bottom sedimentation on macroinver- tebrate distribution and community structure have been well documented (Brusven and Prather 1974, Cordone and Kelley 1961, Cummins and Lauff 1969, Lemly 1982). Bjornn et al. (1977) found that different quan- tities of sediment and the degree to which cobbles were imbedded by fine sediments dif- ferentially affected species within the macroinvertebrate community. Sedimentation can also affect the size of insect populations through degradation of food resources. Reice (1980) reported that leaf litter decomposition was less in silt than on coarse-particle sediments; Herbst (1980) re- ported decreased insect consumption of pre- viously buried leaves. The effects of suspended sediments on stream insects are poorly understood. Field investigations have shown that high sus- pended sediment loads cause increased insect drift (Rosenberg and Wiens 1975, White and Gammon 1977, O'Hop and Wallace 1983). Although most suspended sediments are ap- parently not acutely toxic to aquatic life (Brusven and Hornig 1984, Oxberry et al. 1979), stressful responses, such as catastro- phic drift, have been documented and prompted us to study other sublethal effects. Our study was designed to investigate the effects of suspended inorganic sediment on the degradation of leaf litter by stream insects. MATERIALS AND METHODS Suspended sediment experiments were con- ducted in 10 1-liter glass beakers filled with 0.9 liter of unchlorinated tap water. These vessels were placed in a temperature bath held at5C + 1 to minimize algal growth. A magnetic stirrer was positioned in the bottom of each vessel to main- tain water circulation and oxygen saturation. A 1.2-mm mesh screen was placed 3 cm above the magnetic stirrer in each beaker to protect the insects from injury. Photoperiod was controlled by timers attached to four 1.2-m fluorescent- light tubes suspended 0.6 m above the vessels. For all but one experiment (trial), the dark-light conditions were set at 12-12 h. A 6-18 h dark- light cycle was used to induce feeding by the experimental animals in December. lResearch supported in part by the United States Department of the Interior, Office of Water Research and Technology Agreement 371402. Approved by the director of the Idaho Agricultural Experiment Station as Research Paper 8578. Department of Plant, Soil, and Entomological Sciences, University of Idaho, Moscow, Idaho 83843. 33 34 GREAT BASIN NATURALIST Vol. 46, No. 1 TABLE 1. Mean daily ingestion rates for Hesperophylax occidentalis larvae exposed to two levels of suspended sediment (1.5 and 3.0 g/l) and those for control larvae. (Trial duration: 14 days; individuals per treatment: 5; t and p values calculated using Student's paired t test.) Age Sediment Mean daily ingestion rate (mg/day) class concentration Control Test level insects insects t p Early instar 1.5 g/l 0.42 0.21 2.06 0.11 Early instar 3.0 g/l 1.22 1.19 0.17 0.87 Late instar 1.5 g/l 1.47 0.86 1.38 0.24 Late instar 3.0 g/l 1.81 0.85 2.10 0.10 TABLE 2. Mean daily ingestion rates for Pteronarcys californica naiads exposed to two levels of suspended sediment (1.5 and 3.0 g/l) and those for control naiads (Trial duration: 14 days; individuals per treatment: 5; t and p values calculated using Student’s paired t test). Age Sediment Mean daily ingestion rate (mg/day) class concentration Control Test level insects insects t p First-year 1.5 g/l 1.29 1.13 0.93 0.40 First-year 3.0 g/l 0.57 0.24 2.43 0.07 Second-year 1.5 g/l 1.70 0.76 2.09 0.10 Second-year 3.0 g/l 9.49 9.97 0.93 0.40 The stream insects tested were the cad- disfly Hesperophylax occidentalis (Banks) and the stonefly Pteronarcys californica Newport. These insects are primarily leaf-shredders (Merritt and Cummins 1978). Hesperophylax occidentalis is found in small mountain streams of the western United States (Martin- son and Ward 1982), whereas P. californica is common in larger streams of the region (Elder and Gaufin 1964). Early- and late-instar larvae of H. occidentalis and first- and second-year naiads of P. californica were used to assess age-specific responses to suspended sedi- ments. Insects were collected in the field and acclimated to laboratory conditions for a mini- mum of four days prior to testing. All organ- isms were starved for two days prior to test- ing. Alder (Alnus rubra) leaves were used as the food in all trials. The leaves were conditioned for one month in unchlorinated tap water held at 4 C; in winter feeding trials the leaves were held for two to four months. The suspended sediment material was commercially graded white sand sieved through a 75-~m mesh screen and added to the test chambers as a slurry. Sediments were maintained in suspen- sion by water circulation from the magnetic stirrers. Two suspended sediment concentrations (1.5 g/l and 3.0 g/l) were tested against a con- trol concentration of 0.0 g/l on each of the two nymphal or larval age classes and two insect species for a total of eight trials. For each trial, 12 vessels were arranged in pairs in a continu- ously circulating, _temperature-controlled bath. Five pairs of vessels were used to deter- mine insect feeding rates in suspended sedi- ment and sediment-free environments; the sixth pair of vessels was used to hold and condition leaf disks in suspended and nonsus- pended sediment conditions similar to the other five pairs, but in the absence of insects. Disks placed in the sixth pair of vessels were used as “blanks” and provided a correction factor caused by leaf leaching or decomposi- tion during a trial. One vessel (test vessel) from each pair was randomly selected for sedi- ment introduction, but the other vessel (con- trol vessel) remained free of sediment. This design avoided the segregation of test and control replicates that may occur with a com- pletely randomized design (Hurlbert 1984). Each trial was conducted for 14 days, with one insect placed in each of the 10 experimental vessels for the duration of a trial. Five leaves were used to initially supply leaf disks to the five pairs of test and control vessels, with one leaf per vessel pair dis- | tributed as follows. Two pairs of leaf disks — (each disk 18 mm in diameter) were cut from | the leaf with a cork borer. One disk from the | January 1986 first pair was placed in the test vessel and the pair's other disk in an insect-free container. The second pair of disks was similarly dis- tributed to the matching control vessel and an insect-free container. Leaf consumption was estimated as the difference in ash-free dry weight (AFDW) between the disk of each pair placed in the insect-free container and the corresponding disk placed in a test or control vessel. This procedure allowed direct mea- surement of leaf loss due to insect feeding and is similar to that described by Grafius and Anderson (1979). Depending on leaf utiliza- tion, the residual coarse leaf material was re- placed with new leaf disks at two- to seven-day intervals. RESULTS Mean daily ingestion rates were less among test than control insects for seven of the eight trials, but were not significantly different at p=0.05 (Tables 1 and 2). In five of the eight trials, mean ingestion rates were substantially suppressed (41% to 58%) when compared to the corresponding controls. When averaged over the trial duration, the mean ingestion rate appeared to be unrelated to species, age class, or the two suspended sediment concen- trations tested (1.5 and 3.0 g/l). However, the length of time insects were exposed to sus- pended sediment apparently influenced their feeding rates. Feeding inhibition among test insects was more evident at the end rather than the begin- ning of each trial (Figs. 1-4). The debilitating effect of prolonged exposure to suspended sediment was particularly pronounced among the early-instar H. occidentalis larvae exposed to 3 g/l suspended sediment. Whereas the mean ingestion rate of test and control insects was similar during the first 12 days of exposure to suspended sediment, the leaf consumption rate of test insects was only 12% of the control insect consumption rate during the final 2 days of the trial. DISCUSSION Our results indicate that inert suspended solids, although not acutely toxic at high lev- els, may cause sublethal effects on aquatic invertebrates by reducing feeding activity. HORNIG, BRUSVEN: AQUATIC ECOLOGY 35 Impacts on the biological functioning of large- particle detritivores (shredders) such as the species tested in this study may have serious ramifications on other biotic components of the ecosystem. In small, first-to-third order streams riparian vegetation often shades the stream, suppresses algal growth, and supplies the stream with coarse particulate organic matter. These ecosystems are highly depen- dent on allocthonous material as a primary source of energy (Anderson and Sedell 1979). Processing of large-particle detritus by shred- der invertebrates provides fine-particle detri- tivores energy and nutrients from fecal pro- duction. For example, Short and Maslin (1977) found that in laboratory streams sup- plied with alder leaves, Pteronarcys califor- nica increased the food availability to Hy- dropsyche by 35%—100% and to Simulium by 600%-—700%. Grafius and Anderson (1979) found that although the production of the leaf shredder Lepidostoma quercina was itself a minor component in a small Oregon stream, the feces produced by this insect supported 20%—50% of the more abundant simuliid pop- ulation found in the creek. This is not surpris- ing, because the fecal production rate of Lepi- dostoma has been calculated to be 50 times its growth rate (Grafius and Anderson 1980). We believe the differences we observed in the feeding rates of our control insects may be related to a physio-ecological response keyed to the season when the specimens were col- lected in the field. We were unable to induce noticeable feeding activity in the December- collected, first-year P. californica naiads until the dark-light cycle was altered to 6-18 h. Feeding rates for these insects were still much lower than first-year P. californica naiads col- lected in November. Among the second-year P. californica naiads tested, those collected in October had higher feeding rates than those collected during August. Water temperature and food availability should theoretically make autumn the most opportune time for feeding by leaf-shredding insects. The early-instar H. occidentalis larvae col- lected in mid-February had higher ingestion rates than those collected during early Janu- ary. Field observations at the collection site suggest that greatest larval growth occurs dur- ing early spring. Rapid growth is likely in- duced by seasonal cues such as photoperiod 36 3 = A sc a = = Su = = = oie! rs i) = 4 7 1 14 = 2 = > B (=) =< ciel i= = = ier = 4 7 11 14 Days Fig. 1. Mean daily ingestion rates (mg AFDW/day) for first-year naiads of Pteronarcys californica exposed to: A, 1.5 g/l. B, 3.0 g/l suspended sediment for 14 days. C = control insects, 0.0 g/l suspended sediment; T = test insects. Vertical lines = 95% confidence interval for mean ingestion rate. A Ingestion(mg AFDW/Day) B 2 + : | ch te 2 7 12 Days Fig. 3. Mean daily ingestion rates (mg AFDW/day) for early-instar larvae of Hesperophylax occidentalis exposed to: A, 1.5 g/l. B, 3.0 g/l suspended sediment for 14 days. C = control insects, 0.0 g/l suspended sediment; T = test insects. Vertical lines = 95% confidence interval for mean ingestion rate. Ingestion (mg AFDW/Day) 14 and temperature. Similar growth responses have been reported for other insects by Beck (1980), Hynes (1970), and Lutz (1974). Another factor that may have affected feed- ing rates between trials was the conditioning GREAT BASIN NATURALIST Vol. 46, No. 1 Ingestion(mg AFDW/Day) Ingestion (mg AFDW/Day) o = —_ i) w i a 1 14 4 7 1 Days Fig. 2. Mean daily ingestion rates (mg AFDW/day) for second-year naiads of Pteronarcys californica exposed to: A, 1.5 g/l. B, 3.0 g/l suspended sediment for 14 days. C = control insects, 0.0 g/l suspended sediment; T = test insects. Vertical lines = 95% confidence interval for mean Ingestion (mg AFDW/Day) Ingestion (mg AFDW/Day) co : = 5 7 11 14 Days Fig. 4. Mean daily ingestion rates (mg AF DW/day) for late-instar larvae of Hesperophylax occidentalis exposed to: A, 1.5 g/l. B, 3.0 g/l suspended sediment for 14 days. C = control insects, 0.0 g/l suspended sediment; T = test insects. Vertical lines = 95% confidence interval for mean ingestion rate. time of the leaf material (Golladay et al. 1983). In our study, leaf disks used during our winter trials were conditioned for longer than one month; however, there was little evidence to support differential feeding rates between January 1986 leaves conditioned for one month vs. those conditioned for 2-4 months. The effect of suspended sediment on inver- tebrate ingestion appears to be largely influ- enced by the season and feeding status of the insects. Feeding rates of insects in a relatively active feeding mode, as reflected by the con- trol specimens, may not be as affected by suspended sediment as insects in a less active feeding mode. We speculate that the short- term effects of suspended sediments on feed- ing rates of insects vary seasonally. Our study provides evidence that prolonged exposure to suspended sediments can adversely affect both actively and nonactively feeding insects by reducing their mean daily ingestion rates. Previous research on the sublethal effects of suspended sediments on leaf processing by aquatic invertebrates is minimal and largely limited to zooplankton studies (Arruda et al. 1983, McCabe and O'Brien 1983). These studies demonstrated that suspended sedi- ments can decrease feeding rates of Daphnia spp. by at least 90%. These filter-feeders in- gested large quantities of silt from the water column, which resulted in dense packing of the gut with inorganic particles. A decreased filtering rate is likely caused by excessive gut loading (McCabe and OBrien 1983). Al- though the sediment ingestion rate would likely be lower among nonfilterers, it may have been a factor influencing the early-instar H. occidentalis larvae, which diplayed a sud- den decrease in feeding after 12 days’ expo- sure to 3 g/l suspended sediment in our study. Nontoxic materials, such as inorganic silt, or sublethal concentrations of toxicants may not cause spectacular and immediate impacts on aquatic macroinvertebrates, but they may cause a reduction in secondary production, energy-transfer efficiency, and nutrient cy- cling in stream ecosystems. LITERATURE CITED ANDERSON, N. H., AND J. SEDELL. 1979. Detritus process- ing by macroinvertebrates in stream ecosystems. Ann. Rey. Entomol. 24:351-377. ARRUDA, J. A., G. R. MARZOLF, AND R. T. FAULK. 1983. The role of suspended sediments in the nutrition of. zooplankton in turbid reservoirs. Ecology 64:1225-1235. BECK, S. D. 1980. Insect photoperiodism. 2d ed. Aca- demic Press, Inc., New York. 387 pp. HORNIG, BRUSVEN: AQUATIC ECOLOGY Sif BJORNN, T. C., M. A. BRUSVEN, M. P. MOLNaU, J. A. MILLt- GAN, R. A. KLamtT, E. CHACHO, AND C. SCHAYE. 1977. Transport of granitic sediment in streams and its effects on insects and fish. College of Forestry, Wildlife and Range Sciences. Bull. 17. 43 pp. BRUSVEN, M. A., AND C. E. Hornic. 1984. Effects of sus- pended and deposited volcanic ash on survival and behavior of stream insects. J. Kans. Entomol. Soc. 57:50-62. BRUSVEN, M. A., AND K. V. PRATHER. 1974. Influence of stream sediments on distribution of macroben- thos. J. Entomol. Soc. Brit. Columbia 71:25-32. CoRDONE, A. J., AND D. W. KELLEY. 1961. The influences of inorganic sediment on the aquatic life of streams. California Fish and Game 47:189-228. CuMMINS, K. W., ANDG. H. LaurF. 1969. The influence of substrate particle size on the microdistribution of stream macrobenthos. Hydrobiologia 34:145-181. ELDER, J. A., AND A. R. GAUFIN. 1964. Notes on the occur- rence and distribution of Pteronarcys californica Newport (Plecoptera) within streams. Great Basin Nat. 33:218-220. GoLiapay, S. W., J. R. WEBSTER, AND E. F. BENFIELD. 1983. Factors affecting food utilization by a leaf- shredding aquatic insect: leaf species and condi- tioning time. Holarctic Ecol. 6:157-162. GRraFIUS, E., AND N. H. ANDERSON. 1979. Population dy- namics, bioenergetics, and role of Lepidstoma quericina Ross (Trichoptera: Lepidostomatidae) in an Oregon woodland stream. Ecology 60:433-441. . 1980. Population dynamics, and role of two spe- cies of Lepidostoma (Trichoptera: Lepidostomati- dae) in an Oregon coniferous forest stream. Ecol- ogy 61:808-816. HErssT, G. M. 1980. Effects of burial on food value and consumption of leaf detritus by aquatic inverte- brates in a lowland forest stream. Oikos 35:411- 424. Hur bert, S. H. 1984. Pseudoreplication and the design of ecological field experiments. Ecol. Monogr. 54:187-211. Hynes, H. B. N. 1970. The ecology of running waters. University of Toronto Press, Toronto. 555 pp. LEMLY, A. D. 1982. Modification of benthic insect com- munities in polluted streams: combined effects of sedimentation and nutrient enrichment. Hydrobi- ologia 87:229-245. Lutz, P. E. 1974. Effects of temperature and photoperiod on larval development in Tetragoneuria cynosura (Odonata: Libellulidae). Ecology 55:370-377. MARTINSON, R. J., AND J. V. WarD. 1982. Life history and ecology of Hesperophylax occidentalis (Banks) (Trichoptera: Limnephilidae) from three springs in the Piceance Basin, Colorado. Freshwat. Inver- tebr. Biol. 1(3):41-47. McCaseE, G. O. AND W. J. O'BRIEN. 1983. The effects of suspended silt on feeding and reproduction of Daphnia pulex. Amer. Mid]. Nat. 110:324-337. MERRITT, R. W., AND K. W. Cummins. 1978. An introduc- tion to the aquatic insects of North America. Kendall/Hunt Publishing Company, Dubuque, Iowa. 441 pp. 38 GREAT BASIN NATURALIST O'Hor, J., AND J. B. WALLACE. 1983. Invertebrate drift, discharge, and sediment relations in a southern Appalachian headwater stream. Hydrobiologia 98:71-84. OxBerry, J. R., P. DOUDOROFF, AND D. W. ANDERSON. 1979. Potential toxicity of taconite tailings to aquatic life in Lake Superior. J. Water Poll. Contr. Fed. 50:240-251. REICE, S. R. 1980. The role of substratum in benthic macroinvertebrate microdistribution and _ litter decomposition in a woodland stream. Ecology 61:580-590. Vol. 46, No. 1 ROSENBERG, D. M., AND A. P. WIENS. 1975. Experimental sediment addition studies on the Harris River, N.W.T., Canada: the effect on macroinvertebrate drift. Verh. Internat. Verein. Limnol. 19:1568- 1574. SHoRT, R. A., AND P. E. MASLIN. 1977. Processing of leaf litter by a stream detritivore: effect on nutrient availability to collectors. Ecology 58:935-938. Wuirte, D. S., AND J. R. GAMMON. 1977. The effect of suspended solids on macroinvertebrate drift in an Indiana Creek. Proc. Indiana Acad. Sci. 86:182- 188. EFFECTS OF WATERSHED ALTERATION ON THE BROOK TROUT POPULATION OF A SMALL BLACK HILLS STREAM Timothy Modde’, Henry G. Drewes’, and Mark A. Rumble? ABSTRACT.—The impacts of adjacent landscaping activity and livestock presence on the brook trout (Salvelinus fontinalis) population of a small Black Hills stream were evaluated. Moderate changes in temperature, turbidity, and fecal coliform numbers did not influence brook trout densities. Stream morphometry, particularly factors affecting stream cover, appeared to have the greatest impact on numbers of trout. Brook trout were poor indicators of moderate changes in water quality, but they were adequate indicators of the physical perturbations within the stream. The aquatic biota of streams within the forests of our nation are being increasingly stressed by logging, livestock grazing, mining, road con- struction, and residential development. Impacts of these activities on streams are often mani- fested by changes in water quality or through physical changes in the habitat, such as channel modification or reduction in stream flow. Such environmental changes can produce biological responses that alter composition and abundance of resident species. Organisms sensitive to envi- ronmental change function as indicators of envi- ronmental modification and may be useful in evaluating the magnitude of an impact. Not all aquatic organisms exhibit equal sensi- tivity to the same stresses, however. Aquatic invertebrates are sensitive to water quality changes resulting from organic enrichment (Hilsenhoff 1977, Jones et al. 1981) and toxic pollutants (Hocutt 1975) but are not always effec- tive in evaluating moderate physical modifica- tions (Marsh and Waters 1980). Conversely, be- cause of their mobility, Price (1979) suggested that fishes may not be reliable indicators of change in water quality, although fish frequen- cies do respond to changes in structure of the habitat (Platts 1974, Binns and Eiserman 1979). The objective of this study was to evaluate the response of a resident brook trout (Salvelinus fontinalis ) population to water quality and habi- tat changes in a small Black Hills stream sub- jected to landscaping activities (clearcutting an adjacent slope and pond dredging) and intermit- tent livestock grazing within the watershed. Department of Wildlife and Fisheries Sciences, Box 2206, Brookings, South Dakota 2USDA Forest Service, Rocky. Mountain Forest and Range Experiment Station, Sout 39 METHODS The study area included five sampling sta- tions within the upper Slate Creek watershed (Fig. 1) in Pennington County, South Dakota. Two stations were established above a devel- opment site within the watershed, one on South Slate Creek and the other on Slate Creek proper, both first order streams. A third sampling station was located 0.1 km downstream from the development site on Slate Creek; two additional stations were es- tablished downstream at approximately 1.0 km intervals. Stations above the construction site were not exposed to grazing except for a confined watering site above station 1, whereas stations below the construction site were exposed to grazing during the late fall and winter months. Twelve water quality variables were mea- sured at three-week intervals during the spring and summer of 1981 and 1982. During the period between August 1981 and April 1982, water samples were collected at inter- vals ranging between six and eight weeks. Total hardness, alkalinity, dissolved oxygen, pH, and conductivity were determined on- site with field analysis units. Turbidity, total phosphorus (ortho and total), nitrogen (Kjel- dahl and ammonia), and fecal coliform bacte- ria were determined in the laboratory. Fishes were collected from each station by electrofishing a 61 m section of stream blocked from both directions by 6.4 mm mesh seines. Population estimates were repre- 57007. h Dakota School of Mines Campus, Rapid City, South Dakota 57701. 40 GREAT BASIN NATURALIST Vol. 46, No. 1 ea =H ae ) == Quadrangle Location ~ LEGEND ea Excavated Lake Basin en Slate Creek [| Construction Site =~ Stream Channel —--~ Hard Surface Road ' OC) Sampling Station Fig. 1. Map indicating location of the five sampling stations in relation to the Deerfield Park development site in Pennington County, South Dakota. sented by the total number of fish collected in collection. Fish were sampled at approxi- two consecutive electroshocking passes. mately six-week intervals between 9 June Length and weight of brook trout were 1981 and 29 August 1981 and between 24 May recorded and scale samples taken during each 1982 and 12 August 1982. Because of the ab- January 1986 sence of flow from the electroshocking site at station 2 during 1981, no fish were collected that year. Fish collected from each station on 24 May 1982 were marked with a subcutaneous latex injection to evaluate movement within the stream. Age was determined by analysis of scale annuli. Relative weights (W,) (Anderson 1980) were computed using standard weights pro- posed by Cooper (1961). Stream morphometry characteristics were measured in July 1982 at each of the five elec- trofishing stations. Eleven transects perpendic- ular to the stream bank and spaced 6.1 m apart along each 61-m section of stream were estab- lished. Stream width and water depth at 0.15-m intervals across the transect and at both banks were measured. Data from the 11 transects were pooled, and mean depth, width, and depth at stream-bank interface were calculated. Percent of stream canopy coverage at each transect was estimated by assigning numerical values from 1 to 5 corresponding to intervals of stream canopy cover (1 = 81%-100%, 2 = 61%-80%, 3 = 41%-60%, 4 = 21% -40%, 5 = 0%-20%) which shaded the stream surface. Bottom substrate at each station was also examined. A shovel was used to remove three samples of substrate at each station. Samples were air dried, weighed, and sifted through a series of USA Standard Test- ing sieves. Mean percent by weight of rubble (>76.2/mm), medium/coarse gravel (4.7—76.2/ mm), fine gravel (2.0—4.7/mm), coarse sand (1.0—2.0/mm), medium sand (0.5—1.2/mm), and fine sand/silt (<0.5/mm) was calculated for each station. Analysis of variance was utilized to test for differences in brook trout relative density (num- ber of fishes per 61 m of stream), relative weight (among stations, month, and year), and stream morphometry measurements (among stations). Because the number of brook trout collected at many stations was small, these data were trans- formed by adding 0.5 to the mean for each sta- tion for each date and the square root derived. Following analysis, the transformed data were then squared for presentation of results. Waller- Duncan's K-ratio t-test was employed to define differences. Differences were considered signif- icant at P < 0.05. RESULTS Twelve physicochemical parameters were evaluated on 16 dates from each of the five MODDE ETAL.: BROOK TROUT 4] 40 Lead —o—1981 —-— 1982 —-4-— Average 30 20 = (o} PRECIPITATION (centimeters) eS [o) Spearfish MONTHS Fig. 2. Monthly precipitation levels for the Lead and Spearfish, South Dakota, gauging stations in the Black Hills during 1981 and 1982. sampling stations. Variation in precipitation between 1981 and 1984 (Fig. 2) affected water quality between years. Based upon long-term averages from Lead and Spearfish, precipita- tion in the Black Hills was above normal in 1982, whereas 1981 was a year of lower than normal precipitation. Increased runoff re- sulted in elevated stream flows throughout the summer of 1982. Mean values for dis- solved oxygen and organic phosphorus were significantly higher in 1982 and mean conduc- tivity and pH were significantly lower. Mean fecal coliform number, ammonia nitrogen, and total (Kjeldahl) nitrogen values were higher from all stations in 1982, but these differences were not significant. Seven physicochemical variables differed significantly among stations. Mean turbidity was significantly higher at stations 3, 4, and 5 than upstream at stations 1 and 2 (Table 1). The highest mean turbidity value was at sta- tion 3 (35.7 ntu) just below the development site. Mean temperatures at stations 3 and 4 were significantly higher than upstream of sta- tions 1 and 2. Temperature decreased signifi- 42 GREAT BASIN NATURALIST Vol. 46, No. 1 rn 5 m : ] TABLE 1. Means of water quality parameters among five stations on Slate Creek. Variable (Units of measure) 1 2 Turbidity (NTU) Wd 3.9% Temperature (C) TOME 9. 1° pH Us Wie Conductivity (mhos) 161.0° ey 2 Hardness (mg/1] CaCO’) 110.3° 82.5° Alkalinity (mg/1 CaCO") 101.2° 81.9? Fecal coliform (#/100ml) ee 78.8” Station 3 4 5 35.7° 28.4? 26.9° 12.5? 11.5°° 10.4°° 708 7.58 1 163.3° 163.2° 167.6° 102.5” 105.37> 105.0°° 94.1" 97.8* 100.0° 100.2" 163.9 309. 4° 'Means followed by the same superscript for each variable are not different (P> 0.05) based on Waller Duncan’s K-ratio t-test. TABLE 2. Numbers and species of fishes collected from Slate Creek and South Slate Creek between June 1981 and August 1982. 1981 June July August Station 1 Brook Trout 5 2 1 White sucker 0 1 0 Longnose dace 0 0 0 Fathead minnow 0 (0) 0 Station 2 Brook trout 0 0 0 White sucker 0 0 0 Longnose dace 0 0 0 Fathead minnow 0 0 0 Station 3 Brook trout 7 4 4 White sucker 1 2 2 Longnose dace 1 2 5 Fathead minnow 0 0 0 Station 4 Brook trout 15 44 26 White sucker 0 0 0) Longnose dace 6 ri 2 Fathead minnow 0 0 0 Station 5 Brook trout 19 8 20 White sucker U 2 0 Longnose dace 14 21 53 Fathead minnow 0 (0) 0 1982 x May July August xX Oy, 8 12 6 8.7 0.3 0 0 0 0.0 0.0 0 0 0 0.0 0.0 0 0 0 0.0 0.0 5 8 5 6.0 0.0 0 0 0 0.0 0.0 3 0 2 LG 0.0 0 0 0 0.0 5.0 5 Be 2 4.0 13 0 5 1 2.0 2.7 2 23 23 16.0 0.0 0 1 0 0.3 28.3 9 14 19 14.0 0.0 0 5 5 3.3 5.0 1 1 7 3.0 0.0 0 0 0 0.0 15.7 12 12 ll 11.7 3.0 0 10 21 10.3 29.0 0 6 10 5.3 0.0 0 0 0 0.0 cantly from a mean of 12.5 C at station 3 to 10.4 C downstream at station 5. Mean temperature at station 5 was not significantly different from mean values from stations 1 and 2. Conductivity, pH, hardness, and alkalinity were all signifi- cantly lower at station 2 than at any of the other stations. Mean total coliform numbers at station 5 were significantly higher than at stations 2 and 3. No significant differences were detected among the five stations for phosphorus (or- thophosphate and organic phosphate) or nitro- gen (ammonia and organic), but nutrient levels were the highest at stations 4 and 5. Brook trout, an introduced game species, was the most abundant fish in the study area and was the only species consistently collected at stations both above and below the development site (Table 2). Brook trout composed 53.4% of the fishes collected from the five stations over the two-year period. Other species occurring within the study area were white sucker (Catostomus commersoni) (11.3%), longnose dace (Rhi- nichthys cataractae ) (35.1%), and fathead min- now (Pime-phales promelas ) (0.2%). Differences among stations were observed during both years of the study. Significant dif- . . January 1986 MODDE ETAL.: BROOK TROUT 43 TaBLE3. Mean densities of brook trout per 61m of stream from five stations along Slate Creek during 1981 and 1982.’ Station Number Year 1 2 3 4 5 1981 3.0° 0.5° Bile 28.6° N5rGee 1982 9.0° 6.4" 4.4° 14.2° Qe" Means followed by the same superscript across rows are not different (P > 0.05) from each other based on Waller Duncan’s K-ratio t-test. TABLE 4. Mean values for stream morphometry characteristics among five stations along Slate Creek.! Variable Station (Units of measure) 1 2 3 4 5 Stream depth (cm) 12.9? 27.9? 14.8 94.7° 28.1? Stream-shore depth (cm) bye eee 5.0" 19.0°° Joely Stream cover (%) 3.6" 5.0” 3.6" 1BSE Be Fine sand/silt (%) 2.6" ON 29.6” LE 2266 ric Stream width (cm) 93.3°° 260.7" 94.0°° GIA lieses 106.7° Means followed by the same superscript for a variable are not different (P > 0.05) from each other based on Waller Duncan’s K-ratio t-test. ferences (P S 0.05) were observed both among stations and station-year interaction. Due to the interaction observed, data from each year was subjected to a separate analysis of variance. The results indicated that during 1981, fish were most abundant in downstream stations (i.e., 4 and 5). A weaker trend among stations in trout density was observed during 1982, although the highest numbers of brook trout were still col- lected in the downstream stations (Table 3). Increases in trout density were generally con- sistent with increases in mean stream depth and mean depth at the bank (Table 4). During both years brook trout densities below the develop- ment site were higher at station 4, which had the greatest summer canopy cover and the lowest amount of sand or silt deposition among down- stream stations. Significant differences in stream width among stations did not appear to be re- lated to trout density. Although mean bank depth and mean depth of stream were high at station 2, trout densities were low because flows were not perennial through the 61 m electrofish- ing section. Considerable variation in brook trout (relative weight) W, was observed during the study (Fig. 3). During 1981, when stream flows were low, mean W, decreased from 95.0 in early: June to 79.7 in late August. However, in 1982, when stream flows were higher, W, declined little dur- ing the summer months. Analysis of variance indicated significant (P < 0.05) differences in W, among years, months, and stations. The highest mean W, values were observed at station 2; the lowest were at station 4. The brook trout population of Slate Creek consisted primarily of age-class I and II fish; age-class III fish were collected only in 1982 (Fig. 4). The 1980 year-class was the strongest cohort during both years of the study. Re- cruitment within the study area occurred pri- marily at station 4 and, to a lesser extent, at station 3. During both years of the study, the numbers of young-of-the-year fish collected were lower than the number of yearlings. Age-class I fish composed that largest portion of that Slate Creek brook trout population at all stations during the study, and were always dominant at stations | and 2. No movement of brook trout was observed among stations. Mean recapture rates of fish marked in late May were 41.0% on 1 July and 12.8% on 12 August 1982. DISCUSSION Lotic populations of brook trout are typified by short-lived populations (McAfee 1965) in- habiting headwater streams (Neves and Par- due 1983). Brook trout exhibit a high toler- ance to environmental variation. Lee and Rinne (1980) observed brook trout, a char, to have a slightly higher tolerance to elevated temperature fluctuation than several other trout species, and Brett (1956) reported that brook trout have a greater cold tolerance than several trout species. In addition, brook trout are also tolerant to extremes to both pH (Daye and Garside 1975) and turbidity (Gradall and Swenson 1982). Tolerance to environmental 44 GREAT BASIN NATURALIST 100 90 80 Relative Weight 70 Month Fig. 3. Mean relative weight (W,) values for brook trout by month from Slate Creek and South Slate Creek for 1981 and 1982. change has no doubt led to the success of brook trout stocking within a wide geographi- cal range outside its native distribution (Mac- Crimmon and Campbell 1969). Tolerance to changes in water quality re- sults in low sensitivity to some forms of envi- ronmental perturbation. In the present study, brook trout were poor indicators of the observed changes in turbidity, temperature, and nutrient loading. Natural fish populations have been considered inferior to macroinver- tebrate communities as water quality indica- tors even though individual fish species have been successfully utilized in bioassays (Price 1979). However, several researchers (e.g., Binns and Eiserman 1979, Raleigh 1982, Par- sons et al. 1981) have studied the importance of physical characteristics of streams in deter- mining trout density. Brook trout densities in the present study were responsive to changes in physical characteristics of the stream, par- ticularly stream flow, mean stream depth, depth at the bank, and canopy cover—the major components of cover in a small stream. Both Stewart (1970) and Hunt (1971) similarly reported that depth and cover were dominant factors affecting brook trout densities in streams. Fraley and Graham (1981) observed that cover, substrate, and depth were the pri- mary factors among 30 physical habitat char- acteristics measured that best predicted den- sities of cutthroat trout (Salmo clarki lewisi) and bull trout (Salvelinus confluentus ). Relative weights were higher for brook trout during 1982, when high precipitation resulted in a greater quantity of drift organ- isms available (Drewes 1984). The relation- ship of W, to stations was probably due to intraspecific competition, with the lowest val- Vol. 46, No. 1 ues occurring at the station with the highest relative density of trout. Condition of trout, as expressed by W,, was also influenced more by natural variation than water quality. As primary components of the ichthyofauna in headwater mountain streams, brook trout represent one of the initial biotic components impacted by nonpoint sources of watershed disruption. Because of their tolerance to changes in the water medium, brook trout are not good indicators of moderate alterations in water quality. Brook trout appear to be sensi- tive to changes in stream morphometry and should be adequate indicators of physical dis- ruptions within streams. ACKNOWLEDGMENTS Funding for the project was provided by the U.S. Forest Service through an Eisen- hower Consortium grant and project funds from the Rocky Mountain Forest and Range Experiment Station. Thanks are expressed to Brian Smith, Michele Deisch, and Larry Bea- gle for assisting in field collection and to Dr. Charles G. Scalet, Dr. Lester D. Flake, Ron Koth, and Dick Ford for reviewing this manuscript. LITERATURE CITED ANDERSON, R. O. 1980. Proportional stock density (PSD) and relative weight (W,): interpretive indices for fish pop- ulations and communities. Pages 17-33 in S. Gloss and B. Shupp, eds., Practical fisheries management: more with less in the 1980s. New York Chapter, Amer. Fish. Soc., Bethesda. Maryland. Binns, N. A., AND F. M. EISERMAN. 1979. Quantification of fluvial trout habitat in Wyoming. Trans. Amer. Fish. Soc. 108:215-228. Brett, J. R. 1956. Some principles in the thermal require- ments of fishes. Quar. Rev. of Biol. 31:75—-87. Cooper, E. L. 1961. Growth of wild and hatchery strains of brook trout. Trans. Amer. Fish. Soc. 90:424—438. Daye, P. G., AND E. T. GarsibE. 1975. Lethal levels of pH for brook trout (Salvelinus fontinalis). Canadian J. Zool. 53:639-641. Drewes, H. G. 1984. Factors affecting water quality and macroinvertebrate distribution within a small Black Hills stream. Unpublished thesis, South Dakota State University, Brookings. FRALEY, J. J., AND P. J. GRAHAM. 1981. Physical habitat, geo- logic bedrock types and trout densities in the tribu- taries of the Flathead River Drainage, Montana. Pages 178-185 in N. Armatrout, ed., Acquisition and utilization of aquatic habitat inventory information. Amer. Fish. Soc., Bethesda, Maryland. January 1986 Age Classes © 60 E =} S 3 40 x (e) e al XO) H IC ine) MODDE ETAL.: BROOK TROUT 45 PLL LESS NN WILLLLLLLLLALLLLL LLL AML MLL LL WLP LLLLLL LL LS CHBMM a CLLLLLLLLALLLE LLL LAREN EEN | Stations Fig. 4. Age distribution and abundance of brook trout among stations from the Slate Creek study during 1981 and 1982. GRADALL, K.S., AND W. A. SWENSON. 1982. Responses of brook trout and creek chubs to turbidity. Trans. Amer. Fish. Soc. 111:392-395. HILSENHOFF, W. L. 1977. Use of arthropods to evaluate water quality of streams. Technical Bulletin 100, Wisconsin Dept. Nat. Res., Madison. Hocutt, C. H. 1975. Assessment of a stressed macroinverte- brate community. Water Res. Bull. 11:820—835. Hunt, R. L. 1971. Responses of a brook trout population to habitat development in Lawrence Creek. Tech. Bull. 48, Wisconsin Dept. Nat. Res., Madison. Jones, J. R., B. H. Tracy, J. L. SaBAuGH, D. H. HAZELWOOD, AND M. M. Sarr. 1981. Biotic index tested for ability to assess water quality of Missouri Ozark streams. Trans. Amer. Fish. Soc. 110:627-637. LEE, R. M., AND J. N. RINNE. 1980. Critical thermal maxima of five trout species in the southwestern United States. Trans. Amer. Fish. Soc. 109:632-635. MacCrimMon, H.R., AND J.C. CAMPBELL. 1969. World distri- bution of brook trout (Salvelinus fontinalis). J. Fish. Res. Board of Canada 26:1699-17235. MCAFEE, W. R. 1965. Eastern brook trout. Pages 242-260 in A. Calhoun, ed., Inland fisheries management. Cali- fornia Dept. of Fish and Game, Sacramento. Marcu, P. C., AND T. F. WaTERS. 1980. Effects of agricul- tural drainage development on benthic inverte- brates in undisturbed downstream ranches. Trans. Amer. Fish. Soc. 109:213—223. NEVES, R. J., AND G. B. ParDUE. 1983. Abundance and production of fishes in a small Appalachian stream. Trans. Amer. Fish. Soc. 112:21—26. Parsons, M. G., J. R. MAXWELL, AND D. HELLER. 1981. A predictive fish habitat index model using geomor- phic parameters. Pages 85-91 in N. Armatrout, ed., Acquisition and utilization of aquatic habitat inventory information. Amer. Fish. Soc., Bethesda, Maryland. Piatrs, W. S. 1974. Geomorphic and aquatic conditions influencing salmonids and stream classification. U.S. Forest Service, Surf. Environ. and Mining Prog. Rep., Washington, D.C. Price, D. R. H. 1979. Fish as indicators of river water quality. Chapter 8 in A. James and L. Evison, eds., Biological indicators of water quality. John Wiley and Sons, New York. STEWART, P. A. 1970. Physical factors influencing trout density in a small stream. Unpublished disserta- tion, Colorado State University, Fort Collins. FLORISTIC ANALYSIS OF THE SOUTHWESTERN UNITED STATES Steven P. McLaughlin! ABSTRACT. —A study was made of the distributions of native, terrestrial, vascular plants occurring in 50 local floras from throughout the Basin and Range and Colorado Plateau physiographic provinces of the southwestern United States. The objectives of the study were to objectively define and describe the floristic elements—assemblages of species with roughly coincident geographic distribution—occurring in the southwestern United States and to deter- mine what such assemblages reveal about the floristic history of the region. The total flora (native, terrestrial species only) of the Southwest is estimated at 5,458 species, 77% of which were recorded in | or more of the local floras. Nearly 22% of these species are endemic to the study region. A majority of the species were found to be relatively rare. The average range of a species included only 4 floras, and 90% of the species were recorded from 11 or fewer floras; only 81 species (1.5%) were recorded from 50% or more of the floras. Trees constitute 2% of the regional flora and have the widest average distribution; perennial herbs constitute 59% of the flora and have the most restricted distributions. Factor analysis was used to identify seven floristic elements for the region: a Great Basin element, a Mojavean element, a Colorado Plateau element, a Chihuahuan element, an Apachian element, and a Mogollon element. This factor analysis solution was shown to satisfy criteria of interpretability and consistency. The Mojavean, Colorado Plateau, and Apachian elements are believed to be autochthonous. The other four elements show angen overlap in species composition w rith one or more adjacent regions. Each floristic element is mapped to show its geographic form and distribution. Analysis of these maps shows how the existence of objectively defined floristic elements is not contradictory to either the individualistic view of the distribution of a species or local continuity of vegetation and flora. The rarity of the majority of species and the clear association of floristic elements with rather narrowly circumscribed Holocene environments suggests that many Southwestern species have migrated little and are of rather recent, probably postglacial origin. Geographic “principles” derived from the distribution patterns of relatively few, wide- spread, dominant, usually woody species may not be applicable to entire, regional floras. Plant geographers have long recognized dence between floristic groups and floristic that plant species can be grouped on the basis _ provinces (e.g., Gleason and Cronquist 1964), of similarities among their geographic distri- in which case the floristic group is the charac- butions. Such floristic groups have generally _ teristic flora of the floristic province. More been termed “geographical elements” by Eu- _ often floristic groups are conceived as assem- ropean phytogeographers (Stott 1981, Cain — blages of variable and often overlapping areal 1947). The same concept is embodied in such extent, including a few wide-ranging types terms as “natural floristic areas’ (Raup 1947), and many more narrowly defined types. A “floristic assemblage” (Cain 1944), “floristic good example is the set of “areal types” used group (Gleason and Croquist 1964), “areal by Whittaker and Niering (1964) in their anal- types’ (Whittaker and Niering 1964), “cate- ysis of the flora of the Santa Catalina Moun- gories of geographic origin” (Stebbins 1982), _ tains of southern Arizona. Included are wide- or “directional classes” (Meyer 1978) of Amer- spread and overlapping “temperate,” “wes- ican authors. The term element is used uncrit- tern,” “northern,” and “southwestern” types ically in the American literature asa synonym along with more restricted and largely dis- for taxon or taxa. crete Sonoran,” “Rocky Mountain,” “Plains,” A concept related to that of floristic assem- “Madrean,” and “Chihuahuan” types. blages is that of floristic or biotic provinces. This chaos in terminology regarding floris- Provinces in the sense of Dice (1943) are rea- _ tic assemblages is a consequence of the inade- sonably discrete areas with characteristic quate empirical and theoretical basis for such physiography, climate, vegetation, flora, and concepts in the American tradition. Gleason fauna. There may be a one to one correspon- (1926) challenged the validity of the organis- lOffice ca Arid Lands Studie s, University of Arizona, Tucson, Arizona 85719. 46 January 1986 mic concept of associations as applied to vege- tation by F. E. Clements and argued further that each species has its own individualistic environmental requirements and migrational history. It should follow, therefore, that each community of species is unique and any at- tempt to identify repeatable associations must be highly subjective. The “individualistic con- cept’ and the related “continuum concept” have been accepted by most American plant ecologists (McIntosh 1967). The ideas of Gleason were very broadly applied to plant geography by Mason (1947) and Cain (1944, 1947). Mason (1947) invoked the individualistic nature of the species and the principles of past migrations of species (as revealed in the fossil record) to argue against the existence of persistent and recognizable floras. Cain (1947) believed that floristic ele- ments could be recognized only relative to the particular area under investigation and not as a universal system of floristic regions. He linked the ideas of floristic groups and plant association types, and went on to argue against the objective recognition of associa- tion types, implicitly casting doubt on the objective recognition of floristic assemblages of any form. The current attitude toward floristic assem- blages is well illustrated in a statement by Johnston (1977:356): “Recurrent distribution patterns” may simply be the result of intuitive (but in this case unfortunate?) ten- dencies to lump and generalize, in the way the eye tends to connect totally unrelated star-clusters into meaningless but “recognizable” patterns in the night sky. A recent text on biogeography (Brown and Gibson 1983) discusses the association/contin- uum conflict but makes no reference to the problem of floristic elements. If each plant species is uniquely and inde- pendently distributed, and therefore recogni- tion of floristic assemblages must be inher- ently arbitrary and subjective, why have so many phytogeographers nevertheless recog- nized such groups? Can phytogeographical el- ements be described more objectively and, if so, how can they be reconciled with individu- alistic concepts and what do they mean in terms of the evolutionary histories of their species? The objective of this paper is to an- swer these questions through a floristic analy- MCLAUGHLIN: FLORISTIC ANALYSIS 47 sis of vascular plant distributions recorded in local floras from the southwestern United States. METHODS Study Region The study region includes the Colorado Plateau Physiographic Province and the Basin and Range Physiographic Province north of the U.S.-Mexico International Boundary, both as mapped by Hunt (1967) (Fig. 1). This region is bounded by the Sierra Nevada and southern California transverse ranges on the west; by the Columbia Plateau on the north; and by the Rocky Mountains and southern Great Plains on the east. Only the southern boundary is artificial. This region will be re- ferred to simply as the Southwest. The majority of the region lies between the two great Western mountain systems— Sierra Nevada on the west and Rocky Mountains to the east. The entire region is arid to semiarid. The Great Basin section of the Basin and Range Province experiences hot summers, cold winters, and mostly winter precipitation. The southeastern portion of the Basin and Range Province is at the opposite end of the climatic spectrum for the region, with hot summers, mild winters, and mostly summer precipitation. The most arid portion of the region centers around the lower Colorado River Valley. The study region has generally been recog- nized as a natural one by biogeographers. One of the earliest formal treatments is that of Dice (1943) and a recent one is that of Cronquist (1982). Both recognize a semiarid natural re- gion lying between the Sierra Nevada and the Rocky Mountains that extends into northern Mexico. Cronquist's Great Basin Province es- sentially includes Dice’s Artemisian and Navahonian provinces; Cronquist’s Sonoran Subprovince is composed of Dice's Mohavean and Sonoran provinces; and their Chihuahuan regions are similar. The principal difference is that there is nothing in Cronquist’s treatment that corresponds closely to Dice’s Apachian Province of the uplands of southeastern Ari- zona, southwestern New Mexico, and north- western Sonora. 48 GREAT BASIN NATURALIST PACIFIC Data Base Fifty local floras from the Southwest were examined (Table 1). These floras represent a systematic sample of the regional southwest- ern flora and were selected to provide as thor- ough and uniform coverage of the study re- gion as possible (see Fig. 1). Vol. 46, No. 1 WYOMING YS W \ l CHIHUAHUA l \ a Fig. 1. Map of study region, Southwestern United States. Numbers refer to local floras listed in Table 1; letters in boxes refer to peripheral regions listed in Table 6. "90 34av vuUals Complete bibliographic citations for 33 of the 50 local floras can be found in Bowers’ (1982) annotated bibliography.Flora 42 is based both on the publication referenced in Bowers (1982) and on an unpublished check- list distributed by Natural Bridges National Monument. Flora 13 consists of those plants listed by Twisselman (1967) as occurring in ~ January 1986 MCLAUGHLIN: FLORISTIC ANALYSIS 49 TaBLE 1. Southwestern local floras used in floristic analysis. Map numbers refer to map locations in Figure 1. Map Number of location Flora designation species Reference 1 Sheldon National Wildlife Refuge 489 Rogers & Tiehm (1979) Santa Rosa Mountains 318 Bowers (1982) 3 Ruby—East Humboldt Mountains 507 Lewis (1971) 4 Raft River Mountains 295 Bowers (1982) 5 Stansbury Mountains 476 Taye (1983) 6 Deep Creek Mountains 395 Bowers (1982) i Wheeler Peak area 380 Bowers (1982) 8 Toiyabe Mountains 440 Bowers (1982) 9 White Mountains (California-Nevada) 677 Lloyd & Mitchell (1973) 10 Grapevine Mountains 463 Kurzius (1981)- ll Beaver Dam Mountains 489 Bowers (1982) 12 Spring Range (Charleston Mountains) 603 Bowers (1982) 13 Eastern Kern County 368 Twisselman (1967) 14 Newberry Mountains 385 Holland (1982) 15 Granite Mountains 376 Thorne et al. (1981) 16 Joshua Tree National Monument 534 National Park Service (1973) 17 Western Colorado Desert Region 44] Clemons (1984) 18 Eastern Imperial County 227 McLaughlin & Bowers (unpublished) 19 Hualapai Mountains Planning Unit 623 Bowers (1982) 20 Harcuvar Planning Unit 49] Bowers (1982) 21 Skull Valley Planning Unit 488 Bowers (1982) 22 White Tank Mountains 304 Bowers (1982) 23 Organ Pipe Cactus National Monument A477 Bowers (1982) 24 Sierra Ancha Experimental Forest 655 Bowers (1982) 25 George Whittell Wildlife Preserve 398 Bowers (1982) (Aravaipa Canyon) 26 Saguaro National Monument, Rincon 799 Bowers (unpublished) Mountain Unit Hy Mule Mountains 387 Wentworth (1982) 28 Chiricahua National Monument 619 Bowers (1982) 29 Animas Mountains 620 Bowers (1982) 30 Manzano Mountains 349 Bowers (1982) 31 Jornado Experimental Range 493 Bowers (1982) 32 White Mountains (New Mexico) 895 Bowers (1982) 33 Hueco Tanks State Park 315 Worthington (1980) 34 Carlsbad Caverns—Guadalupe Mountains 543 Bowers (1982) 35 Davis Mountains 461 Sikes and Smith (1975) 36 Big Bend National Park 751 Warnock (1967) 37 Dinosaur National Monument 304 Bowers (1982) 38 Capitol Reef National Monument 402 Bowers (1982) 39 Colorado National Monument 302 Bowers (1982) 40 Bryce Canyon National Park 402 Buchanan & Graybosch (1982) 4] Natural Bridges National Monument 298 Bowers (1982), National Park Service (no date) 42 Mesa Verde 333 Bowers (1982) 43 Lower Grand Canyon area 250 Bowers (1982) 44 Navajo National Monument 252 Bowers (1982) 45 Wupatki National Monument 178 Bowers (1982) 46 Canyon de Chelly National Monument 396 Bowers (1982) 47 Oak Creek Canyon and vicinity 507 Bowers (1982) 48 Petrified Forest National Park 277 Bowers (1982) 49 Mount Taylor 274 Bowers (1982) 50 Datil Mountains 483 Bowers (1982) the arid shrub, creosote bush, and shadscale scrub portions of eastern Kern County, Cali- | fornia. Flora 17 consists of those plants listed | by Clemons (1984) as occurring in the Val- _lecito, Carrizo, and Borrego floristic areas of eastern San Diego County, California. Floras 18 and 26 are from projects in progress. The number of species listed in these local floras varies from a low of 178 to nearly 900 species. The species numbers given in Table 1 50 GREAT BASIN NATURALIST are in every case lower than the numbers given in the original reports, since several classes of species were not included in the present analysis: a. Introduced species. b. Strict aquatics, i.e., floating, submerged, and emergent plants of permanent lotic habitats (ponds, lakes, reservoirs, cattle tanks, etc.) c. Synonyms at the species level. d. Species listed by authors as “likely” to occur in their areas. Aquatic species were excluded because of their limited value in comparative geographic studies. Such species tend to be very wide- spread but are relatively uncommon in dry regions, depending on the occurrence of suit- able (often man-made) habitats. Kartesz and Kartesz (1980) and Welsh et. al (1981) were used to achieve consistency and uniformity in nomenclature and to identify synonymous plant names. In the remainder of this paper, flora of the study region will refer to the na- tive, vascular, terrestrial flora as defined above. In addition to recording the presence of species in each of the 50 local floras, manuals and checklists for the region (Holmgren and Reveal 1966, Cronquist et al. 1972, 1977, 1984, Kearney and Peebles 1969, Martin and Hutchins 1980, Munz 1968, Correll and John- ston 1970) were consulted to determine what additional species occur in the Southwest. Fi- nally, again using regional manuals, I rec- orded occurrences of all southwestern species in each of seven adjacent or peripheral regions (see Fig. 1): (A) Pacific region (roughly equiva- lent to the California floristic province), (B) Columbia Plateau, (C) Rocky Mountain re- gion, (D) Great Plains, (E) Sonoran Desert region, (F) Sierra Madre region, taken here as the area in northern Mexico between the Sonoran and Chihuahuan deserts, and (G) Chihuahuan Desert region. The resulting lists of southwestern species occurring in these peripheral regions were used as floras in the data analysis (see below) to examine the rela- tionships between the Southwest flora and that of the peripheral regions, and to distin- guish allochthonous from autochthonous ele- ments. Data Analysis Similarities in the species composition of the 50 local floras and seven peripheral re- Vol. 46, No. 1 gions were examined using factor analysis. Factor analysis is a family of ordination tech- niques used to examine the pattern and struc- ture of large data matrices. Factor analysis has long been used in sociological and geographic studies, and its applications for plant ecology have been discussed by Goodall (1954) and for phytogeography by Jardine (1972). Since it is essentially a nonhierarchical clustering proce- dure, factor analysis is a particularly appropri- ate tool for examining floristic affinities (Jar- dine 1972). Factor analyses are of two basic types. Con- sider a matrix that lists the presences of s species (rows) in n floras (columns). A Q-mode analysis would examine the relationships in an s x s matrix of correlations or similarities among the s species; an R-mode analysis would examine the relationships in an n x n matrix of correlations or similarities among the n floras. Ideally one would prefer to per- form a Q-mode analysis to investigate floristic affinities, but two problems arise: there is no suitable coefficient or similarity index avail- able to correlate species ranges when most of the species are very rare; and available factor analysis programs are limited to -100 variables (species in this case), a number smaller than the smallest local flora used in this study. I therefore used an R-mode analysis based on similarity indices among the floras and pe- ripheral regions. The R-mode and Q-mode solutions should agree whenever clear-cut patterns actually exist in the data (Bryant et al. 1974). The similarity index used was that of Otsuka (as given in Simpson 1980): IS Gene c,/ (A,B,)"” where: A, = number of species in flora i; B, = number of species in flora j; = number of species common to flo- ras i and j. | © | Since IS6,.,4, can take only positive values be- tween 0 and 1, the analysis used here is, in the terminology of Pielou (1984), one using stan- dardized but noncentered data. With data in this form, the first several factors are usually unipolar, i.e., all or nearly all the factor load- ings (see below) are of the same sign, all posi- tive or all negative. Furthermore, there are as many unipolar factors as there are qualita- tively different clusters of data points (Pielou 1984). | January 1986 MCLAUGHLIN: FLORISTIC ANALYSIS 51 TABLE 2. Statistical summary of the native, terrestrial flora of the Colorado Plateau and Basin and Range provinces of the southwestern United States. Families Endemic species Taxonomic group Genera Species no. % Ferns and fern allies 9 24 120 3 (G2%5) Gymnosperms 3 Uf 36 2 ( 5.6) Monocotyledons 10 136 723 63 ( 8.7) Dicotyledons 105 827 4579 1119 (24.4) Totals 127 994 5458 1187 (21.7) The factoring program used was that of Nie et al. (1975). I used principal components fac- tor extraction with orthogonal (varimax) rota- tion, probably the most commonly used fac- toring technique in ecological studies. Several aspects of the analysis were altered later to check on the consistency of the solution (see below). RESULTS Description of the Flora The flora of the study region is summarized in Tables 2-6. A total of 4,185 species were recorded from the 50 local floras; 1,273 addi- tional species native to the study region were identified from the literature. The estimated total regional flora (5,458 species) is probably somewhat conservative, since the flora of the intermountain region in Utah and Nevada is still being compiled and monographed. The sample of species in the 50 local floras, how- ever, probably represents at least 70%—75% of the regional flora. Table 2 lists total species and endemic spe- cies by major taxonomic group. Within the study region, 21.7% of the species are en- demic. Most of the endemic species are di- cotyledons, and, in fact, nearly a quarter of the dicotyledons are endemic. The proportion of endemics in the study region is much lower than that in the California floristic province to the west (47.7%) but higher than that of other continental areas including Alaska (5.9%), Texas (9.0%), or the northeastern United States (13.5%) (Raven and Axelrod 1978). There is a rather high ratio of species per genus (5.5) in the Southwest, similar again to that of the California floristic province (5.6) and higher than for most other continental areas (3.0—4.0) (Raven and Axelrod 1978). The most common families and genera within the study region are listed in Table 3. Commonness as used in this paper refers to ubiquity rather than abundance (Preston 1948), although ubiquitous taxa are also usu- ally abundant (Hengeveld and Haeck 1982, Brown 1984). The Asteraceae, which is the largest family, accounting for 17.1% of the total species in the flora, is also the most com- mon family, with 78.6 species/flora. Poaceae is the second most common family even though it is represented by more than 100 fewer species than the Fabaceae, the third most common family. The families Fabaceae, Brassicaceae, Scrophulariaceae, _ Boragi- naceae, Polygonaceae, and Cactaceae are characteristic families of the region in that they have both a large number of species and a high proportion (>25%) of their species en- demic within the region. The five most com- mon genera—Eriogonum, Astragalus, Cryp- tantha, Penstemon, and Phacelia —also have a high proportion of endemic species. Several large genera common within but not charac- teristic of the region, including Carex, Jun- cus, and Salix, are mostly found in relatively mesic habitats. The frequency distribution for the 5,458 species occurring in the Southwest is shown in Figure 2. The average number of floras per species is 4.07. Only 81 species (1.5%) were recorded from one-half or more of the local floras (Table 4). Most of these widespread spe- cies have several subspecific taxa within the region, indicating that they are genetically variable species. The widespread species listed in Table 4 are found throughout most of western North America. The vast majority of species in the region, however, are rare. Nearly two-thirds of the species are recorded from three or fewer floras. The distribution patterns of southwestern species, as revealed by their occurrence in the 50 local floras examined, are probably reliable evidence for the rarity of most species in this 59 GREAT BASIN NATURALIST Vol. 46, No. 1 TABLE 3. Commonest families and genera of native, terrestrial plants in the Colorado Plateau and Basin and Range provinces. Species Total per Endemic species Taxon species flora number % 10 MOSTCOMMON FAMILIES: Asteraceae 932 78.6 213 (22.9) Poaceae 359 42.5 22 ( 6.1) Fabaceae 493 28.2, 154 (31.2) Brassicaceae 208 16.6 66 (31.7) Scrophulariaceae 276 16.3 114 (41.3) Boraginaceae 161 13.2 52 (32.3) Polygonaceae 12) 12.8 73 (42.4) Polemoniaceae 144 11.8 37 (25.7) Rosaceae 110 Li, 2 23 (20.9) Cactaceae 111 9.7 33 (29.7) 20 MOsTCOMMON GENERA: Eriogonum 120 9.4 64 (53.3) Astragalus 189 7.4 lll (58.7) Cryptantha 79 6.0 39 (49.9) Penstemon 118 5.3 70 (59.3) Phacelia i 5.2 35 (45.5) Carex 97 5.0 3 ( 3.1) Muhlenbergia 48 4.8 7 (14.6) Opuntia 36 4.7 7 (19.4) Erigeron 72 4.7 29 (40.3) Artemisia 22 3.8 1 ( 4.5) Juncus 26 3.8 0 (0 ) Chamaesyce 40 3.6 6 (15.0) Senecio 50 3.6 9 (18.0) Gilia” 46 B55) 14 (30.4) Bouteloua 19 3.4 pes ( 5.2) Brickellia 36 3.4 12 (33.3) Oenothera” 29 ~ oS) 6 (20.7) Castilleja 45 Bio 13 (28.9) Lupinus 57 3.2 13 (22.8) Salix 28 3.2 I ( 3.6) ‘excluding Ipomopsis, Leptodactylon. Excluding Calyophus, Camissonia. region. Although I use the term local in refer- ence to the floras, most of the floras used in this analysis cover rather extensive areas. 3 Many of the floras are for small to large moun- tain ranges, national parks, and other areas that encompass a diversity of habitats. Within 3 the boundaries covered by each flora there undoubtedly occur habitats suitable for many species not known to occur in the floras. Even 2 if every species occurred on the average in one additional flora, it would not greatly change the statistics of species distributions; | yet it would increase the size of each local flora by -110 species. It is unlikely that 100 species (on the average) could be added to each local Log Number of Species 0 flora, even through very extensive further col- 0 lo 20 30 40 50__ lection. Number of Floras The composition of the flora by life form is Fig. 2. Frequency distribution showing numbers of summarized in Table 5. Trees and shrubs, species occurring in 0-50 local floras from the southwest. though widely distributed, constitute only | ern United States. 14.2% of the total flora. Cacti and succulents | January 1986 MCLAUGHLIN: FLORISTIC ANALYSIS 53 TABLE 4. Most commonly encountered species in 50 local floras from the southwestern United States. a Subspecific % of Subspecific % of Species taxa floras Species taxa floras Artemisia ludoviciana 7 92 Astragalus lentiginosus 37 56 Sitanion hystrix 2 88 Plantago patagonica 3 56 Descurainia pinnata 12 86 Agropyron trachycaulum 4 56 Rhus trilcbata 6 82. Populus tremuloides 0 56 Erigeron divergens 2 76 Comandra umbellata 3 56 Atriplex canescens 3 76 Castilleja linearifolia 0 56 Aristida purpurea” 4 76 = Achillea millefolium 11 54 Gutierrezia sarothrae 2 72 Ceratoides lanata 2 54 Mentzelia albicaulis 2 72 Chenopodium fremontii 2 54 Poa fendleriana 3 70 Juncus arcticus”* 3 54 Mimulus guttatus 6 70 Phoradendron juniperinum 2 54 Vulpia octoflora 3 68 = Mirabilis multiflora 3 54 Salix exigua 0 68 Bromus carinatus” 0 54 Conyza canadensis 3 66 = Koeleria cristata 0 54 Linum lewisii 3 66 Ipomopsis aggregata - 4 54 Oenothera caespitosa 8 66 Amelanchier utahensis 2 54 Bouteloua curtipendula 2 66 Datura inoxia 0 54 Senecio douglasii 3 66 Celtis reticulata 0 54 Oryzopsis hymenoides 2 66 — Baccharis glutinosa 0 52 Holodiscus dumosus 2 66 Senecio multilobatus 0 52 Chrysothamnus nauseosus 20 64 Cryptantha pterocarya 4 52 Lappula redowskii 2 64 Erysimum capitatum 6 52 Sporobolus cryptandrus 0 64 Lepidium lasiocarpum 4 52 Opuntia phaeacantha 10 62 = Aristida adscensionis 3 52 Sporobolus airoides 0 62 Juncus ensifolius” 2 52 Stephanomeria pauciflora 2 62 Ranunculus cymbalaria 3 52 Echinocereus triglochidiatus 8 62 Nicotiana trigonophylla 0 52 Helianthus annuus 0 60 Chrysothamnus viscidiflorus 5 52 Leucelene ericoides 0 60 Pinus ponderosa 3 50 Erioneuron pulchellum 0 60 Pseudotsuga menziesii 2 50 Rosa woodsii 4 60 Gutierrezia microcephala 0 50 Cercocarpus montanus 7 58 Amaranthus blitoides 0 50 Populus fremontii 2 58 Artemisia tridentata 3 50 Erigeron pumilus 6 58 = Stipa comata 2 50 Draba cuneifolia 3 58 — Gilia sinuata 0 50 Allionia incarnata 0 58 Clematis ligusticifolia 3 50 Bouteloua gracilis 2 58 — Fallugia paradoxa 0 50 Castilleja chromosa 2 58 Salix gooddingii 0 50 Artemisia dracunculus 2 56 Physalis hederifolia 2 50 Brickellia californica 2 56 Verbena bracteata 0 50 Symphoricarpos oreophilius 2 56 7 5 7Nomenclature follows Kartesz and Kartesz (1980) except where otherwise footnoted. 2Number of subspecific taxa listed in Kartesz and Kartesz (1980). 3 sensu Cronquist et al. (1977). 4 sensu Hultén (1968). (mostly Agavaceae) make up 3.3% of the flora; annuals constitute 23.5%, a relatively high pro- portion. Perennial herbs, by far the largest group (58.9%), are the least widely distributed of the life forms listed in Table 5. Among herba- ceous plants, species of Poaceae are relatively widespread (5.9 floras/species). Many species of the large genera characteristic of the region are rather uncommon, e.g., Astragalus (2.0 floras/ species), Penstemon (2.2 floras/species), Cryp- tantha (3.1 floras/species), Erigeron (3.2 floras/ species), Phacelia (3.4 floras/species), and Gilia (3.8 floras/species). The number of southwestern species ex- tending into the seven peripheral regions are listed in Table 6. These numbers imply noth- ing about directions of migration; rather they show only the overlap in species composition between the Southwest and adjacent regions. A species occurring in the Southwest and an- other region(s) may be: primarily southwes- tern, barely entering the peripheral region(s); primarily distributed in the peripheral re- gion(s), barely entering the Southwest; or common in both the Southwest and one or more peripheral regions. The greatest overlap 54 GREAT BASIN NATURALIST TABLE 5. Summary of flora by life form. Life form No. species _ Floras/Species Trees 109 8.79 Shrubs 667 4.92 Perennial herbs 3,217 3.65 Annual herbs 1,285 4.24 Cacti and succulents 180 4.09 Total 5,458 4.06 TaBLE6. Species shared between Southwest and adja- cent regions. Letters refer to map locations in Figure 1. Species occurring in Southwest Peripheral region Number Percent of total A. Pacific region 1,544 28.3 B. Columbia Plateau region 1,224 22.4 C. Rocky Mountain region 1,479 folk D. Great Plains region 1,297 23.8 E. Sonoran region 847 15.5 F. Sierra Madre region 1,215 22.3 G. Chihuahuan region 1,119 20.5 occurs with the Pacific peripheral region, the least with the Sonoran Desert peripheral re- gion. Two types of species appear to account for much of the overlap: species of disturbed or weedy habitats, and species of relatively mesic habitats. Factor Analysis Solution In principal components analysis the n vari- ables are expressed in terms of n principal components (factors). By convention, only those factors with eigenvalues >1 (i.e., those factors accounting for at least 1/n of the varia- tion in the data) are retained for rotation to the final solution. Nine such factors were ex- tracted in this analysis. The 8th and 9th factors were bipolar with eigenvalues -1 and could not be simply interpreted. The Ist through 7th factors are depicted in Figures 3-9; they account for 55.4% of the variation in the data. The maps of Figures 3-9 were prepared by plotting the factor loadings for each flora and drawing contours or isolines to show areas of approximately equal factor loadings. Factor loadings are the correlations between the flo- ras and the factors. For this particular analy- sis, the factor loadings can be interpreted as similarity indices between the floras and the factors. The loadings for the peripheral re- gions are shown in the boxes on Figures 3-9. Factor analysis to date has not been used extensively in biogeography, despite its po- Vol. 46, No. 1 tential ability to simplify complex data sets. Apparently, a major reason for this is that both authors and readers often must engage in some rigorous mental gymnastics to interpret the “meaning’ of factors when there is consid- erable sampling error in the data or when patterns among the variables are in fact weak. Compared to most kinds of ecological data, there is relatively little error in determining which species occur in a particular area of limited extent. Ifthe results of the factor anal- ysis can be mapped, the interpretation should be straightforward—either there are simple, reasonable, and easily understood geographi- cal patterns associated with each factor or there are not. The factors mapped in Figures 3—9 are geo- graphically meaningful. Each shows high sim- ilarity with relatively few adjacent local floras. From these centers of high similarity, the magnitude of factor loadings declines more or less continuously in all directions. In form the mapped factors are reminiscent of the “noda” of Poore (1955). Each factor is most strongly represented in a single section of the study region, and these areas are fairly well delim- ited. The factors clearly identify what areas within the Southwest show higher than aver- age overlap in species composition between local floras. It seems to me that these seven factors can be interpreted readily as seven floristic elements in the traditional sense, that is, as clusters or centers of roughly coincident species ranges. Most of the seven floristic elements corre- spond to natural areas that have been recog- nized in the past on more subjective grounds. I will discuss each briefly before proceeding to a discussion of the conceptual and theoretical basis for recognizing floristic elements. 1. GREAT BASIN ELEMENT. —Factor | (Fig. 3) has high loadings with floras in the Great Basin section of the Basin and Range physiographic province. This element shows a distinct repre- sentation in the eastern portion of the Colorado Plateau as well as a recognizable extension into the Mogollon Rim area of central Arizona. The element declines in importance most rapidly to- ward the south in southern Nevada. Beatley (1975) discusses the topographic and climatic factors in this latter area that are correlated with the transition from Great Basin to Mojave Desert vegetation. January 1986 MCLAUGHLIN: FLORISTIC ANALYSIS 5D Figs. 3-6: 3, Plots of loadings for factor 1—Great Basin floristic element. 4, Plot of loadings for factor 2—Colorado Plateau floristic element. 5, Plot of loadings for factor 3—Mojavean floristic element. 6, Plot of loadings for factor 4—Chihuahuan floristic element. The Great Basin element is strongly corre- lated with the flora of the Columbia Plateau pe- ripheral region to the north. Both are regions of sagebrush desert and conifer-forested mountain ranges. The Great Basin element is also strongly correlated with the Rocky Mountain region to the east and the Pacific region to the west— primarily the Sierra Nevada portion—both of which have probably contributed species to the Great Basin. The factor loading with the Rocky Mountain region is the higher of the two, sug- gesting that the contribution from there may have been greater than that from the Sierra Ne- vada, as noted previously by Harper etal. (1978). 56 2. COLORADO PLATEAU ELEMENT.—The second factor (Fig. 4) is clearly associated with the Colorado Plateau. Its greatest representa- tion is in the northern portion of the Colorado Plateau about the confluence of the Green and Colorado rivers. The element extends beyond the limits of the Colorado Plateau into the southeastern Great Basin. Its most abrupt at- tenuation occurs along the southern boundary of the Colorado Plateau in the Mogollon Rim area. Unlike the Great Basin element, the Colo- rado Plateau element is not strongly corre- lated with any of the peripheral regions. The Colorado Plateau is also an area of relatively high endemism (Fig. 10). The high incidence of endemics and the low similarities with pe- ripheral regions identify the Colorado Plateau element as an autochthonous element. 3. MOJAVEAN ELEMENT. —The third factor (Fig. 5) is centered about the Mojave Desert region of southeastern California, northwest- ern Arizona, and southern Nevada. The Mo- javean element, like the Colorado Plateau ele- ment, is weakly correlated with all peripheral regions, including the Sonoran Desert region. The rapid attenuation of the element to the west, where it meets the California floristic province, is notable. The Mojave Desert re- gion has the highest frequency of endemic species in the Southwest (Fig. 10). This ele- ment is clearly autochthonous. 4. CHIHUAHUAN ELEMENT.—The fourth factor (Fig. 6) is identifiable as a Chihuahuan element, located in the Chihuahuan Desert area of Trans-Pecos Texas in the extreme southeastern portion of the study region. It extends up the Rio Grande Valley and into southeastern Arizona. In the latter region spe- cies with Chihuahuan affinities are frequently associated with limestone substrates (Whit- taker and Niering 1968, Wentworth 1982). The Chihuahuan element is strongly corre- lated with the flora of the Chihuahuan periph- eral region and is an extension of the flora of that region. The Chihuahuan element is also moderately correlated with the floras of the Plains and Madrean peripheral regions. 5. SONORAN ELEMENT.—The fifth factor (Fig. 7) is concentrated in southwestern Ari- zona and the Colorado Desert area of extreme southeastern California. It is highly correlated with the flora of the Sonoran Desert periph- GREAT BASIN NATURALIST Vol. 46, No. 1 eral region and weakly correlated with all other peripheral regions. The four desert elements recognized— Great Basin, Mojavean, Chihuahuan, and Sonoran—correspond very closely to the four desert regions of the same names recognized by Shreve (1942). In fact, the formal analysis presented here nicely confirms Shreve’s im- pressions based on vegetation and flora. More recent interpretations that would include the Mojave Desert as a subdivision of the Sonoran Desert (Cronquist 1982; Turner 1982) are not supported. Floristic evidence favors retention of the Mojave Desert as a biogeographic en- tity distinct from the Sonoran Desert to the south. 6. APACHIAN ELEMENT. Factor 6 (Fig. 8) occurs primarily in southeastern Arizona and southwestern New Mexico. It can be termed an Apachian element in the sense of Dice (1943). The Apachian element extends all along the southern boundary of the Colorado Plateau along the Mogollon Rim into north- western Arizona. An extension can easily be recognized in Trans-Pecos Texas and south- ern New Mexico east of the Rio Grande. It is not clear from an analysis of floras from north of the International Border whether this element is largely autochthonous or al- lochthonous. The element could be centered within the mountainous region of southeast- ern Arizona, southwestern new Mexico, and northeastern Sonora, the Apachian Biotic Province of Dice (1943), or it could be a north- ern extension of a broader floristic element centered in the Sierra Madre Occidental of southeastern Sonora, southwestern Chi- huahua, and northwestern Durango, the Sierra Madre Occidental Biotic Province of Goldman and Moore (1945). Two local floras from northwestern Mexico provide additional data. White (1948) prepared a large flora (927 native, terrestrial species) for the Rio de Bavispe region of extreme northeastern Sonora, including the Sierra del Tigre, often considered an outlying section of the Sierra Madre. Maysilles (1959) examined the flora (506 native terrestrial species) of the pine forests of western Durango in the central Sierra Madre. I have calculated the similari- ties between these two Mexican floras and five floras with high loadings on factor 6 (Table 7). The Bavispe flora shows high similarity with January 1986 MCLAUGHLIN: FLORISTIC ANALYSIS O77 Figure 8 Figure /0 Figs. 7-10: 7, Plot of loadings for factor 5—Sonoran floristic element. 8, Plot of loadings for factor 6—Apachian floristic element. 9, Plot of loadings for factor 7—Mogollon floristic element. 10, Distribution of species endemic to the southwestern United States. Isolines are labeled with the proportion of species in local floras occurring only in the study region. all five floras in Table 7; the Western Durango flora shows low similarity with the same five floras. The Bavispe flora, in fact, shows signifi- cantly greater similarity with the Skull Valley flora at the western end of the Mogollon Rim region than it does with the Western Durango flora. These comparisons support an inter- pretation of factor 6 as an autochthonous ele- ment from the southwestern USA and north- eastern Sonora. The Sierra Madre Occiden- tal, like the Sierra Nevada and Rocky Mountains, might best be considered a floris- tic region separate from the Southwest re- gion of this study. More floristic research in 58 GREAT BASIN NATURALIST Vol. 46, No. 1 TABLE 7. Comparison of selected southwestern floras with two floras from the Sierra Madre region of northwestern Mexico. Flora Otsuka Similarity Index Bavispe Region Western Durango 21 Skull Valley Planning Unit 24 Sierra Ancha Experimental Forest 26 Saguaro National Monument 28 Chiricahua National Monument 29 Animas Mountains Western Durango .262 .046 349 118 .459 141 .459 .145 445 .136 172 — northern Mexico is required to clarify rela- tionships in this area. 7. MOGOLLON ELEMENT. —The last factor (Fig. 9) is centered in the mountainous region of central New Mexico, extending into Ari- zona along the Mogollon Rim, across the Kaibab Plateau, and onto the central Wasatch Plateau of southcentral Utah. This element is moderately correlated with the Rocky Moun- tain and Plains peripheral regions. The simi- larity with the Rocky Mountains region is largely due to species occurring in the south- ern Rocky Mountains. I have applied the des- ignation Mogollon’” for this element because it centered about the Mogollon Rim of Ari- zona and the Mogollon Mountains of New Mexico. Axelrod and Raven (1985) have recently re- viewed the history of the Cordilleran Floristic Province, which they define as the Great Basin, Colorado Plateau, and Southern Rocky Mountains. They include the Mogollon area of this analysis in a Madrean Floristic Province. My analysis shows that the Mogol- lon area is more closely related to the Rocky Mountain region than to the Madrean region. Furthermore, the Great Basin area seems to have closer affinities with the northern Rocky Mountains than with the southern Rocky Mountains and Mogollon region. An analysis of a series of local floras situated throughout the Rocky Mountain region would be most helpful in resolving floristic relationships in the Cordilleran Region. The maps of floristic elements can be com- bined to form a single map of floristic areas of the Southwest (Fig. 11), made simply by drawing lines around those floras sharing their highest loading with a particular factor. The position of a boundary between adjacent floras that lie in different floristic areas is deter- mined by the relative magnitudes of the load- ings between both floras and both factors. The map of floristic areas (Fig. 11) does not substi- tute for the maps of floristic elements. Like political units, floristic areas are two-dimen- sional whereas floristic elements, like repre- sentations of rainfall, temperature, or topog- raphy, are essentially three-dimensional. There is a one to one relationship between floristic elements and floristic areas in this analysis, but floristic elements are by no means confined to particular floristic areas. The floristic areas recognized in this study are in close agreement with those of Dice (1943) for the Southwest. The only significant difference is that both my Colorado Plateau and Mogollon floristic areas would be in- cluded in Dice’s Navahonian Biotic Province. The more recent treatment by Cronquist (1982) oversimplifies floristic relationships in the Southwest. Reliability of the Analysis Factor analysis is a nonhierarchical cluster- ing technique that will produce clusters or factors from any data matrix. It is legitimate and important to question how accurately the factors depict the actual relationships among the variables. Does it produce the wrong clus- ters, or does it produce clusters when in fact none exist? Unfortunately there are no simple statisti- cal tests to answer these questions. I think there are two criteria that can be applied to factor analysis solutions. The first is that of interpretability. Stated simply, do the factors make sense? I have shown in the preceding section that the factors can be mapped di- rectly to reveal their geographic meaning and that the particular factors arrived at corre- spond closely to floristic elements and areas in January 1986 Factor | Great Basin Floristic Area % svt Nt@ Mojavegn eT EN Floristic . t ) . Area, ao Factor 5 e\Sonoran® Floristic Area Floristic ®Area } Factor 6\ a Aachen MCLAUGHLIN: FLORISTIC ANALYSIS 59 Mogelan: ; Factory @ Floristic Area |! 4@ S l Ne oN Chihuahuan Floristic Area Fig. 11. Floristic areas of the southwestern United States identified from a factor analysis of similarity coefficients among 50 local floras. the Southwest that have been recognized for at least 40 years. The factors found in this analysis are simply and sensibly inter- pretable. The second criterion is that of consistency. If relationships among the variables are weak or if there are in fact no correlated clusters of variables, small changes in the factoring pro- cedures or in the data are likely to result in substantial changes in the final solution. To check the consistency of the factor analysis solution, I made several changes in the data analysis and compared the results. Changes included using oblique rotation instead of or- thogonal rotation, substituting Simpson’s in- dex of similarity (Simpson 1960) for Otsuka's, 60 and omitting the checklists for the seven pe- ripheral regions from the data base. Each of the alternative analysis produced a set of seven unipolar factors nearly identical to those depicted in Figures 3-9. The factor analysis solution does appear to satisfy the criterion of consistency. Simpson's index did appear to be superior to Otsuka’s index for use in this type of analy- sis. The communalities, defined as the sum of the squared factor loadings for each variable, measure the proportion of variance of each variable accounted for by the final factor anal- ysis solution. With Otsuka’s index the com- munalities were correlated with the size of the flora (r = 0.43, p < .01); with Simpson's index the communalities and flora sizes were not correlated (r = 0.27, p > .05). This means that the unexplained or residual variances were associated with the smaller floras using Otsuka’s index. Intuitively it seems more rea- sonable either for the residual variance to be associated with the larger floras, or for it to be uncorrelated with flora size. Also, the final solution accounts for a larger proportion of the total variance (68.7%) when the similarities are expressed as Simpsons indices. DISCUSSION The flora of the Southwest can be described in terms of seven floristic elements. These elements can be objectively recognized and explicitly mapped. The derived maps (Figs. 3-9) show, however, not discrete units with clearly delineated boundaries, but rather geo- graphic features varying in intensity from a central area of greatest development and di- minishing gradually outward. Each element is most clearly developed in a limited area that can be characterized by topographic and cli- matic features that vary from adjacent areas dominated by other elements. Five of these areas—Great Basin, Colorado Plateau, Mo- jave Desert, Sonoran Desert, and Chihua- huan Desert—are widely recognized natural geographic areas. The other two—Apachian floristic area and Mogollon floristic area—are readily associated with upland regions sepa- rating adjacent lowland desert areas. Floristic elements are objectively defined assemblages, but they are not discrete, bounded units in the sense of Clementsian GREAT BASIN NATURALIST Vol. 46, No. 1 formations or plant association types. Each of the 50 local floras used in the analysis contains representative species of several elements; likewise the species and elemental composi- tion of the floras change in a continuous man- ner along geographic gradients. The situation is analogous to weather fronts and the particu- lar state of the atmosphere at any particular time—the existence of continuous gradients in air pressure across a region is not contradic- tory to the recognition of high and low pres- sure cells in different areas of the region. Somewhat surprisingly the analyses failed to recognize widespread elements. That is, there were no factors that could be associated with extensive groups such as “northern,” “western, “boreal,” “temperate,” or “south- western’ areal types (Whittaker and Niering 1964). There certainly are widespread spe- cies, i.e., those of Table 4. Such species do constitute a significant fraction of any particu- lar local flora, but they constitute a small frac- tion of the regional flora. The average extent of a floristic area in this analysis includes 7 floras, but the average range of a species in- cludes only 4 floras. In fact, only 17% of the species of the region occur in 8 or more floras, beyond the average range of one floristic area, and only 6% occur in 15 or more floras, be- yond the average range of two floristic areas. In many instances one could probably associ- ate many of the subspecific taxa of the rela- tively few widespread species with specific floristic elements. No widespread elements were recognized because the factors reflect general trends among the ranges of the major- ity of species, which are not widespread. The recognition of floristic elements, i.e., coincident patterns in species ranges, is not necessarily inconsistent with the individualis- tic concept of species distributions. The latter view states that no two species distributions are identical since each species has individual physiological tolerances and evolutionary his- tory. Indeed, very few of the over 5,000 spe- cies occurring in the Southwest show an exact correlation of presence and absence in the local floras. The existence of floristic elements indicates that species have nonrandom, over- lapping patterns of area, not that they are found consistently in similar communities, which is what the individualistic concept ad- dresses. Many southwestern species are so January 1986 uncommon that they may consistently occur together in similar areas, i.e., are members of the same local floras yet seldom or never occur together in the same community. The problem of scale or scope is thus seen to be crucial in assessing the validity of associa- tional concepts such as elements or assem- blages of species. Whittaker (1970) presents graphs of overlapping, normal curves repre- senting the abundances of dominant, woody species in local areas to argue against the exis- tence of vegetation association types. The quadrats or plots of the ecologist, however, contain only a minute fraction of a regional flora. As one expands the sample area from plots to local floras to floristic areas, the “grain in species composition becomes finer and patterns emerge. At the scale of conti- nents, I doubt that any biogeographer fails to recognize the existence of distinctive biotas. The best explanation for groups existing at regional levels while continuity prevails at lo- cal levels involves barriers to migration. Such barriers are largely lacking for terrestrial plants at the level of quadrats and plots, but they become more common and more effec- tive as one increases scale. At the level of continents oceans provide very effective bar- riers to migration; at the level of floristic areas topography and climate provide sufficiently effective barriers to promote regional differ- entiation of the flora into floristic elements. Important barriers in the Southwest that serve to segregate one or more floristic ele- ments include the Mogollon Rim along the southern boundary of the Colorado Plateau, the Wasatch Plateau in central Utah, the high- lands of southeastern Arizona and southwest- ern New Mexico (including the rather low-ly- ing continental divide in this area), and the broad zone of increasing basal elevation in southern Nevada. Migration has been depicted in the litera- ture as a process that should effectively pre- clude the formation or recognition of floristic elements. Gleason (1926) argued that both individualistic tolerances and extensive mi- grations of species were inconsistent with the recognition of recurring associations of spe- cies. Good (1931) and Mason (1936) elevated to a “principle” the assertion that extensive | migrations of floras had occurred in the past. Gleason and Cronquist (1964) stated that most MCLAUGHLIN: FLORISTIC ANALYSIS 61 plant species are capable of migrating to all suit- able habitats and that such migrations are suffi- ciently rapid to keep pace with environmental changes. How then can floristic elements be maintained in the face of such extensive migra- tions? The fact is that observed patterns of plant dis- tributions in the Southwest do not provide much evidence of extensive migration as a general phenomenon. the uncommonness of the major- ity of species indicates either that most plants are rather poor migrators or that insufficient time has elapsed for these species to complete their migrations. Both are probably true for south- western species. Two observations seem to support the view that plants can and do migrate to nearly any- where within the range of their physiological tolerances. First, many (but not all) seeds and fruits do possess obvious adaptations for disper- sal. Second, the fossil record provides concrete evidence that some species have migrated widely. Nevertheless, from these observations it does not follow that all or most plant species actually have migrated extensively. Migration is a matter of probabilities. The vast majority of seeds dispersed by plants travel very short distances (Levin and Kerster 1974). The occasional long-distance transport of a single seed or fruit is unlikely to result in successful colonization within a stable, competitive com- munity. Most importantly, the likelihood of suc- cessful migration is dependent on the number of seeds of the species dispersed per unit of time compared to other potential migrators. The ma- jority of plant species in the Southwest occur in three or fewer local floras and may simply not be abundant enough to generate a flux of seed suffi- ciently great to make extensive migration likely, even over considerable spans of time. Most of the actual examples of extensive mi- gration in the fossil record involve dominant, usually woody species. These plants constitute a small fraction of the southwestern regional flora. Derivation of general principles for all species of plants from data on the current and former distri- butions of species such as redwoods, sugar maples, and other forest trees is a questionable practice that may have led to some unwarranted conclusions. If the capacity of most species to migrate to all areas suited to their physiological tolerances were as pervasive as typically assumed, I would 62 expect the average range of a species to be much wider. One could argue that species in the Southwest have very narrow physiological tolerances that effectively restrict them to rel- atively few habitats within a limited area, but that seems rather unlikely given the similar nutritional needs of plants and the apparent lack of coincidence between most species and very specific habitats. It seems much more likely that barriers to migration, competition, and low rates of seed dispersal interact to greatly curtail the ability of most species to migrate. Over the past several years a general model of southwestern phytogeographic history has emerged that includes several postulates: (1) most species present today had evolved by the end of the Tertiary, (2) these species have undergone extensive and intensive migra- tions in response to Pleistocene climate changes, and (3) modern plant distributions reflect both this migrational history and the tolerances of species for modern climatic con- ditions. The concept of great species longevity is supported by Levin and Wilson (1976), who estimated the mean duration for herbaceous species to be 10 million years, with a mean speciation rate of 1.15 lineages per million years. Woody species were estimated to be even more persistent and to have even slower rates of speciation. Consistent with this view, Tidwell et al. (1972) hypothesized that the flora at the beginning of the Pleistocene in the Intermountain Region was basically the same as exists today. The best evidence that many dominant spe- cies have indeed undergone extensive migra- tions in the Southwest at the end of the Pleis- tocene comes from fossil packrat middens (Van Devender and Spaulding 1979, Wells 1983). Many workers have textraploted these results back to previous glacials and inter- glacials, concluding that the repeated climate changes occurring during the Pleistocene in topographically heterogeneous intermoun- tain environments would have kept species populations in constant movement (Cronquist 1978). It is difficult to conceive how recognizable floristic assemblages could survive through the Pleistocene with all this movement, fluc- tuation, and shuffling of populations in re- GREAT BASIN NATURALIST Vol. 46, No. 1 sponse to climate change. My analysis of mod- ern plant distributions in the entire regional flora, however, casts some doubt on the gen- erality of the consensus model outlined above. Most of the local floras included in this analysis are from areas encompassing exten- sive elevational relief. Southwestern moun- tain ranges, particularly in the Basin and Range Province, possess considerable habitat diversity in association with elevation, typi- cally going from desert at the base to pinyon- juniper woodland, pine forest, or even spruce forest at the top (e.g., Lowe 1964). Old spe- cies with extensive migrational histories would have left relictual populations scattered within and between these ranges. Only a sin- gle, small population is required to establish a species as a member of a local flora. The con- sensus model would predict that the ranges of most species would include many such scat- tered, relictual populations, resulting in a modern flora with most species widely dis- tributed throughout the region and little dif- ferentiation of the floristic assemblages within the region. Many species, particularly those of high el- evations, do show distribution patterns con- sistent with the consensus model, but they do not constitute a large fraction of the Southwest flora. For the rare species consisting of small populations in only a few floras, i.e., the ma- jority of species in the Southwest today, ex- tinction, rather than migration, is the more likely response to severe climate change. These species are unlikely to possess suffi- cient ecotypic, genetic diversity or suffi- ciently large populations required to keep pace with the changing environment. Speciation, however, may also be acceler- ated during periods of severe climate change (Stebbins 1947, Axelrod 1981). Many south- western species probably originated in post- glacial time in response to the new, more arid environments created by the changes from Pleistocene to Holocene climates. Such an interpretation has been applied to Atriplex (Stutz 1978) and seems likely for many of the species of large southwestern genera such as Eriogonum, Astragalus, Penstemon, Gilia, Camissonia, Cryptantha, Phacelia, Opuntia, and others. Do we need two models of evolutionary | response to climate change, one for common ee = = _ January 1986 species and one for rare species? I think not. Common plant species consist of many popu- lations of subspecies, varieties, or ecotypes. During periods of severe climate change, some of these ecotypes will migrate, many will go extinct, and some will differentiate to form new ecotypes, varieties, subspecies, and species in new habitats. As a result, the range of the species before the climate change (i.e., in the fossil record) will be different from its range after the climate change. But migration of populations or ecotypes existing before the climate change and persisting through the cli- mate change need account for only part of the total change in the range of a species. In other words, the most likely response of any partic- ular small population to rapid environmental change is extinction; the probability of a spe- cies becoming extinct depends in large part on how many populations it has. The seven floristic elements identified in the present analysis are clearly associated with modern, Holocene environments. They represent centers of postglacial differentiation and speciation within the depleted flora sur- viving at the end of the Pleistocene in the Southwest, partially but effectively isolated from adjacent elements by modern climatic and topographic barriers. My analysis of specific similarities among local floras of the Southwest is consistent with a hypothesis of Holocene differentiation of the modern regional flora. An analysis of generic similarities among these floras might better reflect the Tertiary and Pleistocene migra- tional history of the flora, since the majority of the genera are doubtless of Tertiary age. My interpretation of the evolution of the south- western flora is also consistent with the con- cept of punctuated equilibria, as opposed to slow, gradual phylogenetic change, which would be more consistent with the consensus model discussed above. American plant ecology has been very much a science of dominants. This bias has been carried over into plant geography, which has emphasized the distribution of vegetation almost to the exclusion of flora. I have tried to show in this paper how a study of the entire flora of a region, weighing all species equally, can reveal patterns that may not follow from or be in agreement with principles and models derived solely from study of the dominant MCLAUGHLIN: FLORISTIC ANALYSIS 63 species. Floristics and vegetation analysis are complementary approaches to understanding the distribution and abundance of plants. Phytogeography needs both. ACKNOWLEDGMENTS I thank R. M. Turner, J. H. Brown, J. E. Bowers, and M. A. Kurzius for reading and commenting on_ the manuscript. M. Beauchamp, J. E. Bowers, D. Clemons, R. A. Fletcher, T. R. Van Devender, and R. D. Worthington helped in the acquisition of flo- ras and unpublished checklists. The illustra- tions were prepared by M. A. Kurzius. LITERATURE CITED AXELROD, D. I. 1981. Holocene climatic changes in rela- tion to vegetation disjunction and speciation. Amer. Nat. 117:847-870. AXELROD, D. I., AND P. H. RAVEN. 1985. Origins of the Cordilleran flora. 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The vegetation and flora of the region of the Rio de Bavispe in northeastern Sonora, Mexico. Lloydia 11:229-302. WHITTAKER, R. H. 1970. Communities and ecosystems. Macmillan Co., New York. WHITTAKER, R. H., AND W. A. NIERING. 1964. Vegetation of the Santa Catalina Mountains, Arizona. I. Eco- logical classification and distribution of species. J. Arizona Acad. Sci. 3:9-34. WHITTAKER, R. H., AND W. A. NIERING. 1968. Vegetation of the Santa Catalina Mountains, Arizona. IV. Limestone and acid soils. J. Ecol. 56:523-544. WorTHINGTON, R. D. 1980. Flora of Hueco Tanks State Historical Park. Unpublished rept., Texas Parks and Wildl. Dep. UTAH FLORA: APIACEAE (UMBELLIFERAE) Sherel Goodrich! ApsTRACT.—Eighty-four taxa in 30 genera of the parsley family, Apiaceae (Umbelliferae) are treated for Utah. Four of the genera with one species each that escape from cultivation are included in the key but not in the text. Keys to genera, species and infraspecific taxa are provided, along with detailed descriptions, distributional data, and pertinent comments. Proposed new taxa are Cymopterus acaulis (Pursh) Raf. var. parvus Goodrich and Lomatium scabrum (Coult. & Rose) Mathias var. tripinnatus Goodrich. New combinations include: Cymopterus acaulis (Pursh) Raf. var. fendleri (Gray) Goodrich and var. higginsii (Welsh) Goodrich; Cymopterus purpureus Wats. var. jonesii (Coult. & Rose) Goodrich and var. rosei (Jones) Goodrich; Cymopterus terebinthinus (Hook.).T. & G. var. petraeus (Jones) Goodrich. This paper is another in a series leading to a definitive treatment of the flora of Utah. Pre- vious papers have dealt with the Brassicaceae (Cruciferae), Fabaceae (Leguminosae), Rosa- ceae, Asteraceae (Compositae), Salicaceae, Cactaceae, Chenopodiaceae, and miscella- neous smaller families. The parsley family presents a formidable challenge to students of plant taxonomy and to others who need to identify its members. The family itself is well marked, but some generic lines within it are fraught with difficulty. It is interwoven with look-alikes often of different and sometimes distantly related genera. Flo- ral structures are reduced and uniform. The calyx is lacking or reduced to mere teeth. Petals are only about 1-2 mm long in our taxa except in Heracleum. Color variation of petals is basically restricted to yellow and white, and occasionally purple. Yellow petals very often turn white or cream when dried, and a num- ber of taxa described from dried specimens as having white petals have been proven by field studies to have fresh petals that are yellow. The reduction and uniformity of floral parts requires the use of fruiting and vegetative features for separation of taxa. The features of the mature fruit are quite diagnostic, but this does not help in identification of specimens collected in flowering condition. Vegetative features are used extensively in the keys of this work, but these features are extremely variable. The size of the family (30 genera and 84 taxa in Utah and about 300 genera and ‘USDA Forest Service, Intermountain Research Station, Forest Service, U Ranger District, Ashley National Forest, Vernal, Utah 84078. 66 about 3,000 species worldwide) also contributes to difficulty in identification. Among those of the Old World introduced, cultivated members of the family are caraway (Carum carvi), carrot (Daucus carota ssp. sa- tivus), celery (Apium graveolens), dill (Anethum graveolens), ground elder (Aegopodium poda- graria, parsley (Petroselinum crispum), and parsnip (Pastinaca sativa ssp. sylvestris). Other introductions from the Old World include: hedge parsley (Torilis arvensis), poison hemlock (Conium maculatum), sweet fennel (Foeniculum vulgare), wild carrot (Daucus carota ssp. carota), and wild parsnip (Pastinaca sativa ssp. sativa). The toxic nature of poison hemlock has long been known, and water hemlock (Cicuta maculata) is probably the most violently poi- sonous plant of our native flora. Many other members of the family such as western sweet cicely (Osmorhiza occidentalis ) and spring pars- ley (Cymopterus spp.) are used extensively by livestock and wildlife without apparent harm. Measurements of rays, pedicels, and fruit were taken from specimens with mature fruits. At the end of the discussion of each taxon there are two numbers. The first, in Arabic numerals, indicates the number of Utah specimens exam- ined for the taxon. The second, in Roman nu- merals, indicates the number collected in Utah by the author. ACKNOWLEDGMENTS Appreciation is expressed to Dr. Arthur Cronquist for permitting me to examine his . §. Department of Agriculture, Ogden, Utah 84401. Present address: Vernal Perr : January 1986 Apiaceae manuscript for the Intermountain Flora. His observations of the carpophore in Cymopterus have been especially helpful. Beverly Albee helped with distribution maps for all species. These maps were helpful to understanding possible relationships of taxa as well as distributions. Appreciation is also ex- pressed to the directors and curators of the following herbaria of the state: Brigham Young University, Provo; Forest Service Herbarium, Ogden; Garrett Herbarium, The University of Utah, Salt Lake City; Inter- mountain Herbarium, Utah State University, Logan. I appreciate the loans from each of these herbaria. These specimens are the basis of this work. During the course of this project, numerous specimens have been sent to Dr. Lincoln Constance (a student of the family for more than 40 years) for annotation. His prompt and congenial replies are greatly ap- preciated. The concepts of several taxa are based on his annotations. This work is dedi- cated to him. APIACEAE (UMBELLIFERAE) Parsley Family Annual, biennial, or perennial acaulescent or caulescent herbaceous plants from tap- roots, rhizomes, fibrous or tuberous roots, or caudices; leaves simple to decompound, peti- oles often dilated and partly sheathing at the base or the upper leaves reduced to dilated sheaths; inflorescence mostly of compound umbels, the primary umbels with or without a subtending involucre of bracts, the secondary umbels (umbellets) with or without a subtend- ing involucel of bractlets; flowers mostly regu- lar, perfect, or some of them staminate or sterile; calyx of 5 teeth or lobes, or obsolete, or lacking; petals 5, small, usually inflexed at the tip, mostly white or yellow, occasionally purple; stamens 5, small, alternate with the petals; ovary inferior, bicarpellate, 2- loculed, with 1 ovule per locule, the two styles with or without a conical base (stylopodium); fruit a dry schizocarp of 2 mericarps united by their faces (the commmissure) nearly terete, dorsally compressed (compression parallel to a broad commissure), or laterally compressed (compression, contrary to a narrow commis- sure); mericarps separating at maturity and apically attached to and pendulous from a fine GOODRICH: UTAH FLORA, APIACEAE 67 wirelike entire or bifid to divided carpophore or remaining adherent and then the car- pophore usually lacking or poorly developed and usually adnate to the commissural faces, each mericarp usually 5—nerved, 3 of the nerves dorsal and 2 on the lateral margins, the nerves filiform to winged, or obscure or lack- ing, the intervals between the nerves com- monly with 1 or more oil-tubes, the commis- sural faces often with 2 or more oil-tubes. REFERENCES ARNOW, L., B. ALBEE, AND A. WYCOFF. 1980. Flora of the central Wasatch Front, Utah. Univ. of Utah Print- ing Service, Salt Lake City. 663 pp. HitcHcock, C. L., AND A. CRONQUIST. 1961. Vascular plants of the Pacific Northwest. Part 3: Sax- ifragaceae to Ericaceae. Univ. Washington Publ. Biol. Vol. 17. 614 pp. MaTulias, M. E., AND L. CONSTANCE. 1944—1945. Umbel- liferae. N. Am. Fl. 28B:43—297. il Leaves peltate, simple, orbicular; flowers in a verticellate spikelike inflorescence; plants rhizomatous, of Washington County Hydrocotyle — Leaves not as above; flowers in compound umbels, or globose heads (in a few taxa of Cymopterus ); plants rarely rhizomatous ..... 2 Plants caulescent; pseudoscape lacking; the few to several peduncles mostly shorter than the leafy stem on which they are borne; styles rarely over 1 mm long; stylopodium present and petals white in native taxa except Zizia and in a few taxa keyed both ways — Plants acaulescent, the leaves sometimes whorled atop a pseudoscape, or if subcaules- cent then the usually solitary peduncle longer than the short, leafy stem on which it is borne, and lateral umbels if any mostly borne on the lower 1/3 of the plant; styles often over 1 mm long; stylopodium lacking in all but Podisteria and in taxa keyed both ways; petals yellow, WHIM, OPUS BG acescesoodosobooudduds 4 Leaves simple, pinnate or ternate; leaflets NOSE SSIS, ooponcapadsvqoecce don Key I — Primary leaflets usually petiolulate Leaves ternate or biternate with 3-9 leaf- lets or rarely a few simple, usually only 2-3 per plant; leaflets 1.5—-7 (11.5) em long, entire, linear or nearly so; plants 5-10 cm tall, from a globose or fusiform tuber; petals SEIS) Shs och aia Dim oem ideo auc Orogenia Leaves and leaflets not as above or if so then plants mostly taller and/or petals yellow ..... 5 68 Nt S GREAT BASIN NATURALIST Stylopodium low conic; plants of the Raft River and Uinta mountains, mostly above 2,440 m, sometimes with a lateral umbel in the axil ofa bract or leaf on the upper 1/2 of the stem, glabrous; involucels lacking or of 1-2 linear bractlets; fruit 3-6 mm long; petals White rcskeyonce sewers the asic haaiae poe Ligusticum Stylopodium lacking except in Podistera; plants not as above in all features; involucels mostly present; fruit mostly longer or petals VElOWee Sitsit oes ashen 0 one ears Gitar 6 Key III Key to plants in flower or with young fruits Sa eacehc oes mame ea siete Ceara Alternate Key III Key to plants with mature fruits....... Key I Plants caulescent; the few to several pedun- cles and umbels mostly shorter than the leafy stem; stylopodium present in all but Zizia and two other taxa that are keyed both ways; leaves simple, pinnate, or ternate; leaflets ses- sile. Leaflets entire, linear or linear-elliptic ...... 2 Leaflets toothed and/or lobed, not linear .... 4 Leaves soon withering, some usually decidu- ous shortly after anthesis, some leaflets often more than 1.5 cm long; plants from a tubrous root or fascicle of tuberous roots, the stem readily detached from the tubrous base, from northern Utah; stylopodium present Re Rr tn nepal eee ae Perideridia Leaves more persistent, the leaflets not more than 1.5 cm long or plants of San Juan and Wayne counties; plants from a taproot with a branched crown or caudex; stylopodium lack- Petals and stamens yellow when fresh; leaflets mostly 2—5 cm long; fruit 6—8 mm long; plants of San Juan and Wayne counties er eee bier eRe ty tet | aOR RS Cymopterus beckii Petals and stamens white; leaflets 0.3-2 cm long; fruit 2-4 mm long; plants of Cache Countymr Sere hee Musineon lineare Basal leaves mostly simple, shallowly toothed, cordate at the base; stem leaves usu- ally ternate, not over 3 cm long; petals bright Vie lows rae w ie on ano ot Zizia Leaves pinnate, or if ternate or upper ones simple then over 3 cm long; petals white or VellOWarian ore ree ting einere yee coe comes 5 Leaves ternate, the upper ones sometimes simple, the 3 leaflets 8-36 cm long, about as wide; plants 1-2 m tall or taller, villous- woolly at least on some of the nodes; the larger petals 4—-8.5mmlong............ Heracleum Leaves pinnate, the leaflets less than 8 cm long and much narrower; plants shorter or not villous-woolly; petals smaller .............. 6 6(5). 10(9). Vol. 46, No. 1 Umbels sessile or nearly so; leaflets ovate to suborbicular, 3—lobed to near the middle; fruit about 1.5 mm long; plants cultivated and rarely escaping except in Washington County (celery) hard ee: Apium graveolens L. Umbels not sessile except sometimes the ter- minal one; leaflets variously shaped, but not ovate to suborbicular and lobed to near the middle; fruit more than 1.5 mm long except in Berula, plants varlous= ae eee eee iT Involucre and involucels well developed, sometimes spreading or deflexed, the bracts 1-6, the bractlets (2) 4-12; fruit 1.5-3 mm long, the ribs not winged; plants of very wet places, often growing in water, from fibrous roots Involucre lacking or infrequently of 1 or 2 bracts; involucels often lacking; fruit over 3 mm long or else the ribs winged; plants of various habitats, from a taproot or tuberous roots Stems often sprawling, sometimes stolonifer- ous; leaves with (3) 5-15 opposite pairs of leaflets, these 0.3—4 (6.5) cm long; rays 4-16; ribs of the fruit obscur................ Berula Stems erect, not stoloniferous; leaves with 4—6 opposite pairs of leaflets, these 2—8 (15) cm long; rays 11-24; ribs of the fruit promi- nently corky ........... osm eee Sium Umbels often more than 7 per stem; fruit strongly flattened dorsally, 5-8 mm long, 3-6 mm wide, the lateral ribs slightly winged, the dorsal ones filiform; petals greenish yellow or reddish; plants introduced, cultivated and Wild ).3 he) AEE eee Pastinaca Umbels mostly less than 7 per stem; fruit not strongly flattened dorsally or if so only 3-5 mm long; petals white or greenish; plants na- TIVE: cb ave) nea pita baie EC eee 10 Fruit over 1 cm long; leaves rarely all pinnate; peduncles mostly not subtended by dilated, bladeless sheaths or these greatly reduced Osmorhia Fruit 3-5 mm long, leaves mostly all once pinnate; peduncles often with subtending di- latedisheaths :..: 3... J 0 ae ee eee 11 11(10). Fruit strongly flattened, the dorsal ribs fili- form, the lateral ribs conspicuously winged; plants from fascicles of tuberous roots, of the Abajo and La Sal mountains ......... Oxypolis Fruit rounded in cross section, the dorsal and lateral ribs with small wings; plants from a taproot, widespread Angelica Key II Plants caulescent; the few to several pedun- cles and umbels mostly shorter than the leafy stems; stylopodium present or plants also keyed in Key III; leaves more than once-com- pound; primary leaflets not sessile. January 1986 1. 5(1). At least some of the ultimate leaf segments over 2 cm long, toothed or lobed, but not entire or pinnatifid ...................... 2 Ultimate leaf segments less than 2 cm long or if longer then entire or pinnatifid........... 5 Plants from creeping rhizomes, cultivated and rarely escaping; lower leaves long-peti- oled, often biternate with 9 leaflets but some- times irregularly compound (ground elder) Aegopodium podagraria L. Plants not from creeping rhizomes, seldom cultivated; leaves various Involucels of about 6 bractlets, 1-4 mm long; umbels 6-20 or more per stem, the rays 15-26, 1.5-4 cm long; fruit 2-4 mm long, the TAD SCOLKY geben oigeewets aie encase Sencar. Cicuta Involucels mostly lacking; umbels often fewer than 6 per stem and/or the rays either fewer or longer than above or both; fruit 4-25 mm long, themibsivanious =~ .sas5* 505200050656 4 Fruit (10) 12-25 mm long, bristly pubescent in 2 of 3 species, the dorsal ribs not promi- nent; leaflets often hirtellous; dilated sheaths seldom subtending the peduncles .. Osmorhiza Fruit 4-5 mm long, not bristly pubescent, the dorsal ribs with small wings; leaflets glabrous; peduncles often subtended by dilated blade- less (or nearly so) sheaths Angelica Fruits and ovaries with bristly hairs; involucre often of pinnatifid or compound bracts; plants annual or biennial Fruits and ovaries without bristly hairs; in- volucre mostly of entire bracts; plants mostly bienniallorperenniall race na 8 Involucre lacking or of 1 entire bract; plants with appressed hispid hairs, from Washington (County ee Oe rear one Torilis Involucre of few to several pinnatifid to com- pound bracts; plants glabrous or with spread- ing hairs Bracts of the involucre leaflike, pinnately compound; rays 1-7 (9), 1.5-10 cm long, some much longer than the involucres, some often nearly as long as the peduncles; inflores- cence open; bristly hairs of the fruit hook- ed; plants annual, of Washington County Yabea Bracts of the involucre pinnatifid; rays mostly 10-60 or more, seldom over 3 cm long or if longer then plants biennial, often not much exceeding the involucres, rarely longer than the peduncles; inflorescence congested; bris- tly hairs of the fruit glochidiate at apex; plants WAGES DREAC bet sen pie ame ge Oh ae aioect apn Daucus Involucel and involucre lacking ............ 9 Involucel and sometimes involucre present.. 13 Petals yellow; plants introduced, cultivated and adventive, ultimate segments of leaves filiform, 0.1—4 cm long, about 0.5 mm wide 10(9). 11(9). 12(11). 15(14). 16(15). GOODRICH: UTAH FLorRA, APIACEAE 69 Petals white, or yellow in Lomatium and then plants native; ultimate segments various but ofteniover0!oimnmbwidereen en jaenneeno. o. 11 Plants annual, not glaucous, widely culti- vated; leaves not especially crowded toward the base of the plant, the petiolules of the lowest pair of primary leaflets mostly less than 2 cm long, the ultimate segments 4-20 mm longi (Gill eee Anethum graveolens L. Plants perennial, glaucous, occasionally ad- ventive; leaves sometimes crowded toward the base of the stem, the petiolules of the lowest pair of primary leaflets often over 2 cm long, the ultimate segments 4-40 mm long Foeniculum Plants biennial from taproots, introduced; umbels often 6-12, or more per stem; fruit 3-4 mm long, the ribs filiform, not at all winged Carum Plants perennial from taproots or caudices, native; umbels rarely more than 8 per stem; fruit 3-14 mm long, the lateral and some- times the dorsal ribs winged.............. 12 Petals white; fruit 3-8 mm long, rounded, the dorsal and lateral ribs narrowly winged; sty- lopodium low conic; ultimate leaflets over 50 per leaf, 0.1-1(1.5)cmlong....... Ligusticum Petals yellow when fresh; fruit 8—14 mm long, dorsally flattened, the dorsal ribs filiform, the lateral ribs winged; stylopodium lacking; ulti- mate leaflets usually 3—45 per leaf, 0.3-9 cm longest Stes erat ei theater Lomatium Petals yellow or greenish yellow; plants culti- vated, rarely escaping (Parsley) .......... sareaaeae Petroselinum crispum (Mill.) A. W. Hill Petals white; plants not cultivated ......... 14 . Stems often purple-spotted, usually much branched, mostly with 10-30 or more um- bels; plants 5-30 dm tall, naturalized, weedy in moist or wet places in valleys and foothills, occasionally montane; involucre of 2-6 bracts, 2-6 (15) mm long ............ Conium Stems not purple-spotted with few branches, with (1) 3—7 (12) umbels; plants to 10 dm tall, native, often montane, involucre lacking or Seldombasiabovelnc ne enitene err maonccr 15 At least some of the ultimate leaflets 2-6 cm long and entire; leaves often withering shortly after anthesis; plants from a tuberous root or fascicle of tuberous roots, these easily de- tached from the stem, and often missing in herbarium specimens ...........- Perideridia Ultimate leaflets not over 2 cm long or if so not entire; leaves more persistent than above; plants from a taproot or a cluster of fleshy- fibroustootssso tee aera: 16 Involucels usually with more than 3 bractlets; fruit slightly compressed dorsally; root-crown mostly simple, without old long-persisting petiole-bases; plants rather rare in the eastern halfiofitheistate peer eer er Conioselinum GREAT BASIN NATURALIST Involucels lacking or rarely with more than 3 bractlets; fruit terete or slightly compressed laterally; root-crown simple or branched, usu- ally with fibrous long-persisting petiole- bases; plants common, widespread . Ligusticum Key Ill Plants acaulescent, sometimes with a pseu- doscape, or if subcaulescent then the mostly solitary peduncle longer than the short, leafy stem on which it is borne and lateral umbels if any mostly borne on the lower 1/3 of the plant; styles often over 1 mm long; stylopodium mostly lacking. 1 Fruit strongly flattened dorsally, the dorsal ribs filiform, not winged, the lateral ribs more or less winged, the body 8—15 (20) mm long or if shorter then plants usually pubescent (note: L. cous, L. minimum, and L. scabrum have small fruits and glabrous herbage); involucre lackin goppesr tie pet err as Lomatium Fruit not strongly flattened or if so the dorsal ribs winged, the body usually less than 8 mm long, the wings sometimes to 12 (15) mm long especially in plants with an involucre; plants Plabrousttojbitellousimaeeee ener 2 Stylopodium conic; leaves once compound with palmatifid leaflets; bractlets of involucels with 2-3 or more teeth; plants known from high elevations of the La Sal Mountains where apparentlyarane meaner nee Podistera Stylopodium lacking; leaves either more than once compound or leaflets not palmatifid; bractlets entire or plants not of high eleva- tions of the La Sal Mountains (except Oreoxis DOKE TU tne ier a Maeva EE ce 3 Ribs of fruit not winged or at most with low corky wings; carpophore well developed; leaves pinnate, a few of the leaflets sometimes pinnatifid and nearly bipinnate; plants of Cache, Garfield, and San Juan counties ..... 4 Ribs of the fruit with papery wings or if with low, corky wings then the carpophore lacking; leaves usually more than once compound; plants of broad distribution Ribs of fruit not winged; terminal umbel often subtended by a smaller umbel; petals and stamens white; plants often subcaules- cent, of the Bear River Range, Cache County iG CROCE RR URRon ao eaIn ERR aid Musineon Ribs of fruit with low, corky wings; umbel solitary; petals and stamens yellow; plants strictly acaulescent, of Garfield and San Juan counties (note: our plants may not have a car- 12:0) 0110) 2) eg Reem ee) Ve a Aletes 5(3). Vol. 46, No. 1 Fruit slightly compressed laterally, with low corky wings, 2-5 mm long and with rather conspicuous, persisting calyx teeth, the car- pophore lacking; plants from branched cau- dices, strictly acaulescent, not strongly aro- matic, mostly hirtellous or scabrous throughout or else the bractlets toothed, 1-10 (15) cm tall, montane, mostly above 2,440 m exceptiniOs trotterin eee Oreoxis Fruit compressed dorsally with prominent, more or less papery or corky wings, the calyx teeth obsolete or not persisting or else the carpophore well developed; plants from fibrous often enlarged tuberlike taproots, or if from branched caudices then with one or more of the following features: with one or more cauline leaves and from lower eleva- tions, strongly aromatic, glabrous, regularly over 10 cm tall; bractlets often not toothed wade itia tied 1360 eee Cymopterus Alternate Key III Plants with features of Key III but in flower or with immature fruits. tks 4(3). Leaves pinnate or pinnatifid, rarely trifid with linear entire segments Leaves either more than once-compound or else ternate or ternately divided with toothed to lobed leaflets . =. ..i92: 2 eee eneaReeeeee a ’ Petals and stamens white; terminal umbel sometimes subtended by a smaller axillary umbel; plants of Cache County ..... Musineon Petals and stamens yellow when fresh; umbel solitary; plants of the southern 1/2 of the state Bractlets of the involucel usually with 3 or more teeth or lobes linear-elliptic or oval to obovate; stylopodium present or lacking; plants of the La Sal Mountains ............. 4 Bractlets of the involucel mostly entire, linear or narrowly elliptic; stylopodium lacking; plants not known from the La Sal Mountains Leaflets more or less palmatifid, the major segments again trilobate to palmatifid; sty- lopodium conspicuous; base of plant with few if any persisting leaf bases.......... Podistera Leaflets pinnatifid or trifid; stylopodium lack- ing; base of plant clothed with persisting leaf bases). auc Gee Oreoxis bakeri Leaflets entire, 0.5-2 mm wide, linear- filiform to very narrowly elliptic; plants of Emery, Garfield, Iron, Sevier, and Wayne counties. 8). 44.3-..5 oe ee Lomatium At least some of the leaflets lobed, or if all entire then some over 2 mm wide and elliptic; distributionivanous eee eee eee 6 January 1986 GooprICH: UTAH FLORA, APIACEAE 71 = : aR Gye 6(5). Leaflets 0.3-1.2 cm long; rays 4—8, 2-10 mm — At least some of the leaves with 5—11 opposite long; involucels 4-5 mm long; plants of AS CCE on ee £ Garfield and central San Juan counties .. Aletes a e Din are Sy Dients. ob mountains and deserts, aromatic ornot.............. 14 = Some of the leaflets regularly over 1.2 cm 13(12). Primary leaflets 4-14 mm long, sessile; leaf long; rays (4) 6-13, 5-20 mm long; involucels blades 1-3.5 cm long; plants to about 12 cm 2-15 mm long; plants of Grand and extTeme tall, scabrous-hirtellous throughout or else northern San Juan counties . Lomatium latilobum bractlets of the involucel toothed... .. . Oreoxis 7(1). | Plants pubescent, not more so just below the = At least the lowest pair of primary leaflets a than elsewhere (see also Oreoxis ae 15-35 mm long, sessile or on peti- GLpinaWerey A paeh gece Gee Woes oe Lomatium olules to 15 mm long; plants 8—50 cm tall, a4 Plants glabrous or scabrous, sometimes Pete abe ae oat ve peduunals hirtellous just below the umbel and then with SES eae aie) Coen eee scabrous or hirtellous in the umbel, bractlets of the glabrousjleavesmen anneal eee 8 ; P involucel entire ........ Cymopterus lemmonii 8(7). mn eS pe of the ultimate hee over 2cm 14(12). Lowest pair of primary leaflets seldom over ong and entire or at most toot CO aa eet an sede 9 1/4 as long as the leaf blade, sessile or on — Ultimate leaflets less than 2 cm long or if petiolules to 18 mm long; leaves pinnately longemthenlobedanns a seen sae 10 SABES the blades more or less oblong in (OLUNUUTIVeRUp teats teats aan esters riko tio eNoaNGIe Te oinTet ole are 15 9(8). | Peduncles hirtellous just below the umbel, L stipe aaaerenl anal glabrous below; leaves glabrous, lowest pair rae ; west ey ee carets ue US Ses of primary leaflets sessile or on petiolules less SEs " St a4 S Sree we pence on ue than 2 cm long, the ultimate leaflets to 2.5 cm a One ee ean nae Me long, 1-3 (4) mm wide; plants not aromatic, pel SEED TOTES LESS SS OSS ONES 1 Be Ne OP ae ciate rave ok Ie OWL! prep heed Cee Sry eke eneean mee nee 16 SEN GRTCAGD oe Pep ME CRT nee OER Cymopterus lemmonii _15(14). Segments of leaves 1-12 mm long; some PI tab oa h bractlets of the involucels usually exceeding ec Flants glabrous or 1 scabrous then not more so the flowers; plants widespread, mostly of high just below the umbel than elsewhere; lowest elevations ........ Cymopterus hendersonii pair of primary leaflets either with petiolules pie i longer than above or some of the ultimate Tey Plants growing 792—2,320 m, or if of higher leaflets mostly longer or wider than above; elevations then scabrous and ultimate seg- plants of the northern 1/2 of the state or else cus of leaves 1-4 me long; bractlets of the strongly aromatic ................- Lomatium wae a ely ae the poe D eae ol the southern ol the state, mostly of low 10(8). Plants from a taproot, this sometimes en- to moderate elevations (L. parryi and L. larged and tuberlike, the crown simple or SCabriim Nara ME Te Lomatium or react wie a Hees 16(14). Calyx teeth lacking or to 0.3 mm long; ulti- De LN A ees aed tance ah en mate segments of leaves 0.2-0.3 mm wide ranean one) often conspicuous; leaf blades OTT A ITE Sometimes: with confluent) portions that aresme 9). 2) 9h es et a wider than the ultimate teeth or lobes ..... HL = Calyx ee see ue ea Ses: ments of leaves 0.5—1 (1.5) mm wide ...... =z Plants romvarsimple onmore often branchedi) salu meee ey oe Cymopterus terebinthus often woody caudex, this often clothed with long-persisting leaf bases; pseudoscape lack- ing; leaf blades finely and completely dis- sected so that the ultimate segments are the Aletes Coult. & Rose widest undivided portions of the blade ..... 12 11(10). Taproot very slender, with 1 or more abruptly Perennial, acaulescent, glabrous to pubes- expanded globose or ovoid tuberlike seg- cea : Foor biomuatcnpetios ments; plants strongly aromatic and from Salt POMS MON) ASEM) pina © P »P Lake County north to Cache County or if not late, the leaflets distinct or confluent, often aromatic then of northwestern Box Elder lobed and spinulose-dentate or entire; umbels County (L. ambiguum and L. cous) .. Lomatium compound; involucre lacking; rays few to sev- = Taproot slender or enlarged, if with a tuber- eral, spreading to reflexed; involucel of free or like enlargement then this gradually ex- united bractlets; calyx teeth conspicuous, del- panded! from anatrow portion plantsnota- _toid-ovate; stylopodium lacking; carpophore matic, distribution various (note: rare ow, . : aL STAD ode OeeT GEOL T Saat a anine tam divided to the base, sometimes readily decid wile yee) eee ee Cymopterus uous; fruit oblong to ovoid-oblong, ee terete, the ribs 12(10). Leaves with only about 2—4 opposite pairs of compressed laterally or sup primary leaflets; plants of mountains, mildly if atalllaromaticnneeeee eee error: 13 subequal, prominently corky-winged or ob- SCUre. 72 Aletes macdougalii Coult. & Rose Plants 7-20 cm tall, acaulescent, glabrous or scabrous, from a branched caudex, this more or less clothed with persistent leaf bases; leaves pinnate or some of the leaflets pinnati- fid and nearly bipinnate, with 2—6 opposite pairs of lateral leaflets or lobes, petioles 1.5—7 cm long, blades 1-5 cm long; leaflets 0.3-1.2 cm long, sessile, narrowly elliptic and entire or obovate and with 1—3(5) teeth or lobes; peduncles 5-15 cm long; umbel solitary; rays 4—8, 0.2—1 cm long; bractlets of the involucel about 4—6, 4—5 mm long, linear or linear-el- liptic, more or less united at the base; pedicels 1-2 mm long; calyx teeth about 1-1.5 mm long, narrowly to broadly deltoid; petals yel- low when fresh; styles 1.5—2.5 mm long; fruit 4—6 mm long, the ribs with small, more or less corky wings, the lateral ones about 1 mm wide, the dorsal ones smaller. Rock crevices, rocky slopes, and sandy ground, in pinyon-ju- niper and limber pine-bristlecone pine com- munities at 1,280 to 2,740 m in Garfield and San Juan counties; Arizona, Colorado, and Utah; 8(0). Our plants are referable to ssp. breviradiatus Theobald & Tseng. They could reasonably be included in the genus Cy- mopterus . Angelica L. Perennial, caulescent, single-stemmed herbs from a stout taproot; leaves pinnately to ternately 1-3 times compound, with broad leaflets; the lower blades on elongate petioles, the middle ones often arising directly from a dilated sheath, the upper ones often much reduced or lacking and the leaves reduced to a dilated sheath; umbels compound; involucre and involucel lacking or of narrow scarious or foliaceous bracts or bractlets; calyx teeth minute or obsolete; petals white, seldom pink or yellow; stylopodium broadly conic; car- pophore divided to the base; fruit elliptic- oblong to orbicular strongly compressed dor- sally, the lateral and dorsal ribs with small but obvious wings, or the ribs sometimes all corky-thickened and scarcely winged. 1. Leaflets mostly lanceolate to nearly linear, mostly over 3 times as long as wide; leaf blades oblong in outline; umbels 2—7, with 7-20 rays — Leaflets ovate or broader, or if lanceolate and over 3 times as long as wide then umbels mostly more than 7, and rays 20—40 GREAT BASIN NATURALIST Vol. 46, No. 1 2(1). | Leaves ternate-pinnate, the lowest pair of pri- mary leaflets on petiolules 3.5-12 cm long; leaflets coarsely toothed to lobed, the margins with about 1-3 teeth or lobes per cm; plants of the Deep Creek Mountains A. kingii Leaves pinnate or scarcely ternate-pinnate, the lowest pair of primary leaflets sessile or on petiolules to 1.5 cm long; leaflets finely to rather coarsely toothed, the margins with about 3-7 teeth per cm; plants not known from the Deep Creek Mountains ... A. pinnata Plants not over | m tall, mostly of rocky places above 3,050 m; umbels 1-3; leaflets 1—5 em long, serrate-dentate, rarely lobed; involucels of 1-3 or more linear bractlets about 3—10 mm long; ovaries and fruit glabrous or at most scabrous A. roseana — Plants mostly 1—2 m tall, mostly of wet places below 3,050 m; umbels several; leaflets 3—16 cm long, some usually lobed as well as toothed; involucels lacking; ovaries and young fruithispid to hirsute sane A. wheeleri Angelica kingii (Wats.) Coult. & Rose Great Basin Angelica. [Selinum kingii Wats. ]. Plants (3) 4-12 dm tall, glabrous except scabrous to short-hispid in the inflorescence, from a tap- root, without persisting leaf bases; leaves ter- nate-pinnate with 4—5 (6) opposite pairs of lateral primary leaflets, the lower pairs mostly pinnate, lower petioles to 25 cm long, dilated at the base, the upper ones reduced, lower blades to 40 cm long, oblong in outline, the upper ones reduced; lowest pair of primary leaflets about 2/3—3/4 as long as the leaf blade, ascending and more or less parallel to the primary rachis, on petiolules 3.5—12 cm long; leaflets 2-14 cm long, 4-15 (40) mm wide, lanceolate to nearly linear, coarsely toothed or lobed, the margins with 1—3 teeth or lobes per cm or rarely entire; peduncles mostly 4-17 | cm long; umbels 3-7; involucre lacking; rays 11-20, 1.5—9.5 cm long, scabrous; involucels lacking; pedicels 1-6 mm long, scabrous or short-hispid; petals white, sometimes marked with purple in age; stamens white; styles to about 1.5 mm long; fruit 4-5 mm long, densely hispid, the ribs slightly winged, the lateral wings a little wider than the dorsal | ones. Aspen-fir and streamside communities _ at 2,130 to 2,380 m in the Deep Creek Moun- | tains, Juab County; Nevada, eastern Califor- | nia, and southern Idaho; 5(0). Angelica pinnata Wats. Small-leaved An- | gelica [A. leporina Wats. ]. Plants 4.5-10 (15) dm tall, glabrous or nearly so except scabrous | to hirtellous in the inflorescence, without per- | January 1986 sistent leaf bases, from a taproot and some- times branched crown; leaves pinnate or partly bipinnate with 3 (4) opposite pairs of leaflets, the lowest pair sometimes bipinnate or partly bipinnate, the upper pairs pinnate, lower petioles 5-26 cm long, gradually ex- panded into a dilated, partly sheathing base, the upper ones reduced and the blades some- times sessile on the dilated sheath, blades (5) 9-21 cm long, more or less oblong in outline; leaflets 1.5—13 cm long, 4-37 mm wide, ses- sile, mostly lanceolate, occasionally elliptic, rarely ovate, finely to coarsely serrate, the margins with about 3-7 teeth per cm; pedun- cles 3.5-14 cm long; umbels (1) 2—5; involu- cre lacking; rays 7-14, 2-8.5 cm long, usually scabrous to hirtellous; involucels lacking or very rarely of 1 or more green to scarious linear or nearly linear bractlets to 3 (13) mm long; pedicels 3-7 mm long, glabrous or scabrous; petals white; styles to about 1 mm long; ovary glabrous to hirtellous; fruit 4—5 mm long, glabrous or sparsely hirtellous, the lateral wings about 1 mm wide, the dorsal wings about 0.5 mm wide. Tall forb, oak, maple, aspen, Douglas-fir, spruce-fir, willow, and wet meadow communities, very often along streams or around seeps and springs at 1,520 to 3,290 m in all Utah counties except Box Elder, Carbon, Millard, Morgan, and Rich; eastern Idaho to western Montana, south to Utah and Colorado; 65 (xv). Angelica roseana Henderson Rock Angel- ica. Plants 30-75 cm tall, strongly aromatic, glabrous or scabrous in the inflorescence, from stout taproots; stems stout, hollow, often 1—2 cm in diameter; leaves ternate-pinnate with 3-4 opposite pairs of lateral primary leaflets, the lower pairs bipinnate or ternate and petiolulate, the upper pairs pinnate and sessile; petioles to 8 cm long or lacking on the upper leaves and then blades sessile on a di- lated sheath; blades 5-17 cm long, ovate in outline, the upper ones reduced to lacking and leaves reduced to dilated sheaths; lowest pair of primary leaflets about 3/4 as long as the leaf blade, on petiolules 2.5—5.2 cm long, blades of leaflets (1) 2-5 cm long, ovate to orbicular, sharply serrate-dentate, rarely lobed; peduncles 4—17 cm long, the terminal one often about as thick as the stem, the lat- eral ones often partly enveloped in bladeless, dilated sheaths; umbels 1—3; involucre lack- GOODRICH: UTAH FLORA, APIACEAE 783 ing or occasionally of 1—2 linear bracts to 1.5 cm long; rays 15-30, 3.5-12 cm long, scabrous; bractlets of the involucel 1-3 (rarely more), 3-10 mm long, 0.2—0.5 mm wide, sep- arate, linear; pedicels 4-9 mm long, glabrous or scabrous; petals white; stamens whitish: styles about 2 mm long; ovary glabrous or at most scabrous; fruit ca 5 mm long, the ribs with wings about 1 mm wide. Talus slopes, boulder fields, rock stripes, and rocky ground, above timberline or upper spruce zone at (3,050) 3,200 to 3,570 m across the Uinta Mountains and Mount Timpanogos of the Wasatch Range in Daggett, Duchesne, Summit, Uintah, and Utah counties, Montana to Idaho, south to Colorado and Utah; 14 (iii). Angelica wheeleri Wats. Utah Angelica. [A. dilatata A. Nels. in Coult. & Rose]. Robust plants 1—2 m tall or taller, glabrous except in the inflorescence, mildly if at all aromatic, from stout rootcrowns with large fibrous roots; stems hollow, to 3 cm in diameter; lower leaves ternate-pinnately compound, with 3-5 opposite pairs of lateral primary leaflets, the lower pairs bipinnate or tripinnate and peti- olulate, the upper pairs often pinnate and ses- sile, petioles to 45 cm long, often dilated, blades to 40 cm long, ovate in outline; lowest pair of primary leaflets to 21 cm long, about 1/2 as long as the leaf blade, on petiolules to 5 cm long, blades of leaflets 3-16 cm long, 2-8 cm wide, lanceolate to ovate, serrate and some usually lobed; peduncles 2—29 cm long, often subtended by bladeless or nearly blade- less, dilated sheaths 2-20 cm long; umbels several; involucres lacking or occasionally of 1-2 linear bracts to 2 cm long; rays 20—45, 5-10 cm long, scabrous; involucels none; pedicels 5-12 mm long, glabrate to scabrous- hirsute; petals white; stamens whitish; styles about 1 mm long; ovary and young fruit scat- tered to densely hispid to hirsute, mature fruit 4-5 mm long, densely hispid, the dorsal and lateral ribs conspicuously winged. Boggy or very wet areas often in riparian communi- ties or in seeps and springs at 1,950 to 3,050 m in Cache, Juab, Piute, Salt Lake, Sevier, and Utah counties; endemic to Utah; 7 (ii). A. arguta Nutt. in T. & G. has been reported for Utah, but I have not seen a specimen and suspect that reports are based on A. wheeleri. It is apparently different from A. wheeleri only in the glabrous ovaries and fruit. a | SN Berula Hoftm. Perennial, caulescent, glabrous herbs from fibrous roots, often stoloniferous; leaves pin- nately compound or the submerged ones sometimes with filiform-dissected blades; umbels compound; involucre and involucel usually well developed; calyx teeth minute or obsolete; stylopodium conic; carpophore di- vided to the base, inconspicuous, adnate to the mericarps; fruit elliptic to orbicular, somewhat compressed laterally, glabrous, the ribs inconspicuous. Berula erecta (Huds.) Cov. Cutleaf Water- parsnip. [Siwm erectum Huds. ]. Stems 5-10 dm long or longer, from numerous fibrous roots; leaves pinnate with (3) 5-15 opposite pairs of lateral leaflets, or the submerged leaves (if present) often with filiform-dis- sected blades, petioles to 32 cm long or upper blades sessile on a dilated sheath, blades 2—31 cm long; leaflets 0.3—4 (6.5) cm long, sessile, nearly linear to lanceolate or ovate in outline, toothed to incised or occasionally a few entire; peduncles 1.5-8 cm long; umbels 3-20 or more; bracts of the involucre 1-6, 2—15 (25) mm long, linear or elliptic, entire, toothed, or rarely pinnatifid; rays 4-16, 0.5-2.5 (4) cm long; bractlets of the involucels ca 4—7, 1-7 mm long, linear or elliptic, entire; pedicels 2-7 mm long; petals white; stamens white; styles less than 1 mm long; fruit ca 2 mm long, the ribs obscure. In mud and water of streams, seeps, springs, marshes, swamps, margins of ponds and lakes, and in wet hang- ing gardens at 850 to 2,130 m in all counties of the state except Daggett, Emery, Grand, Iron, Morgan, San Juan, Summit, and Wayne; widespread in Europe, Mediter- ranean regions, and North America. The American plants are referable to var. incisa (Torr.) Cronq.; 64 (vi). Carum L. Biennial, caulescent, glabrous herbs from taproots; leaves pinnately compound; inflo- rescence of compound umbels; involucre and involucel lacking or of a few inconspicuous bracts or bractlets; calyx teeth obsolete; sty- lopodium low conic; carpophore divided to the base; fruit oblong to broadly elliptic- oblong, somewhat compressed laterally, evi- dently ribbed. GREAT BASIN NATURALIST Vol. 46, No. 1 Carum carvi L. Caraway. Plants 3—6 (10) dm tall; leaves 2-3 times pinnate and then often pinnatifid, with about 6—11 opposite or offset pairs of lateral primary leaflets, petioles to 15 cm long, the upper ones reduced and the blades sometimes sessile on a dilated sheath, the blades 5-16 cm long, oblong in outline; primary leaflets from less than 1/4 to about 1/2 as long as the leaf blade, sessile, the ultimate segments 2—8 (15) mm long, 0.5—2 mm wide, linear and entire or obovate and toothed to lobed; peduncles 4-12 cm long, usually sub- tended by a dilated sheath; umbels 6—12 or more; involucre lacking or inconspicuous; rays 6-12 (14), 1.5—8 cm long; involucels lack- ing or of minute scarious teeth; pedicels (5) 8-20 mm long; petals white; filaments white, the anthers pale green or whitish; styles about 0.5—0.85 mm long; fruit 3-4 mm long, the ribs filiform. Introduced, cultivated, the fruits used in flavoring, escaping, and occasionally persisting in mountain brush, meadow and aspen communities, at 1,375 to 2,640 m in Box Elder, Cache, Daggett, Davis, Duchesne, Salt Lake, Sanpete, Sevier, and Summit counties; native to Eurasia, now widespread across the United States; 10 (iv). Cicuta L. Perennial, caulescent, glabrous, violently poisonous herbs, from clusters of fibrous roots, some of these generally tuberous-thick- ened; base of stem thickened, with hollow chambers separated by transverse septa; in- ternodes of stems hollow; leaves 1—3 times pinnate or ternate-pinnate, with well devel- oped leaflets (ours); umbels several, com- pound; involucre wanting or of a few incon- spicuous narrow bracts; involucel of several narrow bractlets or rarely lacking; petals white or greenish; calyx teeth evident; sty- lopodium depressed or low-conic; carpophore divided to the base, deciduous; fruit ovate or orbicular, compressed laterally, the ribs usu- ally prominent and corky. Cicuta maculata L. Water Hemlock. [C. douglasii (DC.) Coult. & Rose, misapplied - the name belongs to plants north and west of _ Utah]. Plants, 6—21 dm tall or taller, with clusters of fibrous roots surmounted by a ]} thickened crown; stems 5—15 mm or more in diameter; leaves pinnate or ternate-pinnate | with 4—7 opposite pairs of lateral primary |} January 1986 leaflets, the lower pairs again pinnate, the upper pairs once pinnate and sessile, lower petioles about 5—40 cm long, the upper ones reduced and the blades often sessile on di- lated sheaths, lowest pair of petiolules 1-3 cm long, leaflets 2-11 cm long, 3-25 mm wide, narrowly lanceolate to lanceolate or occasion- ally linear, finely to coarsely serrate; pedun- cles (2) 4-15 cm long; umbels 6—30 or more; involucre lacking or of | or few linear bracts to 1 cm long; rays 15-26, 1.5-4 cm long; bractlets of the involucels about 6, 1-4 mm long, linear or narrowly deltoid, pale yellow- green or purplish, scarious-margined; pedicels 3-10 mm long; calyx teeth about 0.5 mm long, often pale green with whitish mar- gins; petals white; stamens white; styles 0.5—1 mm long; fruit 2-4 mm long, oval to globose, the ribs prominent, more or less corky, green, often wider than the darker (often purple) intervals. Along streams, rivers, ditches, canals, margins of pond and lakes, in wet meadows and marshes at 1,370 to 2,320 m in Beaver, Cache, Daggett, Duchesne, Kane, Millard, Piute, Salt Lake, San Juan, Sanpete, Summit, Tooele, Uintah, Utah, Wasatch, Washington, Wayne, and Weber counties; widespread in North America; 46 (v.) Some of our plants have leaflets less than 5 times as long as wide (a feature of var. maculata , which is found mostly east of Utah), but in these specimens, as well as others from the state, the styles are not more than 1 mm long. All Utah specimens I have seen belong to var. angustifolia Hook., the common phase in western North America. Conioselinum Hoftm. Perennial more or less caulescent herbs from a taproot or cluster of fleshy-fibrous roots, sometimes with a caudex; leaves pin- nately or ternate-pinnately decompound; in- florescence of compound umbels; involucre lacking or of a few narrow or leafy bracts; involucels of well-developed, narrow, often scarious bractlets; calyx teeth obsolete; petals white; stylopodium conic; carpophore divided to the base or nearly so; fruit elliptic or ellip- tic-oblong, slightly dorsally compressed, glabrous, the lateral ribs evidently thin- ; winged, the dorsal ribs less so and corky. i Conioselinum scopulorum (Gray) Coult. & Rose [Ligusticum scopulorum Gray]. Plants GOODRICH: UTAH FLORA, APIACEAE 75 perennial 3-10 dm tall, glabrous except in the inflorescence, from a fusiform taproot with simple or sparingly branched crown, without persisting leaf bases or these few and weakly persisting; leaves pinnate or ternate-pinnate with (3) 4—5 opposite pairs of lateral primary leaflets, the lower pairs 2—3 times pinnate and petiolulate, the upper pairs pinnate and pin- natifid and sessile or nearly so, petioles 3-23 cm long, blades 3.5-19 cm long, ovate in outline, lowest pair of primary leaflets 1/2—2/3 as long as the leaf blade; on petiolules (0.5) 1—3.5 cm long, ultimate segments 2-15 mm long, 1-5 mm wide; peduncles 3—21 cm long, often subtended by a dilated sheath, this usu- ally with a reduced sessile blade; umbels 1-3; involucre lacking or of | or few linear bracts to 1 cm long; rays 9-15, 1.5-5 cm long; in- volucels of 3-6 linear or linear-filiform bractlets 2-8 mm long; pedicels 4-12 mm long; petals white; stamens white; styles to about 1.3 mm long; fruit 4-6 mm long, lateral ribs narrowly corky winged, the dorsal ones not winged. Apparently rare, along streams at 2,500 to 3,200 m in Daggett, Grand, Garfield, Piute, San Juan, Summit, and Wayne coun- ties; Wyoming to Arizona and New Mexico; 18 (iii). Plants of C. scopulorum are often con- fused with those of Ligusticum porteri. The two taxa differ in the following subtle ways, with features of L. porteri in parentheses; fruit dorsally flattened (nearly _ terete); bractlets of the involucel often 3 or more (0-2, rarely more); terminal umbel solitary or sub- tended by alternate lateral umbels (often sub- tended by opposite or whorled umbels); and plants from a taproot, with a mostly simple crown and with few if any persisting fibrous leaf bases (the crown simple or branched and often with numerous, persistent, fibrous leaf bases). In addition, the rays average shorter and the ultimate segments of the leaves are less conspicuously veined than in those of L. porteri. Conium L. Biennial caulescent glabrous herbs from stout taproots with purple-spotted, freely branching hollow stems; leaves pinnately or ternate-pinnately dissected; umbels com- pound, several or numerous; involucre and involucels of small, lanceolate to ovate bracts or bractlets; calyx teeth obsolete; petals white; 76 GREAT BASIN NATURALIST stylopodium depressed conic; carpophore en- tire; fruit broadly ovoid, somewhat laterally compressed, with prominent, raised, often wavy slightly winged ribs. Conium maculatum L. Poison Hemlock. Plants 5—30 dm tall, glabrous, violently poi- sonous; leaves pinnate or ternate-pinnately decompound with 6—9 opposite pairs of lat- eral primary leaflets, the lower pairs usually twice or more pinnate and then pinnatifid, petiolulate, the upper pairs once pinnate and pinnatifid and sessile, petioles of larger leaves 4—18 cm long; larger leaf blades to 30 cm long, reduced upward and sessile on dilated sheaths, ovate in outline; lowest pair of pri- mary leaflets less than 1/2 to 2/3 as long as the leaf blade, on petiolules 1—5.5 mm long or those of upper leaves shorter; ultimate leaflets pinnatifid, the lobes entire or toothed, the widest confluent portions 2—5 (10) mm wide; peduncles 2—7.5 cm long; umbels many; in- volucral bracts 2-6, 2—6 (15) mm long, entire and ovate or ovate-caudate or ovate-cuspidate to deltoid, green with scarious margins, or rarely pinnatifid; rays 9-16, 1-4 cm long; bractlets of the involucels 4—6, 1-3 mm long, shaped like the involucral bracts; pedicels 2-6 mm long; petals white; stamens white; styles about 0.5 mm long; fruit 2-2.5 mm long, the ribs prominently ridged, narrower than the intervals. Along ditches, streams, rivers, roadsides, and fence lines, in wet and boggy meadows and moist waste places at 1,400 to 2,135 (2,990) m in Box Elder, Cache, Davis, Duchesne, Juab, Rich, Salt Lake, Sanpete, Summit, Tooele, Uintah, Utah, and Weber counties; introduced from Eurasia, now wide- spread in North America; 34 (iv). Cymopterus Raf. Perennial, acaulescent or subcaulescent, glabrous or scabrous herbs from slender to greatly enlarged and tuberlike taproots to branching woody caudices; leaves all basal (these sometimes elevated on an aerial pseu- doscape) or basal and 1—-few cauline mostly on the lower 1/2 of the stems, ternate to pinnate or ternate-pinnately compound, rarely simiple and ternately cleft; umbels solitary to several, open or reduced to globose heads; involucres lacking or well developed; involucels of sepa- rate or united bractlets; pedicels obsolete to well developed; calyx teeth obsolete to con- Vol. 46, No. 1 spicuous; petals white, yellow, or purple; sty- lopodium lacking; carpophore lacking, incon- spicuous and adhering to the inner faces (com- missure) of the mericarps , or well developed and persistent on the pedicel and divided to the base; fruit ovoid to oblong, somewhat flat- tened dorsally, the lateral and usually 1 or more of the dorsal ribs prominently winged. The strongly aromatic members of the group with woody branched caudices and greenish acute conspicuous calyx teeth have been in- cluded in the genus Pteryxia. These plants typically have leaves with 6-10 opposite pairs of lateral primary leaflets. Most of those in Cymopterus are ternate or have only 2—6 op- posite pairs of lateral primary leaflets. How- ever, the caudex and sometimes the number of primary leaflets are repeated in C. bipinna- tus Wats., C. aboriginum Jones, and in other taxa long included in Cymopterus. If high volatile oil content is unique to taxa of the Pteryxia group, this feature, in combination with others might warrant generic segrega- tion. Chemical studies might prove useful in resolving this problem. I, Leaves 1 or 2 times pinnate or a few merely ternate, with entire (rarely bifid) linear or linear-elliptic leaflets 0.5—4 (5.5) cm long and ~ 1-2 (3) mm wide; plants caulescent, of Wayne and San Juan counties, rare ......... C. beckii — Leaves not as above in all features; plants acaulescent or subcaulescent with 1-3 leaves mostly on the lower 1/3 of the stem ......... 2 2(1). | Peduncles rather densely hirtellous just be- low the umbel, mostly glabrous elsewhere; leaves mostly twice pinnate with 2—4 opposite pairs of mostly trifid or pinnatifid primary leaflets, the primary leaflets all sessile or the lower pairs on petiolules to 15 mm long; plants not aromatic, sometimes with 1 or 2 cauline leaves, montane, mostly above 2,400 Teg eogys te aan oe ae C. lemmonii — Peduncles not hirtellous just below the um- bel, sometimes scabrous but then not more so just below the umbel than elsewhere; leaves various but seldom with the above combina- tion of features; plants various ............. 3 3(2). Plants strongly aromatic, from a branched more or less woody caudex, mostly clothed at the base with long-persisting leaf bases and sometimes stem bases, often of rocky places; leaves with (4) 6-10 opposite or offset pairs of lateral primary leaflets, completely and finely dissected so that the ultimate segments (0.3-1.5 mm wide) are the widest undivided parts of the blade; calyx teeth rather promi- nent, about 0.5—1 mm long, acute, greenish (Bteryxia’ group) ss o0-> 3) ae eee 4 | January 1986 Plants not strongly aromatic, from fibrous tap- roots with simple or sparingly branched crowns, without or with few persisting leaf bases, not specific for rocky places; leaves once ternate to pinnately decompound with 2-6 opposite or offset pairs of lateral primary leaflets, sometimes not completely dissected, confluent portions of blades sometimes broader than the ultimate teeth or lobes; calyx teeth to about 0.5 mm long, rarely acute . 5 Lowest pair of primary leaflets (1/5) 1/2—3/4 or more the length of the leaf blade, mostly 3-9 cm long, several times longer than the upper pairs, on petiolules 2-4 cm long; stems fre- quently with | or 2 cauline leaves on the lower 1/3; fruiting styles (2.5) 3-4 mm long, mostly curved or coiled; involucels 2-5 mm long, not exserted beyond the flowers; plants growing 1,400-2,320 (2,560) m C. terebinthinus Lowest pair of primary leaflets 1/4 or less the length of the leaf blade, to 2.7 cm long, often not more than twice as long as some of the upper pairs, sessile or on petiolules to 1 cm long; leaves strictly basal; fruiting styles to 2 (2.5) mm long, straight or nearly so, bractlets of involucels 2-10 mm long, some often exserted beyond the flowers; plants growing at (very rarely 2,285) 2,740—3,660 m sot dctcacoligace arta ic aerate Santee Pee C. hendersonii Involucels scarious, purplish or whitish with purple nerves, the bractlets mostly over 3 mm wide, sometimes united to midlength; involu- cres like the involucels but larger or some- times reduced or lacking; broadest wings of the fruit (2)3—7 mm wide ................. 6 Involucels greenish or the bractlets very nar- row and divided to the base or nearly so; involucres lacking; broadest wings of the fruit to 2.5 mm wide or to 5.5 in C. purpureus.... 8 Rays 1-3.5 cm long, usually at least some exceeding the well- developed to obsolete in- volucre through all stages of phenology, not obscured by the dense mature fruits; lobes of the involucre and involucel not multinerved; carpophore well developed, tending to per- sist on the pedicels after the mericarps have fallen; fruit often oblong in outline, the wings 1.7-3(4) mm wide; plants of the Colorado drainage oe racr ee pie tee C. bulbosus Rays 0.3-1 cm long, rarely longer, not exserted beyond the always well-developed involucre, or if exserted then the lobes of the involucre and involucel multinerved, ob- scured by the dense mature fruits; car- pophore lacking or hairlike and more or less adhering to the faces of the mericarps and not persistent on the pedicels; fruit ovate to orbic- ular in outline, the wings 3-7 mm wide; plants of the Great Basin and Colorado drainaveg yer ne ney rt OPN ete. uae 7 7(6). 10(8). 11(10). GOODRICH: UTAH FLORA, APIACEAE al Lobes of the involucels and usually of the involucres with more than 3 parallel purplish nerves that extend to or near the tip; involucre sometimes reduced to a ring; plants of Kane and Washington counties ates Sieh Sahl hihi eee te C. multinervatus Lobes of the involucels and involucres with a midnerve extending to the tip and sometimes 1 or 2 lateral shorter nerves extending to about midlength or less; involucre well devel- oped; plants widespread ..... C. purpurascens Involucels green and of the same texture as the leaves, seldom scarious-margined, the bractlets mostly 1.5-4 mm wide, plants mostly obscurely viscid and dotted with nu- merous adhering grains of sand especially on the scapes and petioles, growing at 850—1,890 Involucels rarely wholly green, not of the same texture as the leaves, often scarious- margined and/or the bractlets linear or nar- rowly elliptic and not over 1.5 mm wide; plants not viscid, rarely with numerous ad- hering grains of sand, sometimes growing lOO GROIN sconsesocvoevedeusodseede. 10 Leaves once ternate, the 3 leaflets ternately lobed or cleft, the blades with confluent por- tions 5-35 mm wide; outer rays 1-3.3 cm long; bractlets of involucel entire or rarely tridentate; pseudoscape lacking; plants of the southern 1/2 of the state ........ C. newberryi Leaves 2—3 times pinnate with 2 (3) opposite pairs of lateral primary leaflets, some rarely ternate, the blades with confluent portions 1-7 (12) mm wide, rays to 1.3 cm long; bractlets of the involucel often with 2—3 teeth; pseudoscape often present; plants wide- SPreaG rise t SAE etree eee ere C. acaulis Carpophore lacking; pedicels obsolete or to 1 mm long; rays obsolete or short and concealed in the very dense fruits of a globose headlike inflorescence; styles less than 1 mm long, or if rays evident (to 17 mm long) and styles to 2 mm long then leaves ternate without a rachis and with lobes sharply dentate-serrate; plants endemic to the Great Basin, not found above TAOS Onrined: Stas sree er aie al ere eee eal ll Carpophore well developed, divided to the base; pedicels mostly over 1 mm long; rays short or rather long, not concealed in the dense inflorescence; styles mostly 1-3 mm long; leaves 2—3 times pinnate or if ternate then usually with a rachis and lobes not den- tate-serrate; plants of broad distribution, those with rays less than 17 mm long often founckaboverlegsOimieeee Eee e oe 13 Leaves pinnate with 2—3 pairs of lateral pri- mary leaflets, rarely some ternate, the blades narrowly ovate to oblong in outline; rays and pedicels obsolete; inflorescence a dense glo- bose head; wings of fruit more or less spongy thickened; styles 0.5—0.8 mm long; anthers WER eee ae eee ences C. globosus 12(11). 13(10). 14(13). 15(13). GREAT BASIN NATURALIST Leaves ternate or occasionally simple and ter- nately cleft, the blades reniform, orbicular, to ovate in outline; rays and sometimes pedicels more or less evident when young, inflores- cence various; wings of fruit papery; styles various; anthers yellowish or purplish ...... Leaves 2 and opposite, rarely 3, the ultimate lobes crenate; pseudoscape solitary, subter- ranean; peduncles solitary; rays 3-10 mm long, hidden at maturity in the very dense globose headlike umbel; styles about 0.4 mm long; plants of Juab, Sanpete, and Sevier counties C. coulteri Leaves often more than 2, the ultimate lobes dentate; pseudoscapes or stems rarely soli- tary; peduncles 1—4 (6); rays 8-17 mm long, not hidden as above, the umbel not globose and headlike; styles 1.5—2 mm long; plants of Beaver and southwestern Millard counties . . C. basalticus An aerial pseudoscape rather quickly devel- oping, (3.5) 5-24 cm long; leaf blades with 4—6 opposite or offset pairs of lateral primary leaflets; umbels sometimes nodding on re- curved peduncles; petals white or yellow, sel- dom turning light purple; plants mostly mon- tane in central and western Utah Seg ewcrce eae 14 Pseudoscape lacking or mostly subterranean, the aerial portion not over 3 cm long; leaf blades with 2—4 opposite pairs of lateral pri- mary leaflets, or ternate; umbels not nodding; petals white or yellow, often turning dark pur- ple 15 Leaves 3 times pinnate, finely and completely dissected, with the ultimate segments the widest undivided portions of the blades, these to 2 mm long and to 1 mm wide; upper pri- mary leaflets not tending to be confluent with the rachis; petals white; anthers purple; plants of western Utah C. ibapensis Leaves 2 (3) times pinnate, the ultimate lobes or teeth 1-5 mm long, 0.5—3 mm wide, these often not as wide as the confluent portions of the blade that are up to 12 mm wide; upper primary leaflets tending to be confluent with the rachis, and pinnatifid or only lobed; petals yellow or white; anthers yellow or white; plants of northern and central Utah . C. longipes Leaves once pinnately compound with 2 op- posite pairs of lateral primary leaflets, or a few ternate or rarely biternate, glaucous, conflu- ent portions of the blades (3) 6—25 (40) mm wide; petals and stamens bright yellow when fresh, fading to cream or white in herbarium specimens; plants of the Uinta Basin at 1243016 S00hne eases C. duchesnensis Leaves ternate or 2—3 times pinnately com- pound with up to 4 opposite pairs of lateral primary leaflets, glaucous or not, the conflu- ent portions mostly 1-4 mm wide or if wider then the leaves ternate; petals yellow, purple, or white when fresh, if yellow then turning dark purple in herbarium specimens; plants of broad distribution Vol. 46, No. 1 16(15). Petals cream-pink to pale purple when fresh, with light or moderate purple markings in herbarium specimens; rays of the umbel 2-18 mm long; pedicels to 3 mm long; blades of leaves 1-3 cm long, pinnately dissected, the confluent portions rarely over 3 mm wide, plants scabrous, of Garfield, Iron, and Kane counties, at moderate to high elevations .... C. minimus Petals yellow or purplish when fresh, turning dark purple in herbarium specimens; rays, pedicels, and leaves mostly longer than above or the leaves mostly ternate with confluent portions often 5—21 mm wide; plants wide- spread C. purpureus Cymopterus acaulis (Pursh) Raf. Plains Spring-parsley. [Selinum acaule Pursh]. Plants 5-18 (27) cm tall, from a simple or rarely branched, deep seated, nearly linear or slightly to much enlarged fibrous taproot; herbage often more or less viscid and dotted with sand grains; pseudoscapes 1—2 (3) per plant, 0.5—5.5 cm long, often partly or wholly subterranean; leaves basal or more often whorled with the peduncles atop the pseu- doscape, occasionally | or 2 on a pseudoscape- like stem, 2—3 times pinnate, with (1) 2 (3) opposite pairs of lateral primary leaflets; peti- oles 2-8 (11) cm long, blades (1)2—5.5 (7) em long, the confluent portions 1—7(12) mm wide, oblong, ovate, to nearly linear in out- line; primary leaflets 5-35 mm long, gradu- ally reduced upward, pinnate to bipinnatifid with few to several rounded to narrow lobes, the ultimate teeth or lobes to 10 (16) mm long, to 2mm wide; peduncles 1-14, (1.5) 3-14 (19) | cm long; involucres lacking; rays about 6-9, about 1-13 mm long; bractlets of the involucel } 3-8 (11) mm long, ca 1.5—4 mm wide, more or less united at the base, entire or with 2—3 teeth or lobes, green or purple in age, of the texture of the leaves; pedicels to 2 mm long; calyx teeth ca 0.2 mm long, greenish; petals white, yellow, or purple; stamens the color of the petals; styles ca 2.5 mm long; carpophore | lacking; fruit 5-10 mm long, the wings | slightly longer than the body, to 2 mm wide, - slightly corky, some of the dorsal ones some- | times obsolete. With 4 more or less intergrad- | ing varieties in the state. Il, Petals and stamens yellow when fresh, sooner or later fading to white or cream when dried fresh January 1986 2(1). Peduncles mostly shorter than the leaves, to ca 4cm long; wings of the fruit mostly strongly wavey and often erose, to 7 mm long: leaf blades to about 4 cm long; plants seldom over 7 cm tall, of the Great Basin ............. Lae GIS RO OO OTERO Re C. acaulis var. parvus — Peduncles equaling or exceeding the leaves, to 14 (19) cm long; wings of the fruit straight or slightly wavey, mostly entire or obscurely erose, to 10 mm long; leaf blades to 7 cm long; plants often over 7 cm tall, of the Colorado Basins aii cy sees a ahh: C. acaulis var. fendleri Petals and stamens purple; peduncles mostly exceeding the leaves; plants of Kane County C. acaulis var. higginsii — Petals and stamens white; peduncles mostly shorter than or equalling the leaves; plants of Daggett County and the Uinta Basin....... 5 eee Soe A oes Ne C. acaulis var. acaulis Var. acaulis Desert shrub, sagebrush, and juniper communities at 1,432 to 1,980 m in Daggett, Duchesne, and Uintah counties; Saskatche-wan and Minnesota west to Oregon and south to Texas and northern Utah; 21 (vi). Plants with white flowers grow among those with yellow flowers in the Uinta Basin, where it is difficult if not impossible to recognize two taxa. Even when fresh the white flowers do not seem as bright as those of Wyoming, and the Uinta Basin materials seem transitional to var. fendleri. Var. fendleri (Gray) Goodrich, comb. nov. [based on: C. fendleri Gray Mem. Amer. Acad. II. 4: 56. 1849; C. decipiens Jones, type from Cisco]. Desert shrub, blackbrush, sage- brush, and pinyon-juniper communities often on sandy soil at 1,885 to 1,890 m in Duchesne, Carbon, Emery, Garfield, Grand, Kane, San Juan, Uintah, and Wayne counties; Utah and Arizona; 74 (v). This taxon has long been sepa- rated at the species level from C. acaulis. The rather recent discovery of intermediate plants in the Uinta Basin and of other yellow-flow- ered varieties (var. parvus from the Great Basin and var. greeleyorum Grimes & Pack- ard of Oregon) greatly weaken the case for such separation. Var. higginsii (Welsh) Goodrich, comb. nov. [based on: C. higginsii Welsh Great Basin Nat. 35: 377. 1976]. Desert shrub com- munities, often on sandy alluvium of Tropic Shale at about 1,525 m in Kane County; en- demic; 4 (0). The color of the petals persists as a bright purple long following collection, and the color marks this variety as distinct from any in the complex. GOODRICH: UTAH FLORA, APIACEAE 79 Var. parvus Goodrich, var. nov. Similis Cy- mopterus acaulis var. acaulis sed parvioribus saepe, petalis flavis et alis fructus undulatis et erosis valde differt, sed similis var. fendleri in floribus flavis et var. acaulis in scapis brevis. HOLoTyPE: Utah. Tooele Co., 32.7 km 326 degrees NW of Vernon. Skull Valley-Stans- bury Mtns., T6S, R7W, Sec 32, near 1/4 cor- ner with Sec 33, 1,585 m. Juniper-big sage- brush community, stabilized aeolian sand, 7 June 1984, S. Goodrich 20458 (BRY); isotypes UC, NY, RM, CAS, POM, UTC, UT, US. Additional specimens: Tooele Co., Ibid., 7 May 1984, S. Goodrich 20251 (BRY; UT, RM, NY, UTC). Desert shrub, sagebrush, and ju- niper communities, often on aeolian sand, at 1,400 to 1,585 m in Millard and Tooele coun- ties; endemic; 15 (xii). This variety is similar to var. acaulis in the short scape, but differs in the yellow flowers. It is similar to var. fendleri in the yellow flowers but differs in the short scapes, and it is similar to the extralimital var. greeleyorum Grimes & Packard in the short scape and yellow flowers but apparently most closely related to the latter taxon. It differs from var. greeleyorum in the strongly undu- late, erose wings of the smaller fruit. Fruiting material of var. greelyorum was loaned through the kindness of Dr. Patricia Packard. Cymopterus basalticus Jones Dolomite Spring-parsley. Plants 4-15 cm tall, from a taproot with simple or branched crown, glabrous, glaucous, with 1-few mostly etio- lated, subterranean, short pseudoscapelike stems; pseudoscape mostly lacking, if present short and enveloped in bladeless sheaths, the crown sometimes with a few long-persisting leaf bases; leaves basal, (2) 3-9 per plant, ternately divided without a rachis, or occa- sionally simple and ternately cleft, petioles 1.5—5 cm long, blades 1—3.5 cm long, orbicu- lar to reniform, confluent portions 4—32 mm wide; leaflets 5—30 mm long, sessile, orbicu- lar, ternately lobed, the major lobes again lobed, mostly ternately so, the ultimate lobes coarsely dentate; peduncles 1—4 (6) per plant, 3.5-8 (14) cm long; involucre lacking; rays 6-14, 8-17 mm long, usually evident in fruit; bractlets of the involucel 6—8, 2-5 mm long, more or less united at the base, white, pink, or purplish with white scarious margins; pedicels obsolete or nearly so; petals white or purplish; stamens yellowish or purplish; 80 styles 1.5—2 mm long; carpophore lacking; body of fruit 3-6 mm long, the wings 4-7 mm long, 1-2 mm wide, whitish, papery, some of the dorsal ones often reduced. Desert shrub communities, gravelly hills and alluvial fans mostly of dolomite substrate, at 1,705 to 1,985 m in western Beaver and Millard counties; Utah and adjacent White Pine County; Ne- vada, a rather narrow Great Basin endemic; 21 (v). The orbicular to reniform leaf blades without a rachis are unique in the genus. Cymopterus beckii Welsh & Goodrich Pin- nate Spring-parsley. Plants 20-40 cm tall, glabrous, weakly if at all aromatic, caulescent with leaves extending well up the stem, froma taproot with a simple or sparingly branched crown, often clothed at the base with long- persisting leaf bases; leaves 1 or 2 times pin- nate, with 2-3 opposite pairs of lateral leaflets, or the upper ones sometimes ternate, petioles 2-13 cm long, blades 2-10 cm long; leaflets 3-7, 0.5—4 cm long, or the terminal one to 5.5 cm long, 1—2 (3) mm wide, sessile, linear or linear-elliptic, entire or rarely a few bifid; peduncles 4—8 (19) cm long; umbels 1-3 per stem; involucres lacking; rays 6-11, 0.6—1.4 cm long; bractlets of the involucels about 5, 1-5 mm long, to 1 mm wide, green- ish or with narrow scarious margins, mostly separate; pedicels 1-3 mm long; petals and stamens bright yellow when fresh, fading whitish when dried (whitish within 2 years in herbarium specimens); styles 1.2-2.2 mm long; carpophore weak, adhering to the meri- carps; fruit 6-8 mm long, oblong, the lateral wings to about 1 mm wide, the dorsal ones narrower, some often obsolete. Sandy or stoney places, pinyon-juniper-mountain brush communities at 1,700 to 2,150 m in San Juan and Wayne counties; endemic; 8 (iv). Apparently closely allied to C. lemmonii, but differing in entire leaflets, glabrous peduncles and rays, and the slightly longer fruit. Cymopterus bulbosus A. Nels. Onion Spring-parsley. Plants 8—27 cm tall, glabrous, glaucous, from a stout, thickened, often bul- bous, fibrous taproot with a simple or spar- ingly branched crown; pseudoscapes obsolete or 1—2 (3), to 6.5 cm long and often partly or wholly subterranean, enveloped in dilated bladeless sheaths; leaves few to several, basal or whorled atop the pseudoscape with the peduncles, rarely 1 or 2 cauline, (1) 2—3 times GREAT BASIN NATURALIST Vol. 46, No. 1 pinnate, with (2) 3-6 opposite or offset pairs of lateral primary leaflets, the upper pairs often once-pinnate and more or less confluent, blades 2-10 cm long, ovate to oblong or nearly linear, confluent portions 1-5 mm wide, lowest pair of primary leaflets to 4 cm long, sessile or on petiolules to 2(5) mm long, the other primary leaflets progressively re- duced upward, ultimate lobes and teeth 1—8 (12) mm long, about 1-4 mm wide, more or less rounded; peduncles (1) 3-8 (11) per pseu- doscape, 4-18 cm long; involucre obsolete or reduced to a ring or cup, or the bracts well developed, to 13 mm long, translucent, white, more or less united, with a green midrib and occasionally 1 or 2 lateral nerves that extend to about 1/3 the length of the bract; rays 5-15, 1-3.5 cm long, usually ex- ceeding the involucre at all stages of phenol- ogy; involucels 3-10 mm long, the bractlets more or less united at the base, similar in texture and color to the involucre, with a green center and midrib or the midrib some- times purple, rarely with | or 2 lateral nerves extending to about midlength; pedicels 3—9 mm long; calyx teeth about 0.5-1 mm long, scarious, white, like the involucel, with a green midrib to about midlength; petals white, sometimes purplish in age; stamens white, or purple especially in age; styles about 2—4 mm long; carpophore divided to the base, more or less persistent on the pedicel after the mericarps have fallen; body of the fruit 6-11 mm long, to 2 (3) mm wide, the wings (7) 9-13 mm long, 1.7—3 (4) mm wide. Desert shrub and juniper communities at 1,220 to 2,005 m in the Colorado drainage in Carbon, Duch- esne, Emery, Garfield, Grand, San Juan, Uintah, and Wayne counties; Wyoming to New Mexico and Arizona; 95 (xi). See C. pur- purascens . Cymopterus coulteri (Jones) Mathias Two- leaf Spring-parsley. [C. corrugatus var. scop- ulicola Jones; C. corrugatus var. coulteri Jones; Rhysopterus jonesii Coult. & Rose]. Plants 4-11 cm tall, glabrous from a slightly to much enlarged fibrous taproot with simple crown, this giving rise to a solitary, mostly subterranean pseudoscape 2-6 cm _ long; leaves 2 (rarely 3 or 4 and the third and fourth ones usually smaller), opposite, borne at or near ground level, ternate or rarely simple and ternately cleft, petioles 1-3 cm long, January 1986 blades 2—4 cm long, ovate to nearly orbicular in outline, confluent portions (5) 8-38 mm wide; leaflets 7-35 mm long, the lateral ones sessile mostly ternately lobed, the terminal on a winged more or less confluent rachis to 1 em long, ternately cleft, the main lobes again lobed or crenate-toothed; peduncle solitary, _ 2-7 cm long; umbel globose, headlike, 1.5—5 cm across in pressed fruiting specimens; in- -volucre lacking; rays 7-14 or perhaps more, - 3-10 mm long, somewhat evident at anthesis, ‘but hidden by the dense mature fruits; _bractlets of the involucel 2-4 mm long, linear to narrowly ovate, green or purplish in age, often 3—nerved, with whitish or purplish scar- ious margins; pedicels shorter than the bractlets; calyx teeth minute but white and of the texture of the petals, deciduous; petals white; filaments white, anthers purple; styles (including the stigmas) ca 0.4 mm long; car- pophore lacking; body of fruit 5-7 mm long, the wings 7-10 mm long, to about 2 mm wide, papery. Desert shrub, black sagebrush, and juniper communities, often on Arapien shale and other clayey and gravelly barrens or semibarrens at 1,540 to 1,700 m in Juab, San- | pete, Sevier, and Tooele counties; endemic; 28 (vii). The strong tendency for plants to have but 2 leaves is unique in the genus. Cymopterus duchesnensis Jones Uinta Basin Spring-parsley. Plants 7-23 cm tall, from a slender or more often enlarged bulbous taproot with simple or branched crown, glabrous and glaucous, not or weakly aro- matic; stems short, often branched; pseu- doscape lacking; leaves basal or 1-3 or more _ cauline, pinnate with 2 pairs of lateral leaflets, or occasionally a few ternate, or rarely biter- nate, petioles 2.2-11 cm long, blades (2) 3-10.5 mm long, ovate to oblong in outline, the confluent portions (3) 6—25 (40) mm wide; leaflets 1-5, on petiolules 2-32 mm long, ter- nately cleft or divided, the major lobes to3 cm long, (8) 5-15 (20) mm wide, often again toothed or lobed and mostly ternately so, the ultimate teeth or lobes 1—8 mm wide; pedun- cles 1-3 per stem, 7-17 cm long; involucres lacking; rays 6-17, 1.5—4.4 cm long; bractlets of the involucel lacking or more often 1-7, | 1-5 mm long, more or less united at the base, i linear; pedicels (2) 4—9 mm long; petals and ‘istamens bright yellow when fresh, fading to cream or greenish in herbarium specimens, GOODRICH: UTAH FLORA, APIACEAE 81 not turning purple; styles 2-2.2 mm long; carpophore divided to the base, more or less persistent on the pedicel; body of fruit 5-9 mm long, the wings to 11 mm long, 2—2.5 mm wide, undulate to corrugated, more or less papery and not corky. Desert shrub, sage- brush, and juniper communities, sandy clay and clay semibarrens of Duchesne River, Mancos Shale, Morrison, Uinta, and Wasatch formations at_i,430 to 1,860 m, centered in Uintah County and also in eastern Duchesne County, Utah, and extreme western Moffat and Rio Blanco counties, Colorado; 47 (xiii). Cymopterus globosus (Wats.) Wats. Golf- ball Spring-parsley. [C. montanus var. globo- sus Wats. |. Plants 4-10 cm tall, from a slender or thickened fibrous taproot; pseudoscape (1) 2—6 cm long, all or nearly all subterranean, often loosely enveloped in dilated bladeless sheaths; leaves 1 or 2 atop the pseudoscape and some usually arising directly from the fibrous root and then with etiolated subter- ranean petioles, pinnate or bipinnate and then trifid or pinnatifid, with 2-3 opposite pairs of sessile lateral primary leaflets, or rarely ternate, petioles 1.5—6 cm long, blades 2—5 cm long, narrowly ovate to oblong, con- fluent portions 5—10 mm wide, lowest pair of primary leaflets 10-18 mm long, the ultimate lobes to 4 mm long, to 2 mm wide, mostly toothed; peduncles 1 or 2, 3-6 cm long; in- volucre lacking; umbel a globose head, the rays and pedicels obsolete; involucels con- cealed in the dense flowers and fruits; petals white; stamens white; styles 0.5-0.8 mm long; carpophore lacking; body of the fruit about 6 mm long, the wings ca 9 mm long, to ca 2.8 mm wide, wider toward the outside of the head, spongy thickened. Desert shrub communities at 1,400 to 1,525 m in Box Elder, Juab, Millard and Tooele counties; eastern California, Nevada, and western Utah; a Great Basin endemic; 6 (0). Cymopterus hendersonii (Coult. & Rose) Crong. Mountain Rock-parsley [Pseu- doteryxia longiloba Rydb.; Pterxia hender- sonii (Coult. & Rose) Mathias & Const.; Pseu- docymopterus hendersonii Coult. & Rose]. Plants (3) 5—34 cm tall, glabrous, strongly aro- matic, from a branched woody caudex, clothed at the base with old petiole and pe- duncle bases, these sometimes persisting for a few or several years without shredding; leaves 82 basal, bipinnate or occasionally partly tripin- nate with 5—10 opposite or offset pairs of lat- eral primary leaflets, petioles (1) 2-14 cm long, blades (1) 1.5-10 cm long, oblong in outline, finely dissected so that the ultimate segments are the widest undivided parts of the blade, lowest pair of primary leaflets about 1/4 or less the length of the leaf blade, 5-27 mm long, sessile or on petiolules to 1 cm long, upper primary leaflets gradually reduced, the ultimate segments 1-12 mm long, 0.3-1.4 mm wide, acute, with a usually whitish tiny mucro; peduncles 7-30 cm long; umbels com- pact; involucres lacking; rays 6—16, 0.5—2.4 cm long, the inner ones shorter than the outer ones and often abortive; bractlets of the in- volucel 2-6, 2-10 mm long, linear, acute; pedicels 1-5 mm long; calyx teeth about 1 mm long, persisting in fruit, greenish, often red- dish tinged, acute; petals and stamens bright yellow when fresh, fading whitish in herbar- ium specimens; styles to 2 (2.5) mm long; carpophore divided to the base; fruit 4-8 mm long, the wings to about 1 mm wide, some of the dorsal ones sometimes obsolete. Talus, cliffs, ledges, rocky spruce-fir, limber -pine, and alpine communities at (2,285) 2,740 to 3,660 m in Beaver, Box Elder, Cache, Daggett, Duchesne, Grand, Juab, Piute, Salt Lake, San Juan, Sevier, Summit, Tooele, Uintah, and Utah counties; southwestern Montana and central Idaho, south to New Mexico; 66 (x). Cymopterus ibapensis Jones Ibapah Spring- parsley. [C. watsonii (Coult. & Rose) Jones]. Plants 7-25 cm tall, glabrous or granular- scabrous, not or weakly aromatic, from a lin- ear taproot, this hardly if at all swollen, with a simple or occasionally branched crown; pseu- doscapes 1—2 (5) per root, the aboveground portion 3.5-10 cm long, commonly envel- oped at the base by scarious dilated bladeless sheaths; leaves whorled atop the pseu- doscape, rarely some arising directly from the root, tripinnate, with 5-6 opposite or offset pairs of lateral primary leaflets, petioles (1) 1.5—3.5 cm long, blades (2.5) 4-11 cm long, ovate in outline, completely dissected so that the ultimate segments are the widest undi- vided portions of the blade; lowest pair of primary leaflets about 1/2 to over 3/4 as long as the leaf blade, sessile or on petiolules to 2 em long, with 4—6 (8) opposite or offset pairs of GREAT BASIN NATURALIST secondary leaflets, the ultimate segments to 2 Vol. 46, No. 1 | mm long, to about 1 mm wide; peduncles (2) 4—8 per pseudoscape, 2—15 cm long; umbels sometimes nodding on the sometimes re- | curved peduncles; involucre lacking; rays | 10-18, 5-20 mm long; bractlets of the in- | volucels to 4 mm long, to 0.5 mm wide, sepa- rate or nearly so, green with a purple midrib and narrow scarious margins; pedicels 4—6 mm long; calyx teeth to 1 mm long, greenish; petals white; filaments white, anthers purple; styles 1-2 mm long; carpophore divided to | the base; body of fruit 5-8 mm long, the wings _ 6-9 mm long, to 2 mm wide, some of the | | dorsal ones sometimes reduced. Greasewood- | sagebrush, sagebrush-grass, and pinyon-ju- — niper communities at 1,520 to 2,755 m in — Beaver, Box Elder, Iron, Millard, Piute, ei- ther or perhaps both Juab and Tooele, and | Washington counties; southeastern Oregon, and Nevada; 30 (viii). Cymopterus lemmonii (Coult. Dorn Spring-parsley. tanus (Gray) Coult. & Rose; Thaspium mon- tanum Gray; Ligusticum montanum (Gray) — Gray]. Plants 8—50 cm tall, glabrous except on _ the peduncle and in the inflorescence, not or — weakly aromatic, from a taproot with simple | & Rose) | [Pseudocymopterus — lemmonii (Gray) Coult. & Rose; P. tidestromii | Coult. & Rose; P. versicolor Rydb.; P. mon- | | | | | | { | { or branched crown, more or less clothed at the — base with shredded persisting leaf bases; pseudoscape lacking; leaves basal and some-_ times 1 or 2 cauline ones on the lower 1/3 of | the stem and occasionally 1 on the upper 3/4, | mostly bipinnate and then often bifid or pin- | natifid in the lower part, with 2—4 opposite or | offset pairs of lateral primary leaflets, rarely | pinnate in part with entire leaflets; petioles 1-13 cm long, with a dilated base, blades (1) 2-8 cm long, confluent portions to 4 mm } wide; lowest pair of primary leaflets 1/4—2/3 as | long as the leaf blade, sessile, or on petiolules _ to 15 mm long, the ultimate segments 2—20 | mm long, linear or narrowly elliptic; pedun- | cles 1-9 per plant, (4) 9-28 cm long, rather densely hirtellous just below the umbel; in-| volucre lacking, or rarely of 1 or 2 small bracts; rays (5) 9-18, 0.8-2.5 cm long, glabrous, scabrous or hirtellous; bractlets of the in-. volucels 5-11, to 5.5 mm long, linear or nar-_ rowly elliptic, separate or united at the very. base, green or sometimes with a scarious or, January 1986 purplish margin; pedicels obsolete or to 2 mm long; calyx teeth less than 0.5 mm long, decid- uous; petals and stamens bright yellow when fresh, pale or purplish in age; styles about 2 mm long; carpophore apparently lacking to well developed and divided to the base; fruit mostly 3-6 mm long, the wings ca 1.5 mm wide, some of the dorsal ones sometimes ob- solete. Grass-forb, aspen, Douglas fir, and spruce-fir communities, and windswept ridges and raw escarpments especially in limestone, at 2,375 to 3,600 m in Beaver, Emery, Garfield, Grand, Iron, Piute, San Juan, Sanpete, Sevier, Washington, and Wayne counties; southeastern Wyoming to Arizona and Durango, Mexico; 138 (vii). Oc- casional specimens have been confused with Lomatium juniperinum. The following key should aid in separating the two. iL. Peduncles hirtellous just below the umbel; rays 0.8—2 cm long, sometimes scabrous; plants otherwise glabrous, of Sanpete County and southward; petals yellow when fresh, but turning whitish in herbarium specimens; pedicels obsolete or to 2 mm long; lowest pair of primary petiolules to 1.5 cm long Renae Fe bay one Cymopterus lemmonii _ Peduncles glabrous or not any more pubescent than the rest of the plant; rays 1-8 cm long; plants rarely with glabrous herbage, and then with cream or white petals, from Sanpete County and northward; pedicels 3-16 mm long; lowest pair of primary peti- olules 1-3 cm long..... Lomatium juniperinum Cymopterus longipes Wats. Long-stalk Spring-parsley [Peucedanum — lapidosum Jones, type from Echo, Summit County; C. lapidosus (Jones) Jones; Lomatium lapidosum (Jones) Garrett]. Plants 7-30 (50) cm tall, glabrous, not aromatic, from a thickened fi- brous taproot with a simple or sparingly branched crown; pseudoscapes 1—3, 4—24 cm long, mostly aerial, rarely a small portion of it subterranean, more or less enveloped by di- lated bladeless sheaths; leaves whorled atop the pseudoscape, rarely any rising from the fibrous root, (1) 2 (3) times pinnately com- pound, with mostly 4-6 opposite or offset pairs of lateral primary leaflets, the upper pairs often more or less confluent and merely pinnatifid, petioles 1-5 cm long, blades 3-8.5 cm long, oblong to ovate in outline, the con- fluent portions 2-12 mm wide; lowest pair of primary leaflets 1-5 cm long, sessile, or on GOODRICH: UTAH FLORA, APIACEAE 83 petiolules to 5 mm long, the ultimate lobes or teeth about 1-5 mm long, about 0.5—3 mm wide; peduncles 3-18 per pseudoscape, 4-24 cm long; umbels sometimes nodding, on the sometimes recurved peduncles; involucre lacking; rays 4—11, 0.5-3.3 cm long; bractlets of the involucel to 7 mm long, mostly less than 1 mm wide, mostly separate, green with very narrew scarious margins; pedicels 1-12 mm long; calyx teeth 0.2—0.5 mm long; petals yel- low or white when fresh, when yellow fading to white in herbarium specimens; stamens the color of the petals; styles about 2 mm long; carpophore divided to the base; body of the fruit 4—6 mm long, the wings 5-8 mm long, 1-2 mm wide. Mostly in sagebrush-grass communities, but also in pinyon-juniper and mountain brush communities at 1,340 to 3,155 m in Box Elder, Cache, Carbon, Daggett, Davis, Duchesne, Juab, Morgan, Rich, Salt Lake, Sanpete, Summit, Tooele, Uintah, Utah, Wasatch, and Weber counties; southeastern Idaho and northcentral Utah to northwestern Colorado; 160 (xxi). White- flowered specimens are common in the Bear River Range and occasional to the central part of the Wasatch Range. The yellow petals of more southern specimens turn whitish in the herbarium, and the ranges of the two color variants are difficult to determine from herbarium specimens. Other than the color difference, there seems to be no way to tell the phases apart. A third phase, in which the fruits are Lomatium-like (dorsal ribs not or scarsely winged), apparently has white flow- ers. This phase, known from Summit County and adjacent Wyoming, is referable to C. lapi- dosum (Jones) Jones, which may be worthy of the specific status given it by Jones. Cymopterus minimus (Mathias) Mathias Least Spring-parsley. [Aulospermum mini- mum Mathias]. Plants 3-10 (10) cm tall, acaulescent, scabrous, from a slender to much enlarged often deep-seated taproot, with few to several often soboliferous branches; stems mostly subterranean and etiolated; pseu- doscapes lacking or short and subterranean; leaves mostly 2—3 times pinnately dissected, with 3-4 opposite pairs of lateral primary leaflets, petioles 0.5—2 cm long or sometimes much longer with etiolated subterranean por- tions, blades 1-3 cm long, the confluent por- tions 1-6 (10) mm wide, the primary leaflets 84 sessile or the lowest pair on petiolules to 0.5 cm long, the ultimate segments to 3 mm long, to ca 2 mm wide; peduncles 1.5—14 cm long; rays of the umbel mostly 5-10, 0.2—1.8 cm long; bractlets of the involucel about 3-4, 2—4 mm wide; pedicels nearly obsolete or to 3 mm long; calyx teeth minute or lacking; petals cream-pink or pale purple (reputedly white) with whitish margins or with moderately pur- ple markings in herbarium specimens; sta- mens whitish; styles about 2 mm long; car- pophore divided to the base; fruit 4—8 mm long, the wings to 1 mm wide. Ponderosa pine, bristlecone pine, spruce-fir, and per- haps pinyon-juniper communities, at (2,190) 2,440 to 3,170 m, in Garfield, Iron, and Kane counties; endemic; 23 (i). Occasional speci- mens are intermediate to C. purpureus, and more work is needed to establish the range of the small plants with cream-pink or pale pur- ple petals. These plants have been confused with C. purpureus var. rosei, and they are similar in the short rays, short pedicels, small fruit, small leaves, and scabrosity, but the leaves are not ternate, and they are more finely dissected than those of C. purpureus var. rosei. In some features of the leaves and in distribution this taxon is more closely allied with C. purpureus var. purpureus. At the extreme (Cedar Breaks) these plants are very different, but through a series of recent collec- tions from the Markagunt, Paunsaugunt, and Table Cliff plateaus and Escalante Mountains, a rather close relationship to C. purpureus var. purpureus is evident. Perhaps this is only a part of the C. purpureus complex and could be treated as a variety, but no such combina- tion is proposed herein. The color of the petals seems to be diagnostic. Cymopterus multinervatus (Coult. & Rose) Tidestr. Purple-nerved _ Spring-parsley. [Phellopterus multinervatus Coult. & Rose]. Plants 10-15 cm tall, glabrous and glaucous, from a linear or slightly to much enlarged fibrous taproot; pseudoscapes lacking or soli- tary, to 7.5 cm long, partly or mostly subter- ranean, often enveloped by dilated, scarious bladeless sheaths; leaves basal or whorled, with the peduncles atop the pseudoscape, 2—3 times pinnately compound, with 3-5 op- posite pairs of lateral primary leaflets; petioles 1—7.5 cm long; blades 1-7 cm long, ovate in outline, lowest pair of primary leaflets to 4.5 GREAT BASIN NATURALIST Vol. 46, No. 1 cm long, sessile or on petiolules to 5 mm long, the ultimate lobes to 4 (7) mm long, to 2 mm wide, sometimes with small rounded teeth; peduncles 1-8, 4-10 cm long; involucre to 1 (1.5) cm long, the bracts more or less united, sometimes forming a cup or reduced to a ring, greenish basally and centrally, with broad, white scarious margins, multinerved, the nerves purplish, more or less parallel; rays (3) 5-11, 0.3-1 cm long, obscured by the dense fruits, included in or exserted beyond the in- volucre; involucels 5-10 mm long, like the involucre in color, texture, and venation, but never reduced to a ring; pedicels to about 4 mm long, included in the involucel; calyx teeth 0.5—1 mm long or somewhat enlarged and simulating the involucre; petals white or purple; stamens white or purple; styles about 2—3 (4) mm long; carpophore lacking; body of the fruit 7-10 mm long, the wings 12-13 mm long, 5-7 mm wide. Desert shrub and sage- brush communities at 1,220 to 1,525 m in Kane and Washington counties; southern Utah to southern California and to southwest- ern Texas and northern Mexico; 7 (0). Cymopterus newberryi (Wats.) Jones Sweetroot Spring-parsley. | Peucedanum new- berryi Wats.]. Plants 7-18 cm tall, more or less slighty viscid and often dotted with adher- ing sand grains, from a slender or slightly to much enlarged fibrous taproot with simple or rarely branched crown; pseudoscape lacking; leaves arising directly from the fibrous root, ternate, rarely simple and ternately cleft; petioles 3.5-10.5 cm long, (1.6) 2.5—-5 times as long as the blades, often partly subter- ranean, blades 1—4(5.5) cm long, confluent portions 5-35 mm wide; leaflets 1-3.5 cm long, cleft or divided and mostly ternately so, the major lobes again lobed or toothed and often ternately so, the ultimate lobes or teeth to 6 mm long, to 5 mm wide; peduncles 1-10, 5-17 cm long, often partly subterranean; in- volucre lacking; rays 5—16, the central ones often greatly reduced or obsolete, the outer ones 1—3.3 cm long; bractlets of the involucels 3-12 mm long, to3 mm wide, entire, green, or sometimes purplish in age, with texture of the leaves; pedicels about 1 mm long; calyx teeth about 0.5 mm long, deciduous; petals and stamens yellow when fresh, fading to cream or greenish cream in herbarium speci- mens; styles about 2-3 mm long; carpophore | January 1986 lacking; body of fruit 5-8 mm long, the wings 6—10 mm long and 1—1.5 mm wide, more or less corky, some of the dorsal ones obsolete. Desert shrub, blackbrush, sand sagebrush, desert grassland, and juniper communities, mostly on very sandy soil, at 850 to 1,830 m in Beaver, Garfield, Grand, Kane, Millard, San Juan, Washington, and Wayne counties; southern Utah and northern Arizona; 47 (x). Cymopterus purpurascens (Gray) Jones Widewing Spring-parsley [C. montanum var. purpurascens Gray]. Plants 5-15 cm tall, glabrous and glaucous, from a mostly enlarged tuberlike fibrous taproot with simple or spar- ingly branched crown, the crown usually with few to several persistent shredded leaf bases; pseudoscapes lacking or to 3 per plant and to 6 cm long, mostly subterranean, usually envel- oped by scarious dilated bladeless sheaths; leaves basal or more or less whorled atop the pseudoscape, 2-3 times pinnately com- pound, with 3-6 opposite pairs of lateral pri- mary leaflets, the pairs gradually reduced up- ward, petioles 0.6-5 cm long, sometimes longer including etiolated subterranean por- tions, blades 1.2—7 cm long, oblong to ovate in outline, confluent portions to 3 (5) mm wide, lowest pair of primary leaflets (0.4) 1-2 (4) cm long, sessile or on petiolules to 3 mm long, the ultimate lobes or teeth rounded, mostly with narrow-scarious margins; pedun- cles 1-3 per pseudoscape, 3—9 cm long; in- volucre 8—14 mm long, more or less united at the base and sometimes to about midlength, whitish, scarious, the lobes with a greenish or purplish midnerve extending to the tip, and usually 1 or 2 parallel lateral, much shorter nerves; rays about 4—7, rarely longer than 1 cm, mostly shorter than the involucre, hidden in the dense broadly winged fruits; involucels like the involucre but shorter (about 5—7 mm) and usually with the lateral nerves over 1/2 as long as the midnerve, the nerves occasionally branched; pedicels to 5 mm long, mostly con- cealed in the involucels and in the dense fruits; calyx teeth less than 0.5 mm _ long, rounded; petals white or purplish with a green or purplish midvein; filaments white, anthers purple; styles about 2 mm long; carpophore lacking or hairlike and less than 0.02 mm wide, not persisting on the pedicel; body of fruit 6-11 mm long, the wings 9-16 mm long and 3-6.5 mm wide. Desert shrub, sage- GOODRICH: UTAH FLORA, APIACEAE 85 brush, pinyon-juniper, bullgrass, and pon- derosa pine communities, on aeolian sand to heavy clay at 1,065 to 2,745 m in all Utah counties except Daggett, Davis, Grand, Mor- gan, Rich, Summit, Wasatch, and Weber: southeastern Idaho to southeastern California and northwestern New Mexico; 106 (xxi). Cy- mopterus purpurascens is often confused with C. bulbosus, but it is distinguished by a num- ber of features. In addition to those listed in the key, the lobes of the involucre and in- volucel of C. bulbosus mostly are only 1—nerved, this occasionally with 1 or 2 short lateral nerves. The involucre is sometimes reduced to a ring or cup in C. bulbosus, but it is always well developed in C. purpurascens ; C. bulbosus also flowers later (often a month or so) than does C. purpurascens, and it is confined to lowlands mostly of heavy soil, whereas C. purpurascens is found on a wide range of sites and soils. However, rare speci- mens (Neese 7169 and Thorne et al. 1707) show the broad wings of fruit typical of C. purpurascens, but they have rays well over 1 cm long that are exserted beyond the involu- cre, and at least some of the fruits have a well-developed carpophore. Perhaps these specimens indicate rare hybridization of these two taxa. Cymopterus purpureus Wats. Variable Spring-parlsey. Plants 5-26 cm tall, from a slender to much thickened fibrous taproot with a simple or branched crown; stems soli- tary to several, arising at or just below ground level; pseudoscape mostly lacking, mostly less than 2 cm long when present and then usually mostly subterranean; leaves basal or nearly so, ternate or (1) 2—3 times pinnately com- pound, with up to 4 opposite pairs of lateral primary leaflets, petioles 1-7 cm long, blades 1.5—13 cm long, mostly ovate in outline, low- est pair of primary leaflets mostly over 1/2 and to 3/4 as long as the leaf blade, sessile or on petiolules to 32 mm long, the ultimate lobes or teeth acute or rounded; peduncles 1-5, 3-21 cm long; involucre lacking; rays 5-22, 0.2—9.5 cm long; bractlets of the involucel 4—8, 2-4 mm long, separate or united at the base, acute to acuminate, entire; pedicels 1-10 mm long; calyx teeth less than 0.5 mm long, deciduous; petals yellow when fresh, drying dark purple in the field or within a year or 2 in herbarium specimens; stamens yellow 86 when fresh, remaining yellowish or cream or at least pale in herbarium specimens; styles about 2-3 mm long; carpophore divided to the base; body of the fruit about 4-8 mm long, the wings 5-10 (12) mm long, 1.5-4 mm wide, often marked with purple. With 3 more or less integrading varieties in the state. il. Fruiting rays 5—8 (15), 0.2—2 (3) em long; fruiting pedicels 1-5 (7) mm long; wings of fruit 5-8 mm long, to 2 mm wide; leaf blades 1—3.5 (4) cm long, mostly (not always) ternate, the leaflets with rounded lobes; plants glabrous, or more often scabrous, lower to midmon- tane, mostly of central Utah C. purpureus var. rosei — Fruiting rays (8) 12-22, (2) 2.5-7 (9.5) em long; fruiting pedicels 5-10 mm long; wings of fruit 8-10 (12) mm long, (2) 2.5—4 mm wide; leaf blades commonly 3-9 (13) cm long, pin- nately compound, rarely ternate, often with acute ultimate segments, plants mostly glabrous, rarely scabrous, of deserts and lower montane, widespread ............... 2 Plants conspicuously glaucous, of Washing- ton, Iron, and Beaver counties; some wings of fruit apparently thickened and spongy; ulti- mate teeth of leaves acute................ C. purpureus var. jonesii _— Plants not conspicuously glaucous, not of the above counties except Washington; wings of fruit mostly thin and papery; ultimate teeth of leaves acute to rounded C. purpureus var. purpureus Var. jonesii (Coult. & Rose) Goodrich, comb. nov. [based on: Cymopterus jonesii Coult. & Rose Rev. N. Amer. Umbell. 80. 1888; Aulospermum jonesii (Coult. & Rose) Coult. & Rose]. Sagebrush, pinyon-juniper, and mountain brush communities at 1,520 to 1,905 m in Beaver, Iron, and Washington counties; southwestern Utah and adjacent Nevada; 10 (i). Var. purpureus Desert shrub, sagebrush, pinyon-juniper, mountain brush, ponderosa pine, and rarely aspen-fir communities in sandy to heavy clay soils at 1,100 to 2,375 (2,880) m in Carbon, Duchesne, Emery, Garfield, Grand, Kane, San Juan, Uintah, Washington, and Wayne counties; eastern and southern Utah, western Colorado, north- ern Arizona, and northwestern New Mexico; 134 (v). Specimens with rather broad leaflets from the Uinta Basin (Neese et al. 7273, and White and Neese 123) indicate a close rela- tionship to and possible hybridization with C. duchesnensis . GREAT BASIN NATURALIST Vol. 46, No. 1 Var. rosei (Jones) Goodrich, comb. nov. [based on: Aulospermum rosei Jones in Coult. & Rose. Contr. U.S. Nat. Herb. 7: 179. 1900; C. rosei (Jones) Jones]. Pinyon-juniper, sage- brush, mountain brush, bull grass, limber pine, white fir, and rarely desert shrub com- munities, in marly limestone, shaley slopes, and clay or sandy clay soils at (1,615) 1,760 to 2,290 (2,650) m in Duchesne, Juab, Millard, Sanpete, Sevier, and Wasatch counties; en- demic to central Utah; 28 (x). The plants from the Sevier drainage are quite removed from plants of the rest of the complex. In this case the differences seem to be a function of isola- tion, but the Duchesne County materials are not so isolated from plants of var. purpureus. The differences in this case seem to be more a function of ecological stress than of isolation. An independent origin is suggested. Random plants of var. purpureus with features inter- mediate to those of var. rosei are found in other parts of the range of var. purpureus (Neese 5775; Foster 4380, 8338; and N. H. Holmgren et al. 1998). Most of the plants of the Sevier drainage are quite different from those of var. purpureus , but some would be difficult to place without location data (Welsh 12803; Neese & White 2925; and anonymous 1322a, UT 11951). Recognition at varital level seems appropriate. Cymopterus terebinthinus (Hook.) T. & G. Rock Parsley. [Selinum terebinthinum Hook. ; (Pteryxia terebinthina (Hook.) Coult. & Rose]. Plants (12) 15-35 (40) cm tall, glabrous, strongly aromatic, froma heavy, nearly woody root surmounted by a mostly branched caudex, the caudex clothed with leaf bases that often persist for a few years without shredding; pseudoscape lacking; leaves basal and often 1-3 on the lower 1/3 (1/2) of the stem, mostly 2—4 times pinnately or ternate- pinnnately compound, with (4) 6—10 opposite or offset pairs of lateral primary leaflets, peti- oles 2-13 cm long, blades 1.5—14 cm long, finely and completely dissected so that the ultimate segments are the widest undivided part of the blade, lowest pair of primary leaflets mostly 3-9 cm long, (1/5) 1/2—3/4 as long as the leaf blade, on petiolules 2—4 cm long, the ultimate segments 1—5 (7) mm long, 0.5—-1 (1.5) mm wide; peduncles 10-34 cm | long; involucre lacking; rays 7-13, 0.7—5 (8) cm long; bractlets of the involucel (0) 1-5, 2-5 January 1986 mm long, separate or united at the base, lin- ear or linear-subulate; pedicels 2—5(10) mm long; calyx teeth 0.5-0.9 mm long, acute, rather persistent; petals and stamens bright yellow when fresh, fading whitish and rarely yellow for more than 2 years in herbarium specimens; fruiting styles (2—5) 3-4 mm long, mostly curved or coiled; carpophore divided to the base, persisting on the pedicel; body of fruit 5-8 mm long, the wings 6-9 mm long, 0.5-1.5 (2.5) mm wide, some of the dorsal ones sometimes reduced or obsolete. There are two varieties in Utah. 1. Lateral pairs of primary leaflets all longer than their internodes; lowest pair of primary leaflets various but often less than 4 times as long as wide, with the lower secondary leaflets sometimes petiolulate; leaf blades not skeletonlike, the ultimate segments appear- ing to be more numerous or longer or both than in the following variety; plants of the Uinta Basin and northern Utah. ........... C. terebinthinus var. calcareus Upper 4-6 pairs of lateral primary leaflets equal or shorter than their internodes; lowest pair of primary leaflets 4—10 times longer than wide, with sessile secondary leaflets; leaf blades skeletonlike, the ultimate segments appearing to be fewer or shorter or both than in the preceding variety; plants of eastern Utah south of the Uinta Basin C. terebinthinus var. petraeus Var. calcareus (Jones) Crong. [Cymopterus calcareus Jones]. Desert shrub, sagebrush, pinyon-juniper, and mountain brush commu- nities, often in talus, colluvium, and crevices of rock outcrops at 1,445 to 2,320 (2,560) m in Box Elder, Cache, Daggett, Juab, Rich, and Uintah counties; Montana to Colorado, and west to southern Idaho and northeastern Ne- vada; 36 (vii). Some of the Uintah County materials are transitional to plants of var. pe- _ traeus. Plants of var. calcareus are very simi- lar to those of C. terebinthinus var. albiflorus (T. & G.) Jones, which was originally de- scribed as having white flowers. Dried flowers soon turn white or whitish in all of the C. | terebinthinus complex, and the two varieties are both reported from the same regions of Montana and Wyoming. Unfortunately, the t i misnomer var. albiflorus has priority by many years, and our plants might belong to that taxon. Var. petraeus (Jones) Goodrich, comb. nov. [based on: C. petraeus Jones Contr. W. Bot. GOODRICH: UTAH FLORA, APIACEAE 87 8: 32. 1898]. Skeletonleaf Rock-parsley. Desert shrub, blackbrush, and pinyon-ju- niper communities, often in talus, colluvium, crevices of rock outcrops, and in sandy to clayey soil at 1,400 to 2,075 m in Emery, Grand, and San Juan counties; Great Basin in Nevada and southern Idaho, and Colorado drainage in Utah and northwestern Arizona; 16 (i). The disjunction from the Great Basin in Nevada to the Colorado Basin in Utah without occurrence in the Great Basin in Utah is most unusual. An independent origin is suggested for the Utah materials, which at present are known to be separated from plants of var. calcareus only by Desolation Canyon of the Green River. The Colorado drainage materi- als are as skeletonlike or more so than the Nevada materials but are not so distinct as to warrant separate status, even if from an inde- pendent origin. Neither the Nevada nor Utah materials are so distinct from plants of var. calcareus as to warrant recognization at the species level. They fit well into C. terebinthi- nus both morphologically and in volatile oils, and they are surrounded on 3 sides by other varieties of C. terebinthinus. Without distri- bution data some of the San Juan County spec- imens as well as some from central Nevada would be nearly if not wholly impossible to distinguish from some specimens of var. cal- careus of the Uinta Basin. Daucus L. Annual or biennial caulescent herbs from taproots; leaves pinnately dissected; umbels compound; involucre of pinnatifid bracts or lacking; involucel of toothed or entire bracts or lacking; calyx teeth evident to obsolete; petals white or those of the central flower of the umbel or umbellet often purple or rarely all the flowers pink or yellow; stylopodium conic; carpophore entire or bifid at the apex; fruit oblong to ovoid, slightly compressed dor- sally, evidently ribbed dorsally, with two ribs on the commissure, beset with stout spread- ing glochidiate or barbed ribs. iL, Plants biennial, introduced, cultivated and occa- sionally escaping and then somewhat weedy; widespread; bracts of the involucre mostly pinnatifid into mostly entire rather rigid elon- gate segments D. carota 88 Plants annual, native, not cultivated, known from the Virgin Narrows in Arizona, to be expected in Washington County; bracts of the involucre pinnatifid into often lobed or toothed nonrigid segments .. D. pusillus Michx. Daucus carota L. Carrot. Plants 6-10 dm tall, from a taproot; herbage glabrous or hir- sute; leaves in rosettes and cauline, mostly 1-2 times pinnate and then pinnatifid, with about 4—9 opposite or offset pairs of lateral primary leaflets, basal and lower cauline peti- oles to 15 cm long, basal and lower blades 5-15 cm long or more, the upper ones re- duced and sessile on dilated sheaths, lowest pair of primary leaflets about 1/3—1/2 as long as the leaf blade, on petiolules 4-15 mm long, ultimate segments 1-10 mm long, 0.5—2 mm wide, elliptic, or narrowly deltoid, or linear, often acute; peduncles mostly 8—30 cm long; umbels 4—10 or more; involucre of pinnatifid bracts 1-5 cm long, the segments linear and narrow; rays about 15—60 or more, (0.5) 1-6 cm long; involucels similar to the involucre but smaller, or the bractlets entire, 2-16 mm long; fruit 3-4 mm long, bristly hirsute in rows, the hairs or bristles about 2 mm long, minutely glochidate at the apex, the intervals often with shorter simple hairs. Cultivated in all counties of the state, wild (mostly along ditchbanks and waste places) mostly in the more populated counties of the state; intro- duced from Eurasia; 16 (iv). The wild plants (ssp. carota) differ from the cultivated plants [ssp. sativus (Hoffm.) Arcangeli] primarily in the size and flavor of the root. Foeniculum Adans. Biennial or perennial, caulescent herbs with strong odor of anise, glabrous, glaucous, from a taproot; leaves pinnately dissected with fili- form ultimate segments; umbels compouund; involucre and involucel lacking; calyx teeth obsolete; petals yellow; stylopodium conic; carpophore divided to the base; fruit oblong, subterete, or slightly compressed laterally, with prominent ribs. Foeniculum vulgare Mill. Sweet Fennel. Short-lived perennial herbs 1—2 m tall, from a taproot; stems solitary, branched above; leaves to 3-times ternate-pinnately com- pound with about 6—9 opposite pairs of ateral primary leaflets; petioles to about 15 cm long rather abruptly expanded into a dilated GREAT BASIN NATURALIST Vol. 46, No. 1 sheathing base or lacking and blades arising directly from the sheath; larger blades to 30 or 40 cm long, ovate in outline, finely and com- pletely dissected, the elongated filiform ulti- mate segments 4—40 mm long and less than 1 mm wide, the lowest pair of primary leaflets on petiolules often over 2 cm long; peduncles 1.5—6.5 cm long; umbels several; rays 10—40, 2-8 cm long; petals yellow; styles 0.3-0.4 mm long; fruit 3.5-4 mm long. Roadsides and waste places at 850 to 1,465 m in Utah and Washington counties; native of the Mediter- ranean region, widely introduced elsewhere, and in much of the United States especially toward the south, perhaps not well adapted to much of Utah except Washington County; 3 (0). Heracleum L. Biennial or perennial herbs from a taproot or fascicle of fibrous roots, leaves ternately or pinnately compound, with broad toothed or cleft leaflets; umbels compound; involucre lacking or of a few deciduous bracts; involucel lacking or of slender bractlets; flowers of the marginal umbellets generally irregular, the outer petals enlarged and often deeply bilobed; calyx teeth obsolete or minute; sty- lopodium conic; carpophore divided to the base; fruit orbicular to obovate or elliptic, strongly flattened dorsally, usually pubes- | cent, the dorsal ribs narrow, the lateral ribs | broadly winged. Brummit, R. K. 1971. Relationship of Heracleum lanatum Michx. of North America to H. sphondylium of | Europe. Rhodora 73:578—584. Heracleum lanatum Michx. Cow parsnip. [H. ) sphondylium ssp. lanatum (Michx.) A. & D. Love]. Stout single-stemmed perennial herbs 8-25 dm tall, from a taproot or cluster of fibrous roots, glabrate or thinly to densely | villous or villous-hirsute below to villous- | woolly above, especially on the nodes; leaves | ternate or the upper ones simple, petioles to 25 cm long or longer, or lacking on upper leaves with the petiolules and rachis arising / directly from a dilated sheath, blades to 40 cm | long or longer, ovate to orbieular; leaflets | 8-36 cm long or longer, ovate to orbicular, } usually with 3 major lobes that are again lobed | and coarsely toothed; peduncles 5-24 cm | long; involucre lacking or of few mostly linear | | | | | { i q i i January 1986 entire bracts to 2. cm long; rays 12—25, 3.5—12 cm long; involucels of 3-5 linear, subulate or caudate bractlets to 15 mm long; pedicels 6-26 mm long; petals white (2) 4—8.5 mm long, at least some deeply bilobed; filaments white, anthers whitish to dark green or yellow with pollen; styles about 1 mm long, the stig- mas incurved; fruit 8-12 mm long, obovate to obcordate, strongly flattened, the lateral ribs with wings about 1-1.5 mm wide, the dorsal ribs filiform. Aspen, tall forb, fir, oak-maple, willow, streamside, and wet meadow commu- nities and around seeps and springs at 1,430 to 2,930 m in Box Elder, Cache, Carbon, Davis, Duchesne, Juab, Salt Lake, Sanpete, Sevier, Summit, Tooele, Uintah, Utah, Wasatch, and Weber counties; Eurasia and across much of North America; 6] (iii). Hydrocotyle L. Perennial herbs; stems creeping or floating, rooting at the nodes; leaves petiolate, often peltate; inflorescences sessile, or borne on axillary peduncles; involucres small or lack- ing; petals white, greenish, or yellow; calyx minute or lacking; stylopodium conic to de- pressed; fruit orbicular to ellipsoid, more or less flattened laterally, the dorsal surfaces rounded or acute, the ribs obsolete or narrow and acute; carpophore lacking. Hydrocotyle verticillata Thunb. Water Pennywort. Plants glabrous, with slender creeping stems; leaves peltate, suborbicular, 0.5-6 cm wide, shallowly lobed and often crenate; petioles slender, 3-20 cm long or longer; peduncles slender, axillary; flowers apparently verticillate in few to several well- separated whorls; petals pale, small; fruits subsessile, subtruncate at the base, 1.5—2 mm long, 2-3 mm wide. Moist ground or in water at 850 to 1,005 m in Washington County; South America north to Massachussetts and California; 4 (0). Ligusticum L. Perennial caulescent or acaulescent herbs from taproots; leaves ternately or ternate-pin- nately compound or dissected, the lower ones with well-developed petioles, the upper ones with blades arising directly from dilated sheaths; umbels compound; involucre and in- volucel lacking or of a few narrow bracts or bractlets; calyx teeth evident or obscure; GOODRICH: UTAH FLORA, APIACEAE 89 petals white; stamens white; stylopodium low-conic; carpophore divided to the base: fruit oblong to ovate or suborbicular, sub- terete or slightly compressed laterally, the ribs evident, often winged. IL, Ultimate leaf segments more or less linear or very narrowly elliptic, mostly 0.5-3 mm WId OMe TtIFe 1 PROts. < wecred hc. eae ae 2 — Ultimate leaf segments (at least some) elliptic or broader, some usually over 3 mm wide, sometimes toothed orlobed............... 3 Umbels mostly solitary, occasionally 2, rarely 3, never opposite; rays 0.5—3.6 cm long; peti- oles 1.2-13.5 cm long; leaf blades 3-19 em long; plants 10—45 (64) cm tall, of the Uinta Mountainssaer eee ere L. tenuifolium Umbels 2-5 or more, the lateral ones occa- sionally opposite or whorled; rays 2.5—6.5 (8) cm long; petioles 8—32 cm long; leaf blades (9) 12-30 cm long; plants (40) 60-100 cm tall, of central Utah and western Uinta Mountains L. filicinum Umbels 2 or 3, the lateral 1 or 2 alternate, subtended by much reduced leaves; plants of the Raft River Mountains L. grayi — Terminal umbel subtended by often opposite or whorled umbels and 1—3 or more alternate umbels from the axils of reduced or well de- veloped leaves; plants widespread ... L. porteri Ligusticum filicinum Wats. Fernleaf Ligus- ticum. Plants (4.5) 6-13 dm tall, aromatic, glabrous, from a heavy taproot with a simple or branched crown, the crown clothed with fibrous persisting petiole bases; leaves basal and 1-3 cauline, ternate-pinnately 3 times dissected, with 5—6 (7) opposite pairs of lateral primary leaflets, basal petioles 8-32 cm long, blades (9) 12-30 cm long, ovate in outline, lowest pair of primary leaflets 1/2—3/4 as long as the leaf blade, on petiolules 2.5-10 cm long; ultimate leaf segments 1-18 mm long, 0.75—2.5 (3) mm wide, linear, very narrowly elliptic or narrowly deltoid, entire or bifid or trifid; peduncles (5) 10-17 (23) cm long; ter- minal umbel subtended by. 1-3 smaller um- bels, the lateral ones arising from axils of leaves and alternate or the upper ones not from leaf axils and opposite or rarely 3 per node; involucre lacking; rays 7-27, 2.5-6.5 (8) cm long; involucels of 1-3 linear separate usually deciduous bractlets to 5 mm long; pedicels 4—12 mm long; petals white; stamens whitish; styles ca 0.5 mm long; fruit 5-8 mm long. Tall forb, aspen, sagebrush-grass, forb- grass, Douglas fir, and spruce-fir communi- 90 ties at (1,920) 2,377 to 3,110 m in Cache, Duchesne, Juab, Morgan, Sanpete, Summit, Tooele, Utah, and Wasatch counties; Idaho, Montana, Utah and Wyoming; 46 (xx). This taxon is rather easily confused with L. porteri (q.Vv). Ligusticum grayi Coult. & Rose Grays Ligusticum. Plants 3-6 (9.5) dm tall, glabrous, aromatic, from a stout taproot with simple or branched crown, the crown clothed with fibrous persisting petiole bases; leaves basal and usually 1-3 much reduced cauline ones, ternate-pinnately twice compound and then pinnatifid with (2) 3-5 opposite pairs of lateral primary leaflets, petioles (2.5) 4-34 cm long, blades 4-26 cm long, ovate in outline, lowest pair of primary leaflets 1/2—3/4 as long as the blade, on petiolules 1-6.5 cm long, the larger secondary leaflets pinnatifid with the larger lobes again bilobate or trilobate; pe- duncles 2—55 (90) cm long; terminal umbel subtended by 1-2 alternate umbels arising from the axils of much reduced leaves; involu- cre lacking or rarely of 1 linear mostly decidu- ous bract to about | cm long; rays 8—18, 1.2—4 cm long; involucels lacking or of 1-5 linear bractlets to 4.5 mm long; pedicels 4-10 mm long; petals white; stamens whitish; styles 0.8—1.1 mm long; fruit 4-6 mm long. Forb- grass and fir communities and snowflush areas at 2,650 to 2,900 m in the Raft River Moun- tains, Box Elder County; Washington to Cali- fornia and east to Idaho and northwestern Utah; 3 (iii). Ligusticum porteri Coult. & Rose Southern Ligusticum. [L. brevilobum Rydb., type from the Aquarius Plateau]. Similar to L. filicinum, but leaves with broader ultimate segments, these (1.5) 3—8 mm wide, and with the termi- nal umbel often subtended by a whorl of 3-8 lateral umbels, and occasionally with up to 12 or more umbels, but sometimes with the lat- eral umbels only 2 and opposite, but not alter- nate. Sagebrush, oak, aspen, Douglas-fir, spruce, fir, and occasionally in open forb-grass communities at 2,255 to 3,171 m in Beaver, Carbon, Duchesne, Garfield, Grand, Iron, Juab, Kane, Millard, Piute, San Juan, San- pete, Sevier, Uintah, and Utah counties; southern Wyoming to northern Mexico, west to Idaho and Arizona; 58 (x). Plants of this taxon are sometimes mistaken for Co- nioselinum scopulorum (q.v.). The separation GREAT BASIN NATURALIST Vol. 46, No. 1 of L. porteri from L. filicinum is made diffi- cult by a rather extensive overlap in distribu- tion and lack of definitive morphology from Utah County south to Sevier County. Other- wise the ranges of the two taxa are essentially discrete in Utah, but occasional specimens from scattered locations throughout the state would be difficult to place without location data. Ligusticum tenuifolium Wats. Small Ligus- ticum; Slender-leaf Ligusticum. [L. filicinum var. tenuifolium (Wats.) Mathias & Const. ]. Plants 11-64 cm tall, glabrous mildly aromatic from a taproot, the crown more or less cov- ered by short shredded old leaf bases; leaves basal and sometimes 1 or 2 cauline, ternate and then 2—3 times pinnate with 5—7 pairs of lateral primary leaflets; petioles 1.2-13.5 em long; blades 3-19 cm long, completely dis- sected, ovate in outline; lowest pair of primary leaflets about 1/2 to 2/3 as long as the blade, on petiolules (0.5) 1-4 cm long, the upper pri- mary leaflets progressively reduced, the ulti- mate segments 2-9 mm long, 0.5-1.5 (2.5) mm wide; scapes or peduncles 10—45 (61) em long; involucre lacking; umbel solitary or the terminal one sometimes subtended by a lat- eral | (very rarely 2) that usually arises from the axil of a reduced leaf; rays 6—15, 0.5—3.6 cm long; involucels lacking or of 1—3 filiform- linear bractlets to 3 mm long; pedicels 2—4 mm long; calyx obsolete; petals about 1 mm long, white, sometimes tinged with light pur- ple in age; stylopodium evident, conic; styles 0.5—0.8 mm; fruit about 3-5 mm long. Moist and wet meadows, along streams in lodgepole pine and Engelmann spruce woods at 2,440 to 3,420 m, common across the Uinta Moun- tains, in Daggett, Duchesne, Summit, Uin- tah, and Wasatch counties; northeast Oregon to western Montana south to Colorado and Uinta Mountains of Utah; 37 (xviii). Through a series of features (none of which are exclu- sive), plants of L. tenuifolium are readily dis- tinguished from those of L. filicinum. The two taxa are sympatric in the western Uinta Mountains, where somewhat intermediate specimens occur, but the range of overlap is small and few specimens seem intermediate. Lomatium Raf. Plants perennial, acaulescent or caulescent, occasionally with a short pseudoscape, glab- January 1986 rous or pubescent, from a slender tap root that sometimes has | or more tuberlike segments, or from a thickened, woody branching cau- dex, sometimes clothed at the base with marcescent material; stems simple or rarely branched and thus peduncles and umbels mostly solitary; leaves pinnate or pinnately to ternate-pinnately compound, sheaths often dilated especially in lower leaves, petioles well developed and distinct or confluent with and poorly differentiated from the sheath, or lacking and the petiolules arising directly from the sheath, ultimate segments extremely variable; involucre lacking or inconspicuous; rays few to many, spreading to ascending, the central ones often shorter and sterile; in- volucel mostly of separate or partly united bractlets, rarely lacking; pedicels slender or stout, the central ones often shorter and ster- ile; petals small, yellow, white, greenish yel- low or purplish; calyx teeth obsolete or small, or conspicuous in a very few species; styles slender, often curved or coiled; stylopodium lacking; carpophore divided to the base; fruit linear to orbicular or obovate, flattened dor- sally, glabrous or pubescent, dorsal ribs fili- form or obsolete or occasionally with rudi- ‘mentary wings at base. Note: The genus is closely related to the genus Cymopterus , and the filiform, wingless, dorsal ribs of the fruit seem the only consistent difference from Cy- mopterus. The dependability of this separa- tion is somewhat weakened by the tendency for lack of dorsal wings in some taxa of Cy- mopterus . 1. Leaves once-pinnate and/or the ultimate seg- ments over 15 mm long and less than 50 per leaf; plants glabrous and/or petals yellow WAIST OS] SY BOR a den Sa rey bes ER oe Seen 2 — Leaves more than once-compound, the ulti- mate segments not over 15 mm long and mostly over 50 per leaf, or if a few ultimate segments over 15 mm long or less than 50 per leaf then plants pubescent and petals white (1). Leaves once-pinnate or partly bipinnate, the leaflets sessile and more or less confluent with the rachis; plants from stout, more or less woody caudices, clothed at the base with long-persisting leaf bases, from thé southern Donte ss tated ener ieee lacie aelsn ete: 3 GOODRICH: UTAH FLORA, APIACEAE 91 Leaves more than once-compound, the pri- mary leaflets mostly with well-developed petiolules, not confluent with the rachis: (plants from taproots or small caudices, not much if at all clothed at the base with long- persisting leaf-bases, from the northern 1/2 of the state except in L. nuttallii and then plants keyedinothways) itera aseee eee ee 6 Leaflets lanceolate to elliptic, 2-12 mm wide, some always over 5 mm wide; plants of Grand and northern San Juan counties ... L. latilobum Leaflets linear, not over 4 mm wide: distribu- tion not as above Leaves with 1—7 elongate, terete leaflets that simulate the rachis in diameter and shape, these 1-18 cm long, at least some commonly over 5 cm long in each leaf; calyx teeth green- ish, acute, somewhat persistant, about 1 mm long; fruit 8-12 mm long; plants of Emery, Garfield, and eastern Sevier counties, mostly below2lGimbon rary acer L. junceum Leaves either with more than 7 leaflets and/or leaflets less than 5 cm long and more or less flattened and at least slightly wider than the rachis; calyx teeth not over 0.6 mm long, scar- ious or greenish; fruit various; distribution not as above, mostly of higher elevations ....... 5 Plants 2—12 (17) cm tall, of Garfield and Iron counties; fruit 4-7 mm long, leaflets 3-13 per leaf, 0.2-1.5(2)cmlong......... L. minimum Plants 15—30 cm tall or taller, not known from the above counties; fruit 5-15 mm long; leaflets sometimes more than 13 per leaf, sometimes over2cmlong ........ L. nuttallii Ultimate leaflets ovate to nearly orbicular ora few broadly elliptic, less than 3 times as long as wide, at least some dentate-toothed on the upper 1/4; rays (4) 8-19 cm long; peduncle often swollen just beneath the umbel; plants of the Deep Creek Mountains and western BoxailderCountyasne eee oee L. nudicaule Ultimate leaflets linear to elliptic, 3 or more times longer than wide, entire; rays 0.5—10 em long: peduncle not swollen just beneath the umbel: distribution not as above ........ i Plants caulescent, glabrous; peduncles to 13 cm long; involucel lacking; root very slender, sometimes with | or more globose or fusiform tuberous segments; ultimate leaflets and rays WET UMC WEll 653 oo04c00000000% L. ambiguum Plants acaulescent or if caulescent then the peduncles mostly over 13 cm long and plants puberulent; involucels present, to 1 cm long; root not as above; ultimate leaflets and rays WAIMOUS ococooboovcvcrpccston00g900000N0 8 8(7). 10(9). 11(10). 12(11 GREAT BASIN NATURALIST Plants from thickened woody branching cau- dices, glabrous, strongly aromatic, often of rocky places, escarpments, or semibarrens; caudex often clothed with old long-persisting leaf bases; leaves strictly basal, the ultimate leaflets 0.3-5(6.5) cm long; 0.5—2(4) mm wide; lateral wings of the fruit to 1 mm wide L. nuttallii Plants from taproots or small caudices, pu- berulent at least on the peduncle, not strongly aromatic, mostly growing in loamy soil; old leaf bases lacking or weakly persisting; leaves basal and sometimes 1-3 cauline, the ulti- mate leaflets 1-13 cm long, 1—6(15) mm wide; lateral wings of the fruit 1-2 mm wide L. triternatum Larger mature leaves with blades (10) 15-30 cm long, ternate-pinnately compound, the larger ultimate segments 2-3 mm _ wide; plants 30—130 cm tall, peduncles fistulose, (3) 4—6 (10) mm thick at the base.... L. dissectum Blades of leaves 2-11 cm long or iflonger then either not at all ternate or with ultimate seg- ments not over | mm wide; plants rarely over 50 cm tall; peduncles fistulose or not, often less than 4 mm thick Plants pubescent; petals white or yellow ... 11 Plants glabrous or at most scabrous; petals yellow or if white then plants keyed both ways Ovaries and fruit glabrous or occasionally somewhat scabrous; plants of the central part and the northern 1/2 of eastern Utah (rare specimens of L. nevadense from southern Utah will key here) Ovaries and young fruit rather densely pubescent older fruit sometimes glabrous but often retaining some hirtellous hairs; plants of the central, western, and southern parts of Utah Bractlets of the involucel about 10, the longer ones 4-10 mm long, pubescent; herbage more or less villous; leaves with about 4 oppo- site pairs of primary lateral leaflets, the lowest pair sessile or on petiolules to 1 cm long; mature fruit 9-12 (15) mm long Bractlets of the involucel 1-5, 1-4.5 mm long, glabrous; herbage glabrate to puberu- lent; leaf blades with (3) 4—6 opposite pairs of lateral primary leaflets, the lowest pair on petiolules 1-3 cm long; mature fruit 5-8 (11) mm long (rare specimens of L. nevadense with glabrous ovaries will key here, but then the primary leaflets all sessile or nearly so) ..... L. juniperinum 13(11). 15(10). 16(15). 17(16). Vol. 46, No. 1 Longer ultimate segments of leaves 5—27 mm long, some often over 1 mm wide; leaves mostly with 4 opposite pairs of sessile or nearly sessile lateral primary leaflets; root slender, usually with a deep-seated globose or fusiform tuberous segment; bractlets of the involucel glabrous or sparingly pubescent; petals white L. nevadense Ultimate segments of leaves 1-5 mm long, to 1 mm wide; leaves mostly with 5-8 opposite pairs of sessile or petiolulate lateral primary leaflets; root not as above; bractlets of the involucel pubescent; petals yellow, rarely white Petals and anthers yellow; leaves often con- spicuously ternate-pinnately compound, the lowest pair of primary leaflets nearly sessile or on petiolules to 5 cm long, some of these often arising directly from a dilated sheath with the petiole lacking or only to 2.5 cm long; plants common L. foeniculaceum Petals white; anthers purple or whitish; leaves pinnately or scarcely ternate-pinnately com- pound, the lowest pair of primary leaflets ses- sile or on petiolules to 1.5 cm long, these rarely arising directly from a dilated sheath, but rather from a petiole 1-4 (6) cm long; plants apparently rare, known only from west- ern Millard County L. ravenii Lowest pair of primary leaflets less than 1/3 as long as the leaf blade, sessile or on petiolules to 1.2cm long; leaves strictly basal, the blades never arising directly from dilated sheaths; plants clothed at the base with old long-per- sisting leaf bases, of the southern 1/2 of the state Lowest pair of primary leaflets 1/3 to 3/4 as long as the leaf blade, sessile or on petiolules to 7.5 cm long; leaves basal and sometimes a few on the lower part of stems, the blades often arising directly from dilated sheaths; (plants not clothed at the base with old leaf bases and of the northern 1/2 of the state except sometimes in distribution of L. grayi) Mature pedicels 1-10 mm long; fruit 4-9 mm long; leaves and peduncles scabrous, the blades 2-7 cm long; plants of the Great Basin and Washington County The longer mature pedicels 10-20 mm long; fruit (6) 8-20 mm long; leaves and peduncles glabrous or plants of the Colorado Basin ... . Leaf blades 3-7 cm long, the ultimate seg- ments 2—4 mm long; fruit 8-10 mm long; rays 4—6, 1-3 cm long; plants densely scabrous, 10-15 cm tall, of western Colorado, to be expected in Utah in extreme eastern Grand and San Juan counties L. eastwoodeae (Coult. & Rose) Macbr. L. scabrum | 17 | January 1986 GOODRICH: UTAH FLORA, APIACEAE 93 —= Leaf blades 7-24 cm long, the ultimate seg- ments 1-15 mm long; fruit to 20 mm long; plants glabrous, glaucous, 8—40 cm tall, of the Colorado Basin and Washington County ... Sc NA Ea iNet ata ark 8 RRS Rae oe See area L. parryi 18(15). Bractlets of the involucel broadly elliptic to obovate, to 3 mm wide; pedicels 1-2 mm long; ultimate segments of leaves 2-13 mm long, 0.5-—4 mm wide, dimorphic, plants of northwestern Box Elder County ...... L. cous — Bractlets of the involucel linear to subulate, not over 1 mm wide; pedicels 2-18 mm long; ultimate segments of leaves 1-7 mm long, 0.2-1.5 mm wide, not much if at all dimor- [DLOVIS) 6.5 is sie Zpca Sant O sain Re A Ona ak dearest ena 19 19(18). Petals white or cream; ultimate segments of leaves 0.5—1.5 mm wide; plants not aromatic; pedicels 3-16 mm long; leaves with 3—6 op- posite pairs of lateral primary leaflets; rare AA VOWS KOANS oos6¢o0cbcdsoe L. juniperinum — Petals yellow when fresh; ultimate segments of leaves 0.6—0.2 mm wide; plants strongly aromatic, glabrous; pedicels various; leaves Valls OUS tracer kes sheen oceans een ae ee 20 20(19). Fruit 2-4 mm wide, the wings 0.4—0.6 mm wide; pedicels 2-5 mm long; rays very un- equal; inflorescence open during flowering; leaves with 5-6 opposite pairs of lateral pri- mary leaflets; plants not clothed at the base with old leaf bases, from a slender root, this often with 1 or more tuberlike segment .... siSA Solo ee OTR a Ne Se ee. L. bicolor = Fruit 5-8 mm wide, the wings about 1.5—2 mm wide; pedicels 5-18 mm long; rays sube- qual; inflorescence congested at flowering time; leaves with about 7—10 opposite pairs of lateral primary leaflets; plants often clothed at the base with old long-persisting leaf bases, from a more or less woody branched caudex Sh GickiseeReesRueo es Cas cxe RS ies alin L. grayi Lomatium ambiguum (Nutt.) Coult. & Rose Wyeth Biscuitroot. [Eulophus ambiguus Nutt.]. Plants caulescent, 10-40 cm tall, glabrous without persisting leaf bases; root very slender, sometimes with 1 or more gla- | bose or elongate tuberlike segments; leaves 'ternately or ternate-pinnately compound, petioles to 2 cm long or lacking and blades arising from a dilated sheath 1.5—4 cm long, _ blades 4-15 cm long, ovate in outline, lowest _ pair of primary leaflets mostly over 1/2 as long ! as the blade with petiolules 1—4 cm long, ulti- mate segments about 15—45, 0.3—9 cm long, 1-4 mm wide, often very unequal in the same leaf; peduncle 2.5—-13 cm long; involucre lack- ing; rays 0.5—6.5 cm long, very unequal in the ‘same umbel; involucel lacking; pedicels 2-12 i: mm long; petals and stamens yellow, fading in ‘herbarium specimens; styles about 1 mm long; fruit 8-10 mm long, 2—3 mm wide, lat- eral wings about 0.5 mm wide, dorsal ribs filiform. Sagebrush and mountain brush com- munities at 1,525 to 1,980 m in Cache, Salt Lake, Utah, and Weber counties; Washington and adjacent British Columbia to Montana and south to Utah and Wyoming; 14 (0). The narrow fruits, unequal rays, and slender root with globose or elongate tuberlike segments are features also found in plants of L. bicolor, but the larger leaf segments are conspicuously different from the very narrow and shorter ones of plants of L. bicolor. Lomatium bicolor (Wats.) Coult. & Rose Wasatch Biscuitroot. [Peucedanum bicolor Wats. |. Plants 10—50 cm tall, acaulescent or caulescent, aromatic, glabrous, without per- sisting leaf bases or these few and weakly per- sisting; from a slender taproot, this often with one or more tuberlike segments; leaves ter- nate-pinnately decompound, petioles mostly lacking and the blades arising directly from a dilated sheath, blades 4—12 cm long, ovate in outline, finely and completely dissected, the lowest pair of primary leaflets mostly over 1/2 as long as the leaf blade, with petiolules 2.5-6 cm long, each with 5—6 opposite or offset pairs of secondary leaflets, ultimate segments mostly over 300, 1—4 (6) mm long, 0.2—0.6 mm wide; peduncle 10—28 cm long; rays 3-12 (20), 1-8 (11) cm long, very unequal in the same umbel; involucel lacking or of 1—8 linear separate bractlets; pedicels 2-5 mm long; petals and stamens yellow; styles about 1 mm long; fruit 8-11 mm long, 2—4 mm wide, con- gested; lateral wings 0.4—0.6 mm wide, dorsal ribs filiform. Sagebrush, mountain brush, as- pen, and meadow communities at 1,525 to 2,438 m in Cache, Morgan, Rich, Salt Lake, and Weber counties; southeastern Idaho and northern Utah; 25 (0). This taxon has been included in L. leptocarpum (T. & G.) Coult. & Rose. Plants of the two taxa differ only in size of ultimate segments of leaves. Perhaps they are not distinct at the species level. L. bicolor has priority at the species level. Lomatium cous (Wats.) Coult. & Rose Cous Biscuitroot. [Peucedanum cous Wats. |. Plants 5-15 (25) em tall, not or weakly aromatic, glabrous (ours), from a globose or fusiform tuberous root, this sometimes deep-seated and giving rise to a subterranean pseu- doscape; leaves basal and sometimes 1-2 cauline on the lower 1/3 or rarely to midlength 94 of the stem, 2—3 times pinnately or ternate- pinnately compound, blades 4—8 (11) cm long, mostly borne on dilated sheaths with the petioles obsolete or short or some leaves origi- nating from the deep-seated tuber and then with etiolated, mostly subterranean petioles to 11 cm long, lowest pair of primary leaflets about 1/2 to over 3/4 as long as the leaf blade, sessile or with petiolules to 42 mm long, ulti- mate segments or lobes 2-13 mm long, 0.5—4 mm wide, as many as 200 or more, linear to elliptic; peduncles 1-7, 3-18 cm long; involu- cre lacking or of a solitary bract to 7 (10) mm long; rays 6-15, 0.4-5 cm long, strongly di- morphic in the same umbel; bractlets of the involucel about 6—10, 3-5 mm long, to 3 mm wide, broadly elliptic, ovate or obovate, greenish, sometimes with yellowish or scari- ous margins; pedicels 1-2 mm long; calyx teeth obsolete; petals and stamens yellow when fresh, fading to white in herbarium specimens; styles about 1.5 mm long; fruit 6—9 mm long, the lateral wings to about 1 mm wide, the dorsal ribs filiform or very obscurely winged. Sagebrush-grass communities at 2,440 to 2,560 m, in the Grouse Creek and Raft River Mountains, Box Elder County; Or- egon to Montana, south to northwestern Utah and northeastern Nevada; 2 (ii). Lomatium dissectum (Nutt.) Mathias & Const. Giant Lomatium. [Leptotaenia dis- secta Nutt.]. Plants 30-130 cm tall, mostly short caulescent, puberulent or rarely glabrous, from a woody thickened taproot or caudex, without old leaf bases or these short- persistent and soon shredding; leaves pin- nately or ternate-pinnately decompound, with 5—9 opposite or offset pairs of primary leaflets, or the upper cauline leaves much reduced, petioles 3-20 cm long, often lacking on cauline leaves and then the blades sessile on a dilated sheath, blades 10-30 cm long or smaller on cauline leaves, ovate in outline, the lowest pair of primary leaflets usually over 1/2 as long as the leaf blade, with petiolules 2.5-12 cm long, ultimate segments numer- ous, 1-12 mm long, 0.5—3 mm wide; pedun- cles 15—50 (90) cm long; involucre lacking or rarely of 1-3 rather quickly deciduous bracts; rays 9-27, 2—7 (12) cm long; bractlets of the involucel 3-6 mm long, or occasionally much longer and foliaceous; pedicels 3—10 (15) mm long; petals and stamens yellow, yellow- GREAT BASIN NATURALIST Vol. 46, No. 1 green, or purplish; styles about 1.5 mm long; fruit 9-15 (20) mm long, 6—10 mm wide, lat- eral wings about 1-2 mm wide, dorsal ribs filiform. Sagebrush, pinyon-juniper, oak- maple, aspen-fir, riparian, and rarely grease- wood-desert shrub communities, from rock outcrops to deep loamy soil, at 1,280 to 2,650 (3,170) min Beaver, Box Elder, Cache, Duch- esne, Iron, Juab, Millard, Morgan, Rich, Salt Lake, Sanpete, Summit, Tooele, Uintah, Utah, Washington, and Weber counties; southern British Columbia and Alberta south to southern California, Arizona and Colorado; 91 (xiii). Utah materials are referable to var. eatonii (Coult. & Rose) Crongq. |Leptotaenia eatonii Coult. & Rose]. Sometimes the leaves are mistaken for those of Ligusticum porteri, but the mostly solitary umbel is strikingly dif ferent from the usually opposite or whorled lateral umbels in addition to the terminal one in the Ligusticum. Lomatium foeniculaceum (Nutt.) Coult. & Rose Desert-parsley. [Ferula foeniculacea Nutt. ]. Plants 5-25 (38) cm tall, acaulescent, densely pubescent throughout, from a more or less branching caudex and deep taproot, often clothed at the base with persisting leaf bases; leaves ternate-pinnately dissected, with 6-8 opposite pairs of lateral primary leaflets, petioles to 2.5 cm long or lacking and the blade arising from a dilated sheath, blades 2-13 cm long, completely and finely dis- sected, ovate in outline, the lowest pair of primary leaflets over 1/2 as long as the blade, sessile or with petiolules to 5 cm long, ulti- mate segments numerous, often more than 500, 1-3 (5) mm long, 0.5—1 (2.5) mm wide; peduncles 4—30 cm long; rays 5-20, 0.2—7 em long; bractlets of the involucel 2—5 (6) mm long, separate or united at the very base, lin- ear; pedicels 2-12 mm long; petals and an- thers yellow (rarely white) when fresh and mostly remaining yellow for many years in herbarium specimens or occasionally turning purplish; styles about 1.5—2 mm long; fruit 5-10 mm long, 3-7 mm wide, lateral wings about 1—2 mm wide, dorsal ribs filiform. Sage- brush (mostly black sagebrush), pinyon-ju- niper, and mountain brush communities at 1,250 to 2,635 m in Beaver, Daggett, Box Elder, Emery, Juab, Kane, Millard, Sanpete, and Tooele counties; Manitoba to Missouri and Texas, west to southeastern Oregon and January 1986 California; 71 (xvi). Most Utah materials are referable to var. macdougalii (Coult. & Rose) Cronq. Some plants from western Utah with ciliolate petals are referable to var. fimbriata (Theobald) Boivin, but this feature seems variable in some populations. Lomatium grayi (Coult. & Rose) Coult. & Rose Milfoil Lomatium. [L. millefolium (Wats.) Macbr. |. Plants (8) 15—40 (80) cm tall, acaulescent or subcaulescent, strongly aro- matic, glabrous, from a simple or branched caudex and thick taproot often clothed at the base with old, mostly shredded fibrous leaf bases; leaves ternate-pinnately dissected, with about 7-10 opposite pairs of lateral pri- mary leaflets, petioles to 14 cm long or lacking and the blades arising from a dilated sheath 1-16 cm long, blades 7—16 (2) cm long, finely and completely dissected, ovate in outline, the lowest pair of primary leaflets from 1/2 to as long as the blade, with petiolules 1—7.5 cm long, ultimate segments several hundred or a thousand or more, extremely fine, 1—3 (6) mm long, 0.2—0.3 mm wide; peduncle 10—45 (70) cm tall; rays 10-26, 1.5-6 (8) cm long; bractlets of the involuces 3-5 mm long, lin- ear, separate or united at the base; pedicels 5-13 (18) mm long; petals and stamens yellow when fresh, soon fading whitish when dried; styles 1.5—2.5 mm long; fruit 6-12 mm long, 5-8 mm wide, lateral wings about 2 mm wide, dorsal ribs filiform. With 2 intergrading but more or less geographically distinct varieties. 1. Fruit 6—9 (10) mm long, the lateral wings to about 1.5 mm wide; leaves rather openly dis- sected, with a few hundred ultimate seg- ments; plants usually with greater buildup of old leaf bases that persist a little longer before shredding than in the following variety, aver- aging smaller, 8—20 (35) cm tall, of the west- ern tier of counties from Box Elder County south to Beaver County aii: Man Jaen Ok ea ae L. grayi var. depauperatum — Fruit 8-12 mm long, the lateral wings to about 2 mm wide; leaves with congested and numerous ultimate segments, these several hundred ora thousand or more; plants usually with less buildup of old persistent leaf bases, these usually shredding within a year, averag- ing larger, 15—40(80) cm tall, of more easterly distribution and only in the eastern 1/4 of the western tier of counties where more or less transitional with the preceding variety ..... L. grayii var. grayi (Jones) Mathias var. depauperata Var. depauperatum [Cogswellia millefolia GOODRICH: UTAH FLORA, APIACEAE 95 Jones]. Desert shrub, pinyon-juniper, and mountain brush communities at 1,525 to 2,835 m in Beaver, Box Elder, Juab, Millard, and Tooele counties; western Utah and adja- cent Nevada; 58 (xiii). Var. grayi Sagebrush, pinyon-juniper, mountainbrush, ponderosa pine, and Dou- glas-fir communities at 1,340 to 2,745 m in Box Elder, Cache, Daggett, Davis, Du- chesne, Grand, Juab, Morgan, Rich, Salt Lake, San Juan, Sanpete, Summit, Tooele, Uintah, Utah, and Weber counties; Washing- ton to northeastern Nevada and east to Idaho and southwestern Colorado; 121 (xxiii) Lomatium junceum Barneby & N. Holm- gren Rush-lomatium. Plants (6) 10—37 cm tall, acaulescent, glabrous, from a simple to much branched woody caudex, clothed at the base with old petioles, some of which often persist for a few years before shredding; leaves rush- like, trifid or pinnatifid or rarely reduced to a petiole and a linear bladeless rachis, with 1-7 linear segments, petioles 3-15 mm long with a short sheath at the base, blades 3-17 cm long, the segments 1-18 cm long, about 1-2 mm wide, terete and similar to the rachis and petioles in diameter; peduncles 5—25 cm long; rays of umbels 6-13, 1.5—3 cm long; bractlets of the involucel 1.5—3 mm long, separate or united at the base, linear; pedicels 4-11 mm long; calyx teeth to about 1 mm long, acutish, somewhat persistent; petals and stamens bright yellow or cream, quickly fading to white when frozen or dried; styles about 2-3 mm long; fruit 8-12 mm long, 5—7 mm wide, lateral wings 1-2 mm wide, dorsal ribs fili- form. Desert shrub, sagebrush, pinyon-ju- niper, ponderosa pine, and Douglas-fir com- munities at 1,615 to 2,485 m in Emery, Garfield, Sevier, and Wayne counties; en- demic; 21 (iii). Lomatium juniperinum (Jones) Coult. & Rose Juniper Lomatium. [Peucedanum ju- niperinum Jones]. Plants 8-32 cm tall, acaulescent or with 1—3 leaves on the lower part of the stems, more or less hirtellous, occasionally glabrate and rarely glabrous, of- ten with a short pseudoscape, from a taproot with simple or sparingly branched crown, not clothed at the base with old leaf bases or these weakly persisting; leaves ternate-pinnately dissected, with (3)4—6 opposite or offset pairs of lateral primary leaflets, petioles to 8 cm 96 GREAT BASIN NATURALIST long or lacking and the blades arising directly from dilated sheaths 1-4 cm long, blades 2.5—8 (11) cm long, ovate in outline, the low- est pair of lateral primary leaflets 1/3 to about as long as the leaf blade, with petiolules to 3 cm long, the ultimate segments about 50-400, 1-7 mm long, 0.7-1.5 mm wide; peduncles 6—29 cm long; rays of the umbel 3-12, 1-8 cm long; bractlets of the involucel about 1-5, 1—4.5 mm long, linear, separate or united at the base; pedicels 3-16 mm long; petals white, cream, or yellow; anthers white, ochroleucus, purple, or yellow; styles about 1—2 mm long; fruit 5—8 (11) mm long, 3-6 mm wide, glabrous, or scabrous to sparsely hirtel- lous especially when young; lateral wings 0.5-1.5 mm wide, the dorsal ribs filiform. Sagebrush, pinyon-juniper, forb-grass, as- pen, Douglas-fir, and alpine (Mt. Nebo) com- munities at 1,830 to 3,230 m in Carbon, Daggett, Duchesne, Grand, Juab, Sanpete, Summit, Uintah, Utah, and Wasatch coun- ties; southwestern Wyoming and adjacent Idaho to Utah and extreme northwestern Col- orado; 50 (xviii). Quite variable as to color of petals and anthers. Plants with yellow petals and anthers are known only from the west and north side of the Uinta Mountains and West Tavaputs Plateau. Those with white petals and white to purplish anthers are found in the Wasatch Mountains, south slope of the Uinta Mountains, Tavaputs Plateau, and to the north end of the Wasatch Plateau. Plants are quite pubescent except a few specimens from the Wasatch Mountains and Wasatch Plateau. The plants are sometimes confused with those of L. nevadense. The following key and dis- cussion should help separate the two taxa. Ihe Leaves ternate-pinnately compound, the lowest pair of primary leaflets on petiolules mostly 1-3 cm long; plants of northern Sanpete County and northward and_ eastward L. juniperinum — Leaves pinnately compound, the lowest pair of primary leaflets sessile or on petiolules mostly less than 1 cm long; plants of Millard County and southward and westward ............. In Utah the difference is more obvious be- cause most of the Utah plants of L. nevadense have: usually densely hirtellous or puberulent ovaries and young fruits, with the pubescence often remaining in some of the mature fruits; Vol. 46, No. 1 ultimate segments of leaves about 20—100, dimorphic, with the larger ones 7-27 mm long, and slender roots with fusiform tuberous enlargements. Plants of L. juniperinum have: glabrous or scabrous ovaries and young fruits, with the scabrousity lacking or scattered in mature fruits; ultimate segments of leaves 50—400, 1-7 mm long, and the root does not have tuberous enlargements. Sometimes the plants are also confused with those of Cy- mopterus lemmonii. (q.v.) Lomatium latilobum (Rydb.) Mathias Canyonlands Lomatium. [Cynomarathrum latilobum Rydb.]. Plants (6) 10—30 cm tall, acaulescent, glabrous, from a_ branched woody caudex, clothed at the base with old persistent leaf bases; leaves pinnate with 3—4 (5) pairs of lateral leaflets; petioles 2-16 cm long; blades 1-10 cm long, oblong in outline, leaflets 1-4 cm long, 2-12 mm wide, sessile, entire or a few bifid or trifid; peduncles 4-27 cm long; rays of the umbel 4-13, 0.5-2 cm long; bractlets of the involucel 2-15 mm long, 0.5-2 mm wide, linear or elliptic, separate; pedicels 1-4 mm long; calyx teeth 1-1.5 mm long, acute; petals yellow when fresh, drying white; styles 2-3 mm long; fruit 8-12 mm long, 3-7 mm wide, the lateral wings about 1 mm wide, the dorsal ribs filiform. Pinyon-ju- niper communities, and in hanging gardens, sandstone ledges, and sandy soil, at ca 1,525 m, in southern Grand and adjacent San Juan counties, also Mesa County, Colorado; 17 (i). Lomatium macrocarpum (H. & A.) Coult. & Rose Big-seed Lomatium. [Ferula macro- carpa H. & A.]. Plants 12-30 cm tall, acaules- cent or subcaulescent with leaves mostly on | the lower 1/4 of the stem, more or less tomen- | tose-villous or glabrate, from a thickened tap- | root with a simple or sparingly branched | crown with few or no persisting leaf bases; leaves pinnately or ternate-pinnately dis- sected, with about 4 opposite pairs of lateral primary leaflets, petioles often long tapering — into a dilated sheath, the petiole and sheath | about 3-6 cm long, blades 3-6 cm long, ovate in outline, the lowest pair of primary leaflets | 1/2 to 3/4 as long as the leaf blade, sessile or — with petiolules to 1 cm long, ultimate seg- ments about 30—300 or more, 1.5—6 mm long, | 0.5-2 mm wide, elliptic or linear; peduncles | 8-26 cm long; rays of the umbel 6-18, 1-4 (6.5) cm long; bractlets of the involucel about | { | January 1986 10, 2-10 mm long, separate or united at the base, pubescent; pedicels 2—5 mm long; calyx teeth to about 0.5 mm long; petals white or purplish in age; anthers white; styles about 2-3 mm long; fruit 9-12 (15) mm long, 4—5 mm wide, glabrous, the lateral wings 1—1.5 mm wide, the dorsal ribs filiform. Desert shrub, sagebrush, and pinyon-juniper com- munities at 1,480 to 2,550 m in Daggett, Juab, Millard, Sanpete, Tooele, and Uintah coun- ties; southern British Columbia to California and east to Manitoba and Colorado; 29 (xi). Lomatium minimum (Mathias) Mathias Least Lomatium. [Cogswellia minima Mathias]. Plants 2-12 (17) em tall, acaules- cent, glabrous or scabrous, from a branched caudex, the caudex branches clothed with persisting leaf bases; leaves once-pinnatifid or rarely trifid, with (3) 5-9 (13) segments, peti- oles to 2 cm long, blades 1—2.5 cm long, the segments 2—15 (20) mm long, 0.5—2 mm wide; peduncles to 10 (16) cm long; rays of the um- bel 3-6, 0.3—2.3 (3.2) cm long; bractlets of the involucel 2—4 mm long, linear-subulate, sepa- rate; pedicels 1-3 mm long; calyx teeth to 0.6 mm long, acute, greenish or purplish in age with scarious margins; petals and stamens yel- low, drying to cream; styles about 1.5-2 mm long; fruit 4-7 mm long, 3—4 mm wide, lateral wings 0.5—1 mm wide, dorsal ribs mostly fili- form. Forb-grass, ponderosa pine, and bris- tlecone pine communities, often on exposed ridges and raw escarpments, often on lime- stone at 2,165 to 3,170 m in Garfield, Iron, and Kane counties; endemic; 20 (0). Appear- ing much like a diminutive form of L. nuttallii var. alpinum. Lomatium nevadense (Wats.) Coult. & Rose Nevada Lomatium. [Peucedanum nevadense Wats. |. Plants 10—36 cm tall, acaulescent or with | or 2 leaves on the lower part of stems, more or less pubescent throughout, from a slender root frequently with a fusiform tuber- ous segment, with or without persisting leaf- bases; leaves 2—3 times pinnately compound, with about 4 opposite pairs of lateral primary leaflets, petioles to 7.5 cm long or often lack- ing and the blade sessile on a dilated sheath, _ blades 2.5-9 cm long, ovate in outline; the lowest pair of primary leaflets about 1/2 to nearly as long as the blade, sessile or on peti~ ‘ olules to 5 mm long, the ultimate segments about 20—80, 1-27 mm long, 0.5—3 mm wide; GOODRICH: UTAH FLORA, APIACEAE 97 peduncles 7-33 cm long: rays of the umbel 7-12, sometimes with as few as 3 of them fertile, 1.5-4 cm long; bractlets of the in- volucel 2—3 mm long, lanceolate, linear-ellip- tic, or narrowly obovate; pedicels 4-11 mm long; petals and stamens white; styles about 1—1.5 mm long; fruit 5-10 mm long, 3-7 mm wide, densely puberulent or rarely glabrous when young to glabrate; lateral wings 0.8—2 mm wide, dorsai ribs filiform. Desert shrub, sagebrush, pinyon-juniper, mountain brush, and ponderosa pine communities at 1,524 to 2,285 m in Beaver, Garfield, Iron, Kane, Mil- lard, and Washington counties; Oregon to California east to Colorado and Arizona; 44 (viii). Our materials are perhaps referable to var. parishii (Coult. & Rose) Jepson [Peucedanum parishii Coult. & Rose]. This variety has been keyed as having glabrous fruits, and the Utah materials with pubescent fruits have been referred to as var. nevadense . However, the Utah plants have dimorphic ultimate leaf segments 1-27 mm long. This is a feature of var. parishii. The ultimate leaf segments of var. nevadense are only 2-3 mm long. A specimen from Navajo Mountain, San Juan County (Albee 4463 UT), has uniformly small leaf segments and glabrous fruits. Lomatium nudicaule (Pursh) Coult. & Rose Naked-stem Lomatium. [Smyrnium nudi- caule Pursh]. Plants 20—45 cm tall, acaules- cent, glabrous, from a taproot, without persis- tent leaf bases or these few and weakly persisting; leaves ternate or biternate; with 3-11 distinct leaflets, petioles to 6 cm long, or obsolete and the blades arising from a dilated sheath, blades 4—10 cm long, ovate in outline, leaflets 2-5 cm long mostly 1-5.5 cm wide, ovate or orbicular to reniform, coarsely toothed toward the apex; peduncles 15-27 cm tall, sometimes swollen at the apex; rays of the umbel 7-27, 8—10 cm long; involucel lacking; pedicels 3-10 mm long; petals yellow; styles about 1—2 mm long; fruit 8-12 mm long, 2-5 mm wide, the lateral wings about 0.5 mm wide, dorsal ribs filiform. Sagebrush, pinyon- juniper, and mountain brush communities at 1,585 to 2,530 m in Box Elder, Juab, and Tooele counties; southern British Columbia to central California and east to southwestern Alberta and Utah; 6 (iii). Lomatium nuttallii (Gray) Macbr. Stinking Lomatium. Plants 15—50 cm tall, acaulescent, 98 GREAT BASIN NATURALIST glabrous, strongly aromatic, from a branched caudex, the caudex clothed with persistent leaf bases; leaves pinnatifid to bipinnate or ternate-pinnately compound, petioles gradu- ally expanded into a dilated sheath, with the sheath 2—21 cm long, blades 2-15 cm long, usually oblong in outline, ultimate leaflets or segments about 7—30, 0.3-6.5 cm long, 0.5—2 (4) mm wide; peduncles 12—47 cm long; rays of the umbel (3) 5—12, 1-5 cm long; bractlets of the involucel 3—10 mm long; pedicels 2—10 mm long; calyx teeth about 0.5 mm long, rather scarious; petals and stamens yellow, soon turning pale to white in herbarium speci- mens; styles about 2-3 mm long; fruit 5-15 mm long, 3—5 mm wide, lateral wings about 0.5-1 mm wide, the dorsal ribs filiform or somewhat prominent with rudimentary wings. There are two geographically sepa- rated varieties that are distinct morphologi- cally, but some of the smaller specimens of var. nuttallii are much like those of var. alpinum. They are separated as follows: 1. Fruit 5-8 mm long, the lateral wings ca 0.5 mm wide; pedicels 4-10 mm long; umbels with only 3-6 rays; leaves once pinnatifid with ses- sile segments or some of the lower pairs of segments bipinnatifid; plants of Washington and western Millard counties ............. MEETS PRE St pads L. nuttallii var. alpinum _ Fruit 10-15 mm long, the lateral wings about 1 mm wide; pedicels 2-6 mm long; umbels with up to 12 rays; leaves pinnatifid to ternate- pinnately compound, with the lowest pair of primary leaflets on petiolules (1) 2.5-9 cm long; plants of Sevier and eastern Millard counties and northward . L. nuttallii var. nuttallii Var. alpinum (Wats.) Mathias Pinyon-ju- niper and mountain brush communities at 2,225 to 2,440 m in Millard and Washington counties; western Nevada and southwestern Utah: 4 (0). Var. nuttallii [Peuceda-num graveolens Wats.]. Sagebrush, bullgrass, mountain brush, Douglas-fir, limber pine, and spruce- fir communities, often in rocky places, mostly on limestone and other basic substrates, sometimes in raw snowflush areas, at 1,980 to 3,200 m in Cache, Davis, Duchesne, Millard, Rich, Salt Lake, Sanpete, Sevier, Summit, Tooele, Utah, Wasatch, and Weber counties; western Nevada, and western Wyoming; 85 (Xvili). Vol. 46, No. 1 Lomatium parryi (Wats.) Macbr. Parry Lo- matium. [Peucedanum parryi Wats.; Cogswellia cottami Jones]. Plants 8-40 cm tall, acaulescent, glabrous, from a branched caudex, clothed at the base with persisting leaf bases; leaves bipinnatifid or partly tripin- natifid, with mostly 7—9 opposite pairs of pri- mary leaflets or the upper leaflets simple, petioles 3-16 cm long, terete, often persisting for a few years without shredding, blades 7-24 cm long, lowest pair of lateral primary leaflets 1/10-1/4 as long as the leaf blade, sessile or with petiolules to 1.2 cm long, the ultimate segments mostly 50-150, 1-15 mm long, 1-2 mm wide, acute; peduncles 5-32 cm long; rays of the umbel 8-13, 1-5 cm long; bractlets of the involucels 3-10 mm long, en- tire, tridentate or rarely pinnatifid, spreading to reflexed in age; pedicels 1-2 cm long; petals yellow, turning white in herbarium speci- mens; styles about 2-4 mm long; fruit 6—20 mm long, 5-10 mm wide, the lateral wings 1—3 mm wide, the dorsal ribs filiform. Desert shrub, blackbrush, pinyon-juniper, and mountain brush communities at 975 to 2,320 m in Emery, Garfield,-Grand, Iron, Kane, San Juan, and Washington counties; Utah to eastern California; 65 (v). | Lomatium ravenii Mathias & Const. Raven | Lomatium. Plants 4—23 cm tall, acaulescent, densely hirtellous throughout, from a taproot, with a simple or branched crown, usually | clothed at the base by shredded leaf bases; | leaves ternate-bipinnate or 2—3 times pin-_ nately dissected, with 5—7 (8) opposite pairs of — primary leaflets, petioles to 6 cm long or lack- | ing and blades arising from dilated sheaths to 2 cm long, blades 1.5—8 cm long, finely and completely dissected, the lowest pair of pri- mary leaflets usually over 1/2 as long as the | leaf blade, sessile or with petiolules to 1.5 cm | long, the ultimate segments about 300-600 or more, 1-5 mm long, 0.5—1 mm wide; pedun- | cles 2.5—21 cm long; rays of the umbel nearly | obsolete or to 3.7 cm long; bractlets of the — involucel 1-3 mm long, linear, pubescent; pedicels 1-8 mm long; petals white; anthers | purple; styles about 1-2 mm long; fruit 6-10 | mm long, 3-6 mm wide, pubescent, the lat- eral ribs with wings 0.5-1 mm long, dorsal ribs filiform. Pinyon-juniper-mahogany com- | munities, at ca 2,380 m in western Millard County; Great Basin from southeastern Ore-_ January 1986 gon and southwestern Idaho to northern Cali- fornia, central Nevada and western Utah; | (i). Except for the white petals, purple anthers and sometimes slightly less pubescent foliage, plants of this taxon could pass for plants of L. foeniculaceum. Fruiting specimens may be difficult to distinguish. Lomatium scabrum (Coult. & Rose) Mathias Rough Lomatium; Cliff Lomatium. [Cynomarathrum scabrum Coult. & Rose]. Plants 6—25 (34) cm tall, acaulescent, mostly scabrous, from a branched caudex, clothed at the base by persistent leaf bases; leaves bipin- nately to tripinnately dissected, with (5) 7-11 opposite pairs of lateral primary leaflets, peti- oles 1-7 (10) cm long, blades (1.5) 2-11 cm long, lowest pair of primary leaflets less than 1/3 as long as the leaf blade and seldom over 1/4 as long, sessile or nearly so, the ultimate segments about 50—400 or more, 1-4 mm long, 0.4—2 mm wide; peduncles 5—25 (32) cm long; rays of the umbel 4-11, 0.5—2 (3) cm long; bractlets of the involucel 1-4 mm long, linear; pedicels 1-5 (10) mm long; petals and stamens mostly yellow or occasionally white when fresh, fading white in herbarium speci- mens; styles about 2-3 mm long; fruit 4-7 mm long, 3-4 mm wide, the lateral wings to 1 mm wide, the dorsal ribs filiform or some- times with a rudimentary wing at the base. There are 2 intergrading varieties as follows: 1. Leaves mostly bipinnately dissected, with about 50—110(140) ultimate segments; fruit 4-8 mm long; plants mostly found above 1,615m_L. scabrum var. scabrum Leaves tripinnately dissected, with about 150—400 or more ultimate segments; fruit 6-9 mm long; plants mostly found below 1,615 m po 0000d5b0000G00 L. scabrum var. tripinnatum Var. scabrum Desert shrub, pinyon-ju- niper, mountain brush, and white fir commu- _ nities, mostly on limestone and dolomite out- vada; 55 crops at 1,615 to 2,684 m in Beaver, I[ron, Juab, and Millard counties and adjacent Ne- (xii). Some specimens, especially from Iron County, are wholly transitional to the following variety. Var. tripinnatum Goodrich, var. nov. _ Similis Lomatio scabro var. scabro sed foliis tripinnatifidis et segmentis ultimo plus nu- ~ merosis differt. HOLOTYPE: Utah. Washington { Co., T41S, RI6W, Sec 10, SE1/4, 14.2 km 340: + degrees NW of St. George, Lava Ridge-Snow GOODRICH: UTAH FLORA, APIACEAE 99 Canyon, 1,280 m elev., Mahonia-Fraxinus- Coleogyne-Arctostaphylos comm., on sand- stone, 8 May 1984, S. Goodrich 20282 (BRY); isotypes ARIZ, RM, CAS, UC, POM, NY, UT, UTC, MO, US, WS. Additional speci- mens: Washington Co., T41S, R17 W, Sec 8, NE1/4, 22.5 km 309 degrees NW of St. George, 8 May 1984, S. Goodrich 20268 (ARIZ, BRY,CAS, MO,NY,POM,RM,UC,US, WS). Blackbrush and pinyon-juniper commu- nities, often on sandstone or in sandy places at 792 to 1,475 (2,170) m, in Washington County; and adjacent Arizona; 29 (ii). Lomatium triternatum (Pursh) Coult. & Rose Ternate Lomatium. [Seseli triternatum Pursh]. Plants 20—70 cm tall, acaulescent or subcaulescent, mostly hirtellous throughout except on the fruit, from a taproot with simple or sparingly branched crown, not clothed at the base with persistent leaf bases or only weakly so; leaves ternate-pinnately com- pound with (3) 9-21 leaflets or segments, petioles up to 23 cm long including the dilated sheathing base, or reduced to the sheath; blades 4—20 cm long, ovate in outline, the lowest primary leaflets often over 1/2 as long as the leaf blade, ultimate leaflets or segments 1-13 cm long, 1-15 mm wide; peduncles 15-55 cm long; rays of the umbel 4—20, 2-10 cm long; bractlets of the involucel about 6-10, 1-10 mm long, about 0.1-0.5 mm wide; pedicels 2-7 mm long; petals and stamens bright yellow when fresh but fading to white in herbarium specimens; styles about 1—-1.5 mm long; fruit 8—15 mm long, 4-11 mm wide, the lateral wings 1—2.5 (4) mm long, the dorsal ribs filiform. There are 2 subspecies. In Utah they can be separated as follows: ie Ultimate leaflets or segments linear, over 10 times as long as wide, to 13 cm long, 1—6 (10) mm wide; fruit broadly elliptic, the mature wings as broad or nearly as broad as the body L. triternatum ssp. platycarpum Ultimate leaflets elliptic, 3-9 times as long as wide, 2—6 cm long, (3) 6-15 mm wide; fruit rather narrowly elliptic to nearly linear, the mature wings seldon more than half as wide as the body. ..... L. triternatum ssp. triternatum Ssp. platycarpum (Torr.) Cronq. [L. sim- plex (Nutt.) Macbr.]. Sagebrush-grass, pinyon-juniper, mountain brush, ponderosa pine, lodgepole pine, and dry meadow com- munities at 1,310 to 2,895 m in Box Elder, 100 Cache, Daggett, Duchesne, Morgan, Rich, San Juan, Summit, Uintah, Utah, Wasatch, and Weber counties; southern British Colum- bia and Montana to Idaho and Colorado; 116 (viii). Ssp. triternatum Mountain brush and aspen communities, sometimes on heavy clay soils with Wyethia at 1,580 to 2,590 m in Weber and Summit counties; southern Alberta and British Columbia to Utah; 8 (0). Utah speci- mens are referable to var. anomalus (Jones) Cronq. [L. anomalum Jones]. Musineon Raf. Perennial plants with leaves mostly at or near the base, from a thickened taproot with a simple or branched crown or caudex; leaves 1 or more times pinnately or ternate-pinnately compound; umbel compound; involucre usu- ally lacking; involucel of several separate or basally united bractlets; calyx teeth well de- veloped, ovate; petals and stamens white or yellow; stylopodium lacking; carpophore en- tire to deeply cleft; fruit ovoid to linear oblong, somewhat laterally compressed, evi- dently ribbed. Musineon lineare (Rydb.) Mathias Rydberg Musineon. [Daucophyllum lineare Rydb. Aletes tenuifolia Coult. & Rose]. Plants 5.5— 25 cm tall, caulescent or subcaulescent, glabrous, from a mostly branched caudex, more or less clothed at the base with long-per- sisting leaf bases; leaves mostly on the lower 1/3 of the plants, ternate or more often pin- nate, with 2—4 opposite pairs of lateral leaflets, petioles 0.5—6 (14) cm long, blades 1—5.3 cm long; leaflets, sessile, entire or bifid, trifid or rarely pinnatifid, ultimate leaflets or lobes 3-20 mm long; peduncles 5—22 cm long, very slender; umbel solitary; rays about 5-10, 1-5 mm long; involucels of about 3 linear or narrowly elliptic bractlets 4—10 mm long; pedicels about 1 mm long; calyx teeth about 0.5 mm long, greenish or purplish with scarious margins; petals and stamens white; styles about 1 mm long; fruit 2-4 mm long, minutely scabrous, the ribs evident but not winged. Limestone cliffs in the Bear River Range, Cache County; endemic; 4 (0). Oreoxis Raf. Caespitose, acaulescent herbs from branched woody caudices, these usually GREAT BASIN NATURALIST Vol. 46, No. 1 clothed with long-persisting leaf bases; leaves pinnate or bipinnate; umbels compound; in- volucre mostly lacking; bractlets of the in- volucel more or less united at the base, usu- ally exceeding the flowers; calyx teeth conspicuous; petals and stamens yellow at least when fresh; stylopodium lacking; car- pophore lacking; fruit oblong to ovoid-oblong, slightly compressed laterally, the ribs corky- winged. Plants of the genus could reasonably be included in Cymopterus, and with the re- cent discovery of the low elevation O. trotteri such inclusion will probably be necessary. il, Bractlets obovate, toothed at the apex, usually purplish, plants of the La Sal Mountains simnth dani hat eee O. bakeri = Bractlets linear or narrowly elliptic, entire, acute to acuminate; plants more widely dis- tributed Plants pulvinate caespitose, forming clumps to 30 cm wide, from low elevations in Grand County; caudex clothed with a thatch of terete leaf bases; ultimate segments elliptic to cuneate-ovate O. trotteri — Plants caespitose but hardly pulvinate, from high elevations, widespread; caudex clothed with short, more or less flattened leaf bases; ultimate segments of leaves linear to linear- elliptic O. alpina Oreoxis alpina (Gray) Coult. & Rose Alpine Oreoxis [Cymopterus alpinus Gray]. Plants 2.5-11.5 cm _ tall, — scabrous-hirtellous throughout, from a branched caudex, the caudex clothed with persisting leaf bases; leaves all basal, mostly bipinnate, with ca 4 opposite pairs of sessile or nearly sessile lat- eral primary leaflets, the upper pairs and those of smaller leaves sometimes once pin- nate and then trifid to pinnatifid, petioles 0.5-2.5 cm long, blades 1—3.5 cm long oblong in outline, lowest pair of primary leaflets 4-14 mm long, the ultimate segments about 1-6 mm long, 0.4-1.5 mm wide, linear to nar- rowly elliptic; peduncles 2-10.5 cm long; um- bel solitary; involucre lacking; rays 4-7, 1-6 mm long; involucels of 5—9 bractlets 1-4 mm long, united at the base; pedicels obsolete or to about 0.3 mm long; calyx teeth 0.6—1 mm long, green; petals and stamens yellow when fresh, fading to white or cream or purple tinged within a few years in herbarium speci- mens; styles 1.7—2 (3) mm long; fruit 4—5 mm long, the ribs with low corky wings to about 0.7 mm wide. Forb-grass, limber pine, January 1986 spruce, and alpine communities, and raw es- carpments and barren ridges at 2,440 to 3,475 m in Duchesne, Garfield, Grand, San Juan, San- pete, Summit, and Wayne counties; Wyoming to New Mexico and Arizona; 27 (vi). Oreoxis bakeri Coult. & Rose Plants 1-12 cm tall, slightly puberulent at base of umbels and rays; leaves basal, bipinnate for the most part or pinnate with pinnatifid or trifid leaflets, with 3-4 opposite pairs of lateral primary leaflets, the petioles 0.8—2.5 cm long; blades 0.8—5 cm long, lowest pair of primary leaflets to about 1 cm long, sessile or nearly so, the ultimate segments to 7 mm long, to 1 mm wide; peduncles 1-11 cm long; umbels solitary, involucre lacking; rays 3-8, 3-5 mm long; bractlets of the involucel united at base, 3-5 mm long, nearly linear-ellip- tic to obovate, usually 3—-toothed at the apex; petals and stamens yellow at least when fresh; styles to about 1 mm long; fruit 2-4 mm long, the ribs with low corky wings to 0.75 mm wide. Alpine forb-grass communities, at ca 3,660 m, La Sal Mountains, in Grand and San Juan coun- ties; Colorado, Utah, and New Mexico; 4 (0). Oreoxis trotteri Welsh & Goodrich Plants pul- vinate-caespitose, forming clumps to 30 cm wide, 4—8 cm tall, scabrous and more or less glandular, from a branching caudex, this clothed with a thatch of persistent, terete leaf bases and peduncles; leaves all basal, bipinnate, with ca 4 opposite pairs of sessile, lateral, primary leaflets, the upper pairs and those of the smaller leaves sometimes once-pinnate and then trifid or pin- natifid; petioles 1-3.5 cm long; blades 1.5-2.3 cm long, oblong in outline, the lowest pair of primary leaflets 3.5-5 mm long, the ultimate segments 1-3.5 mm long, 1-3 mm wide, elliptic to cuneate-ovate; peduncles 4—7.5 cm long; um- _ bel solitary; involucre lacking; rays 5-7, 3-5 mm _ long; involucels of 4—7 linear-subulate bractlets 2-3.5 mm long, distinct or essentially so; pedicels obsolete or to ca 1 mm long; calyx teeth ca 1 mm long, green or purplish; petals and stamens yellow; styles 1-1.2 mm long; fruit 2.8-4.8 (5) mm long, the ribs with low, corky wings to 0.7 mm wide. Mixed juniper and warm _ desert shrub community at ca 1,464 m in Grand County; endemic; 2 (0). Orogenia Wats. Perennial acaulescent glabrous low herbs from a fusiform or globose root; leaves ternate or GOODRICH: UTAH FLORA, APIACEAE 101 biternate with linear entire leaflets; umbel compound; involucre lacking or of a few linear minute scarious bractlets; calyx teeth obso- lete; petals and stamens white or purplish; stylopodium lacking; carpophore lacking; fruit oblong to oval, nearly round in cross section, the dorsal ribs evident or obsolete, the lateral ones corky-winged but inflexed into the com- missure, a corky riblike projection also run- ning the length of the commissural faces of each mericarp. Orogenia linearifolia Wats. Indian Potato. Plants 5-10 (13) cm tall, glabrous, not aro- matic, from a globose or fusiform root, with a fragile etiolated subterranean pseudoscape- like stem easily detached from the tuberous root; leaves borne at ground level or a few arising from the tuberous root with etiolated petioles, ternate or biternate, blades 3-8 (12.5) cm long, the 3-9 leaflets 1.5—5 (11.5) cm long, 1-11 mm wide, linear, entire, the lowest pair of petiolules to 2 cm long; pedun- cles 3-8 cm long, usually a little longer than the subterranean stem; involucre lacking; rays 3-12, but rarely more than 5 of them fertile, 0.3-3 cm long; involucel proper apparently lacking, but some of the pedicels usually bear- ing a linear bractlet to 4 mm long; pedicels nearly obsolete or to 2 mm long; petals white; filaments white, anthers pale or dark purple; styles about 1 mm long; fruit about 4-6 mm long; dorsal ribs filiform. Sagebrush-grass, oak, maple, aspen, ponderosa pine, white fir, and rarely desert shrub communities, mostly flowering at the edge of melting snow at 1,370 to 2,805 m in Beaver, Box Elder, Juab, Mil- lard, Morgan, Salt Lake, San Juan, Sanpete, Sevier, Summit, Tooele, Uintah, Utah, Wasatch, Washington, and Weber counties; Washington to Montana, south to Utah and Colorado; 50 (vii). Osmorhiza Raf. Perennial caulescent usually pubescent herbs from taproots with simple or branched crowns; leaves ternately or pinnately 1-3 times compound with well-marked leaflets; umbels compound; involucre lacking or of 1 or a few narrow foliaceous bracts; involucel lack- ing or of several foliaceous reflexed bractlets; calyx teeth obsolete; petals and stamens white, greenish white, yellow, pink, or pur- ple; stylopodium, conic to depressed; car- 102 pophore bifid less than 1/2 its length; fruit linear or clavate, somewhat compressed later- ally, bristly hispid to glabrous, the ribs nar- row. I. Ovaries and fruit glabrous, generally obtuse at both ends; petals and stamens yellow or greenish yellow; leaves (1)2 times pinnately or ternate-pinnately compound; plants strongly aromatic, usually with more than 2 stems O. occidentalis — Ovaries and fruit bristly hispid, with long, pointed bristly hispid tails; petals white or greenish white; leaves biternate; plants not strongly aromatic, often with solitary stems .. 2 Mature fruit including tails mostly 16-25 mm long, the apex concavely pointed into a 1-2 mm long beak; the most divergent rays spreading 30 degrees to 65 degrees from the peduncle; fruiting pedicels mostly ascending- spreading; plants most common below 2,470 HGIENY, a ei lala Meo tte eae cra See arnold O. chilensis — Mature fruit including tails mostly 13-18 mm long, the apex convex and obtuse; the most divergent rays spreading 40 degrees to 90 degrees from the peduncle; fruiting pedicels horizontally spreading to ascending; plants common above as well as below 2,470 m O. depauperata Osmorhiza chilensis H. & A. [O. nuda Torr.|. Stems often solitary, 18—75 cm tall, from a taproot, without long-persisting leaf bases; herbage not strongly aromatic; leaves basal and 2—3 cauline, biternate, usually with 9 distinct leaflets, petioles about 3-16 cm long, or cauline leaves sessile, blades 5—15 cm long, lateral primary leaflets about as long as the central one, with petiolules (1) 2-5.5 cm long, blades of leaflets 1—4 (5.5) cm long, ellip- tic to ovate, lobed to cleft, and toothed, ciliate and often pubescent on nerves below and sometimes scattered pubescent between the nerves; peduncles 5-34 cm long; umbels 1-5; involucre lacking; rays 3—7, 2.5-9 (13) cm long, ascending or spreading-ascending, glabrous to hirtellous; involucels lacking; pedicels 5-22 (30) mm long, ascending; petals and stamens greenish white; styles less than 0.5 mm long; fruit, including the tails, 16-25 mm long, linear-clavate, bristly-hispid, the beak concavely pointed, 1-2 mm long, the concave beak usually evident in young fruits. Oak, maple, aspen, Douglas-fir, white fir, narrowleaf cottonwood, and riparian commu- nities at 1,520 to 2,470 (2,680) m in Box Elder, Cache, Daggett, Davis, Duchesne, Juab, Mil- GREAT BASIN NATURALIST Vol. 46, No. 1 lard, Salt Lake, Sanpete, Tooele, Uintah, Utah, Wasatch, Washington, and Weber counties; Alaska to California, east to Alberta and Arizona, also Great Lakes region and in Argentina and Chile; 54 (viii). See O. depau- perata. Osmorhiza depauperata Phil. Blunt-fruit Sweet-cicely. [O. obtusa (Coult. & Rose) Fern.]. Stems mostly solitary, 14-63 (77) cm tall, often with a slight ring of hairs at the nodes, from a taproot, without persisting leaf bases; herbage not strongly aromatic; leaves basal and 1—3 cauline, biternate, usually with 9 distinct leaflets, or the upper cauline ones once-ternate, petioles (1) 3-17 cm long, often with dilated, ciliate bases, blades (2) 4-11 cm long, the lateral primary leaflets about equal to the central one or a little shorter, with petiolules (0.5) 1—4 cm long, blades of leaflets 1-4 (5.5) cm long, elliptic to ovate, lobed to cleft and toothed, ciliate and often pubescent on nerves below and sometimes scattered pubescent between nerves; peduncles 3.5—15 (22.5) cm long; umbels 3—6; involucre lacking, or rarely of a solitary bract to 12 mm long; rays 3-5, 1.5-8.5 cm long, spreading to divari- cate, involucels lacking or infrequently of 1 or 2 separate ciliolate bractlets to 3 mm long; pedicels 5-20 mm long, spreading to divari- cate; petals greenish white; styles about 0.2 mm long; fruit, including the tails (11), 13-18 mm long, linear-clavate, the beak convex-ob- tuse. Oak, maple, aspen, ponderosa pine, Douglas-fir, lodgepole pine, spruce-fir, ripar- ian, and rarely pinyon-juniper and sagebrush communities at 1,980 to 3,200 m in all Utah counties except Cache, Davis, Emery, Mor- gan, Piute, Rich, and Wayne; Alaska to Cali- fornia, east to South Dakota and New Mexico, also in the Great Lakes Region, also Chile and Argentina; 110 (xv). Very much like O. chilen- sis but generally at higher elevations and with more widely spreading rays and pedicels and shorter fruits, and much more common and widespread in Utah than is O. chilensis. Al- though the two taxa are similar and have about the same distribution in North and South America, the differences seem quite constant. Osmorhiza occidentalis (Nutt.) Torr. West- ern Sweet-cicely. [Glycosma occidentalis Nutt. ex T. & G.]. Plants 6-13 dm tall, from a taproot, with few or no persisting leaf-bases; _ strongly aromatic, leaves (1)2 times pinnate or _ January 1986 the upper cauline ones ternate pinnately com- pound, with 3-4 pairs of opposite lateral pri- mary leaflets, petioles of lower leaves 4—30 cm long or longer, the upper ones reduced, | lower blades to 25 cm long or longer, the / upper ones much reduced, the lowest pair of | primary leaflets usually again pinnate, usually ‘over 1/2 as long as the leaf blade, with peti- ‘olules 1-3.5 cm long, ultimate leaflets 1-9 cm | long 0.5—4 cm wide, lanceolate to lance-ellip- | tic or ovate coarsely toothed and some often | lobed; peduncles 6—20 cm long; umbels about 3-5; involucre lacking or occasionally of 1—2 linear or filiform bracts to 16 mm long; rays GOODRICH: UTAH FLORA, APIACEAE 103 lacking; rays 5-14, 1-5 (7) cm long, ascend- ing; involucels lacking; pedicels 3-10 mm long; petals and stamens white; styles mostly less than 1 mm long; fruit 3-5 mm long. Streambanks, on Abajo and La Sal mountains in San Juan County; Wyoming south to New Mexico; 4 (0). Pastinaca L. Biennial or perennial caulescent herbs from large taproots; leaves pinnately compound, with broad-toothed to pinnatifid leaflets; um- bels compound; involucre and involucel usu- ally lacking; calyx teeth obsolete; petals yel- 7-13, 2-6.5 cm long, involucels lacking; \ pedicels 2-7 mm long, calyx obsolete; petals / greenish white or greenish yellow, about 1-2 low or red; stylopodium depressed-conic; carpophore divided to the base; fruit elliptic mm long; stylopodium low; styles about ‘0.75-1 mm long; carpophore divided to the | base; fruit 16-20 mm long, 2-3 mm wide, linear, glabrous. Tall forb, aspen, oak-maple, ‘spruce-fir, riparian, and infrequently in sage- | brush communities at 1,765 to 3,170 m in Box ‘Elder, Cache, Carbon, Duchesne, Iron, Juab, Millard, Morgan, Salt Lake, San Juan, Sanpete, Sevier, Summit, Tooele, Utah, Wasatch, Washington, and Weber counties; southern British Columbia and Alberta south to California and Colorado; 67 (xvi). Oxypolis Raf. Perennial, caulescent, glabrous herbs from fascicled tuberous roots; leaves pinnate; um- bels compound; involucre and involucel lack- ing; rays ascending; calyx teeth conspicuous; petals white to purple; stylopodium conic; carpophore divided to the base; fruit oblong to oval, strongly flattened dorsally, dorsal ribs filiform, lateral ribs broadly winged. Oxypolis fendleri (Gray) Heller [Arche- mora fendleri Gray]. Plants 6—8 dm tall, with- out persisting leaf-bases; leaves pinnate with _ 2-5 pairs of opposite lateral leaflets, the up- per ones sometimes reduced to bladeless or ‘nearly bladeless sheaths, the petioles (3) 5-15 cm long or the upper blades sessile on a di- lated sheath, blades 7-17 cm long, oblong in outline, leaflets sessile, 2-5 cm long, ovate to orbicular, shallowly to deeply crenate-den- ‘ tate or serrate or rarely incised, or those of the to obovate, strongly flattened dorsally, the dorsal ribs filiform, the lateral ones narrowly winged. Pastinaca sativa L. Parsnip. Biennial caulescent aromatic herbs 8—15 dm tall, from a taproot; leaves pinnate or partly bipinnate in some of the lower leaflets, with 3-6 opposite or offset pairs of lateral leaflets; petioles 3-15 mm long or lacking and the blade sessile on a dilated sheath; blades 12-35 cm long or longer, oblong in outline; leaflets sessile and sometimes confluent or the lower ones some- times on petiolules to 1.7 cm long, the blades 2.5-12 cm long, lanceolate to ovate, coarsely serrate, and often lobed; umbels 6—15 or more, the terminal one sessile or pedunculate but shorter than the 2 immediately lateral ones, the lateral umbels alternate or opposite or on opposite branches supporting 2 or more umbels; involucre lacking or of 1—few linear entire or occasionally toothed or lobed bracts to 2 (4) cm long; rays 9-25, 0.8-8.5 cm long; involucels lacking or infrequently of 1—2 lin- ear bractlets to 2 mm long; pedicels 4-20 mm long; petals greenish yellow or reddish; styles less than 0.5 to about 1 mm long; fruit 5—8 mm long, 3-6 mm wide, broadly elliptic to orbicu- lar or obovate, strongly flattened dorsally the dorsal ribs filiform, lateral ribs slightly winged. Ditch banks, roadsides, fence lines, gardens, fields, margins of ponds and lakes, and moist floodplains at 1,370 to 2,365 m, probably cultivated in all counties of the state, escaping and persisting, introduced from Eu- rope, now widely established in North Amer- ica; 23 (x). The cultivated plants (ssp. sativa) differ from the wild plants [ssp. sylvestris iupper leaves lanceolate to linear and some- | times entire; peduncles (1) 4—20 cm long; um- ‘bels usually 4 or more per stem; involucre 104 (Mill.) Roua & Camus] in having larger roots. Some of the wild plants might be recent es- capes from cultivation. Perideridia Reichenb. Perennial, caulescent herbs from a fusiform or tuberous root or fascicle of tuberous roots, these often deep-seated and easily detached from the rather fragile etiolated subterranean portion of the stem and often lacking in herbarium specimens; leaves ternate, pin- nate, or ternate-pinnately compound, the up- per ones sometimes reduced to a simple, lin- ear rachis; petioles sheathing; umbels com- pound; involucre lacking or of mostly few more or less scarious bracts; involucel lacking or of 1—few bractlets; calyx teeth inconspicu- ous, of the texture and color of the petals (in ours); petals white when fresh; stamen white; stylopodium conic or low conic; carpophore divided to the base; fruit linear-oblong to or- bicular, scarcely compressed or lightly so at right angles to the commissure, with filiform ribs. il. Bractlets of the involucel scarious, as wide or to 5 times as wide as the pedicels, 3-5 mm long, linear to ovate, often caudate; styles 1-2 mm long after anthesis; longest rays rarely over 2 cm long; lower leaves mostly ternate- pinnately 2 or more times compound with petiolulate primary leaflets, the ultimate divi- sions commonly 10—50 or more per leaf .... P. bolanderi Bractlets of the involucel not scarious or with narrow scarious margins, only about as wide as the pedicels, to 3 mm long, linear or linear subulate; styles to about 1 mm long after an- thesis; longest rays commonly 2—3(4) cm long; leaves ternate or pinnate; leaflets or pinnatifid divisions sessile, simple, commonly 3-5, rarely over 10 per leaf P. gairdneri Perideridia bolanderi (Gray) Nels. & Macbr. Yampah. [Podosciadium bolanderi Gray]. Plants 23—40 cm tall, glabrous, with- out long-persisting leaf bases;. leaves often crowded on the lower part of the stem, ter- nate-pinnately 2 or more times compound, with petiolulate primary leaflets, the upper ones reduced, and sometimes simple and lin- ear, often withered before or shortly after an- thesis, petioles to 4 cm long or lacking and the petiolules arising directly from a dilated sheath, blades 4-12 cm long, the ultimate leaflets strongly dimorphic, 0.2—8 cm long, GREAT BASIN NATURALIST Vol. 46, No. 1 mostly 10-50 or more per leaf on the lower leaves; peduncles (2) 5-14 cm long; umbels 2—6 per stem; involucre lacking or usually of 1—4 scarious bracts to 5 mm long; rays 4—12, 1—2 cm long; bractlets of the involucels about 4—8, 3-5 mm long, to 2.5 mm wide, linear to ovate and often caudate, with pale yellow- green midrib, this often flanked on either side by purple and then by conspicuous scarious margins; pedicels 3-6 mm long; petals white; styles 1-2 mm long, spreading to recurved; fruit 3—4 (5) mm long, some of the ribs usually conspicuously ridged. Sagebrush, juniper, mountain brush, and stream-side communi- ties, sometimes in snowflush areas at 1,524 to 2,320 m in the western 1/2 of Box Elder County and Deep Creek Mountains., Juab County; eastern Oregon and western Idaho south to California and Utah; 12 (iii). Some- times confused with P. gairdneri, but in addi- tion to the several features listed in the key it differs as follows (features of P. gairdneri in parentheses): peduncles mostly 5-14 cm long (mostly 1-5 cm long); fruit 3-5 mm long, oblong, the ribs ridged (2-3 mm long, orbicu- lar, the ribs obscure). Perideridia gairdneri (H. & A.) Mathias False Yarrow [Atenia gairdneri H. & A.; Carum garrettii A. Nels. in Coult. & Rose, type from the Wasatch Mountains]. Plants 15-75 cm tall, glabrous, without long-persist- ing leaf bases; leaves 2—5 per stem, ternate or pinnate, with up to about 5, rarely more, ses- sile leaflets, the upper ones reduced and often simple and linear; leaflets to 13 cm long, mostly confluent with the rachis, linear and hardly wider than the petiole, occasionally expanded to 11 mm wide; peduncles (1) 2-5 (7) em long; umbels 2—5 per stem; involucre lacking or occasionally of 1 or 2 linear bracts to 6 mm long; rays 7-16, 0.7-4 cm long; | bractlets of the involucels lacking or more | often 1-6, 1-3 mm long, linear or linear-sub- | ulate, hardly if at all wider and conspicuously shorter than the pedicels, not marked with purple or if so then the whole bractlet mostly | purple; pedicels 3-5 mm long; petals white or turning purplish; styles to 1 mm long, re-_ curved; fruit 2-3 mm long, orbicular, the ribs _ obscure. Sagebrush, forb-grass-silver sage- brush, meadow, oak, maple, aspen, and wil- . low-streamside communities at 1,680 to 2,685 - m in eastern Box Elder, Cache, Daggett, — January 1986 Juab, Salt Lake, Sanpete, Summit, Utah, and Wasatch counties; British Columbia to southern California, east to Saskatchewan and New Mex- ico; 25 (v). Our plants belong to ssp. borealis Chuang & Const. Podistera Wats. Perennial acaulescent glabrous plants from taproots or branched caudices; leaves pinnate with deeply lobed leaflets; umbel solitary, com- pound, compact; involucre wanting; involucel of toothed bractlets; calyx teeth conspicuous, ovate; petals greenish yellow; stylopodium conic; carpophore stout, undivided; fruit oval, slightly flattened laterally, the ribs filiform to prominent. Podistera eastwoodiae (Coult. & Rose) Mathias & Const. [Ligusticum eastwoodae Coult. & Rose]. Plants 7-20 (30) cm tall, acaules- cent, without or with few long-persisting leaf bases; leaves pinnate, with 4-6 pairs of sessile lateral leaflets, petioles 1.5—7 cm long; blades 2.5-7.5 cm long, oblong in outline; leaflets about 1-2 cm long, ovate to obovate in outline, ternately or palmately lobed or cleft, the larger lobes again toothed or lobed; peduncles (7) 10-20 (30) cm tall; involucre lacking; rays 5-8, 2-8 mm long; bractlets of the involucel 4—6 mm long, often exceeding the flowers and fruit, ovate ~ or obovate, with 2—3 teeth or lobes, with the texture and color of the leaves; pedicels 1-2 mm long; petals greenish yellow, turning purple; styles about 1 mm long; fruit about 3-4 mm long, ~ the ribs evident but not winged. Apparently rare at upper elevations of the La Sal Mountains, San Juan County; Colorado, New Mexico, and Utah; 0 (0). Sium L. Perennial, caulescent herbs from fascicles of fibrous roots; leaves mostly pinnately compound or decompound, with well-marked, toothed to _ pinnatifid leaflets; umbels compound; involucre _ of entire or incised, often reflexed bracts; in- volucel of narrow bractlets; calyx teeth minute or obsolete; petals white, stylopodium depressed _ or rarely conic; carpophore divided to the base (but threadlike and adnate to the faces of the - mnericarps in our plants); fruit elliptic to orbicu- | lar, slightly compressed laterally and somewhat ‘constricted at the commissure, the subequal ribs prominent and corky but hardly winged. GOODRICH: UTAH FLORA, APIACEAE 105 Sium suave Walt. Hemlock Water-parsnip. Plants 5-10 dm tall; leaves pinnate or occa- sionally partly bipinnate, with 4—6 opposite pairs of sessile lateral leaflets, lower petioles to 25 cm long, often septate, the upper ones smaller and sometimes reduced to a dilated sheath, lower blades 14-32 cm long, the up- per ones reduced; leaflets 2-8(15) cm long, (1) 3-8 (20) mm wide, linear to lanceolate, sharply and uniformly serrate to pinnatifid with linear segments; peduncles 4-10 cm long; umbels 3-11 or more per stem; involu- cre of about 1-6 separate, often reflexed bracts 2-9 mm long; rays 11-24, 1.5-3 em long; involucels of (2) 5-12 separate bractlets 2—5 mm long; pedicels 2-8 mm long; petals and stamens white; styles about 1 mm long; fruit 2-3 mm long, the ribs prominent. Mud flats, marshlands, wet meadows, along streams and shorelines, and in ponds and lakes at 1,365 to 2,990 m in Garfield, Piute, Rich, Salt Lake, Sanpete, Sevier, Utah, and Wayne counties; southern British Columbia to Newfoundland, south to California and Vir- ginia; 15 (i). Often confused with Cicuta and frequently found with that genus in herbaria, but conspicuously different by the merely pinnate leaves. Torilis Adans. Annual caulescent hispid or pubescent herbs from slender taproots; leaves 1—2 times pinnate or pinnately decompound, petioles sheathing; umbels compound, capitate or open, sessile or pedunculate; involucre lack- ing or of afew small bracts; involucel of several linear or filiform bractlets; calyx teeth evident to obsolete; petals white; stylopodium thick, conic; carpophore bifid or cleft ca 1/3—1/2 its length; fruit ovoid or oblong, flattened later- ally, tuberculate or prickly, the primary ribs filiform, setulose, the lateral ribs displaced onto the commissural surface, the intervals covered with glochidiate prickles or tuber- cles. Torilis arvensis (Huds.) Link Hedge Pars- ley. [Caucalis arvensis Huds.]. Plants 3-10 dm tall, divaricately branched, appressed- hispid throughout, retrorsely so on the stems and antrorsely so on the leaves and rays; leaves 2—3 times pinnate, or the upper ones once-pinnate, the ultimate leaflets 5-60 mm long, 2-20 mm wide, ovate to linear lance- 106 olate, acute or acuminate, regularly incised or divided: peduncles 2-12 cm long; involucre lacking or of a single small bract; rays 2—10, 0.5—2.5 cm long; involucel of several subulate bractlets longer than the pedicels; pedicels 1—4 mm long; petals white; styles short; fruit ovoid-oblong, 3-5 mm long, the mericarps densely covered with straight glochidiate prickles with minute retrorse-barbs, these spreading almost at right angles and about as long as the fruit is wide. LaVerkin in orchard, (Barnum 1316 BRY); Washington County; ad- ventive; introduced from southern and cen- tral Europe; 1(0). Yabea K.-Pol. Annual caulescent herbs from taproots; leaves pinnate or dissected; umbels com- pound; involucre of a few entire or dissected, usually somewhat scarious bractlets; calyx teeth evident; petals white; stylopodium thick and conic; carpophore entire or bifid at the apex; fruit oblong or ovoid, somewhat com- pressed laterally, with spreading uncinate prickles along alternating ribs, and bristly- hairy on the other ribs. Yabea microcarpa (H. & A.) K.-Pol. Califor- nia Hedge-parsley. (Caulis microcarpa H. & A.). Plants annual, caulescent, 8—40 cm tall, pubescent with spreading hispid hairs, froma slender taproot; leaves 2—3(4) times pinnate or ternate-pinnate, with about 3—4 opposite pairs of lateral primary leaflets, blades 1-5 cm long, oblong or ovate in outline, on petioles 1—4.5 cm long or the upper ones sessile, low- est pair of primary leaflets about 1/2 as long as the leaf blade, sessile or petiolulate, ultimate segments 1-8 mm long, 0.5—2 mm wide; pe- duncles 3-10 cm long; umbels 1—4, involucre resembling the upper leaves or a little smaller; rays (1) 2-7 (9), 1.5—10 cm long, often about as long as the peduncles; involucels sim- ilar to the involucre, but usually reduced, GREAT BASIN NATURALIST Vol. 46, No. 1 sometimes much reduced and the bractlets only pinnatifid or entire; pedicels 5-15 cm long; petals white; stamens white; styles very short; carpophore bifid for about 1/5 its length; fruit 3-7 mm long. The one specimen seen (Atwood 4871 BRY) is from the Pine Valley Mountains, Washington County; British Columbia south to Baja California, east to Idaho and Arizona; 1(0). Zizia Koch Perennial glabrous or subglabrous herbs with basal and cauline leaves, from a short caudex and a cluster of fleshy-fibrous roots; leaves simple or ternate, with toothed blades or leaflets; umbels compound; involucre lack- ing or obsolete; involucel of a few inconspicu- ous bractlets; calyx teeth well developed; petals bright yellow; stylopodium lacking; car- pophore bifid about 1/2 its length; fruit oblong or broadly elliptic, somewhat laterally com- pressed, the ribs prominent but not winged. Zizia aptera (Gray) Fern. [Thaspium trifo- liatum var. apterum Gray]. Perennial, caules- cent, glabrous herbs 15-50 cm tall, from a taproot or fascicle of roots, without long-per- sisting leaf bases; basal leaves simple, rarely ternate, petioles 3-18 cm long, blades 1.5—5 cm long, ovate to nearly orbicular, cordate, crenate-serrate; cauline leaves ternate, not over 3 cm long, the leaflets sessile or on peti- olules to 4mm long; peduncles 6—12 cm long; umbels 1 or 2 per stem; involucre lacking or obsolete; rays 10-17, 0.5—-2 cm long; in- volucels of about 4—6 bractlets, to about 2 mm long, separate or united at the base; pedicels 1-3 mm long; petals yellow; stamens yellow; styles about 1 mm long; fruit about 2 mm long, the ribs prominent. Willow-streamside and meadow communities at 2,130 to 2,440 m in Sanpete, Sevier, Summit, Utah, and Wasatch counties; widespread in the United States and Canada; 12 (iii). ( QUERCUS (FAGACEAE) IN THE UTAH FLORA Stanley L. Welsh! ABSTRACT.—Reviewed are the oak taxa as they are presently understood in Utah. Keys and descriptions are | included, occurrences are cited, and problems of hybridization are discussed. Named as new varieties from Utah are Quercus gambelii Nutt. var. bonina Welsh and Quercus havardii Rydb. var. tuckeri Welsh. Both varieties occur in southeastern Utah. Quercus eastwoodiae Rydb. is proposed as a hybrid. The native oaks have been a source of con- fusion almost from the beginning of botanical exploration, and a huge bibliography has ac- cumulated dealing with the oaks of Utah and ‘the West (Harper et al. 1985). Collection of the materials serving in typification of the ear- liest known portion of the complex of species existing in Utah confounded interpretation from the beginning. The first epithet in our oaks and a name that has long plagued Utah plant taxonomy, Quercus undulata Torr., was published in 1828 and is based on material taken by Dr. James on the Long Expedition in the summer of 1820 (Tucker 1971). The exact nature of the type material has remained ob- .scure for reasons reviewed by Tucker (1971), but the material was evidently taken in Hard- ing County, New Mexico, a place where Q. gambelii Nutt. and Q. grisea Liebm. coexist. Although indicating that “the type shows scant evidence of the influence of Q. gambelii (aside from characters of foliar trichomes), Tucker (1971) concludes that the locality from which the type was taken contains a mixture of intermediates between the parental taxa and that the “‘species —Quercus undulata— is in fact a variable complex derived from hybridi- -zation.” The observations made by Tucker suggest, however, quite a different applica- ‘tion of the name, i.e., that it should replace Q. grisea, a later synonym. Interpretation of the type specimen, not the population that it came from, is crucial in typification. The cloud still remains, but fortunately it is beyond the bounds of Utah. The Utah oaks belong to three main popula- tion complexes: Q. gambelii Nutt., Q. turbin- ella Greene, and Q. havardii Rydb. Within those complexes the species concepts are mostly clear and unarguable, but they have no really apparent barriers to hybridization, and interme- diates are known between nearly all of them. The following taxonomic treatment is based on the examination of more than 300 specimens in Utah herbaria and more than three decades of experience with oaks in the field. Numbers following the descriptions of the taxa and hybrids indicate the number examined (in Arabic numerals) and the number collected by me (in Roman numerals). Quercus L. Trees or shrubs; wood hard, ring-porous, with prominent rays; leaves alternate, lobed, toothed, or entire; staminate flowers in usually pendulous, naked catkins; bracts caducous; calyx with 2-8 lobes; stamens 3-12; pistillate flowers with a subtrilocular, 6-ovuled ovary; stigma 3- lobed, enclosed by a scaly involucre, this hard- ened and cuplike, surrounding the base of the nut or acorn. I, Leaves evergreen, the lobes of teeth spines- cent, or seldom entire; plants of Washington, and, less commonly, of Kane and San Juan counties, hybridizing with the following Q. turbinella Leaves deciduous (persistent in some hy- brids), the lobes variously angled or rounded, but seldom, if ever, spinescent; plants of broad or other distribution................--.06-+- 2 Leaf lobes typically 1-2 times longer than the width of the leaf axis, rounded to obtuse or less commonly acute and often bilobed apically; plants broadly distributed Q. gambelii ‘Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 107 108 — Leaf lobes seldom, if ever, as long as the width of the leaf axis, acute to acuminate apically .....3 4(3). Plants deciduous, colonial in sandy sites, typi- cally with branch ends protruding above the substrate 1-5 dm; acorns mostly over 15 mm long and about as broad; hybridizing with the former along canyons.............. Q. havardii — Plants semievergreen, mainly 10—30 dm tall or more, or, if deciduous, the leaves typically hairy above and densely so beneath, forming clones within and adjacent to stands of Q. gam- belii, and occurring sporadically where Q. gambelii and Q. turbinella or Q. havardii coex- ist; acorns typically less than 15 mm long and less than 10 mm wide, if formed atall ....... Bae Bie Q. pauciloba and Q. eastwoodiae (hybrids) Quercus gambelii Nutt. [Q. stellata var. utahensis A. DC., type from west of Salt Lake City (?); Q. utahensis (A. DC.) Rydb.]. Clonal, deciduous shrubs or small trees, or less commonly trees to 10 m tall and with a trunk diameter to 6 dm or more thick, spread- ing by rhizomes; leaves densely grayish or yellowish stellate hairy on both surfaces when young, in age stellate hairy and paler beneath but glabrate and green and subglossy above, 2.4-17 cm long, 1-11 cm wide, obovate to elliptic in outline, the sinuses obliquely de- scending about 1/4—3/4 to the midrib, the lat- eral lobes (0) 2-10, oblong to lance-oblong, entire or notched apically and sometimes again laterally; staminate catkins 3.5-5 cm long; involucral cup 3-10 mm long, 10-17 mm wide, short-pedunculate to subsessile, ca 1/4—1/2 the length of the acorn, clothed with imbricate, densely hairy scales; acorns 8—18 mm long, 7-15 mm thick. Mountain brush, sagebrush, pinyon-juniper, ponderosa pine, and aspen communities at 1,125 to 2,745 m in all Utah counties except Daggett and Rich (?); Wyoming, Colorado, New Mexico, Arizona, Nevada, Texas, and Mexico; 206 (xxi). Gambel oak is central to a series of prob- lematical taxa, belonging in a broad sense to the Q. undulata Torr. complex, in which ev- ery degree of consanguinity is recognized. All our indigenous oaks are portions of the com- plex, and all form intermediates wherever contact is or has been made. Viewed broadly, all could be regarded as phases of Q. undu- lata, sensu latissimo. Problems related to such an approach involve similar circum- stances of hybridization with species belong- ing to other oak groups away from the Q. GREAT BASIN NATURALIST Vol. 46, No. 1 undulatum centrum. Ultimate consolidation of all intergrading groups would lead to absur- dity. There is one variant within Q. gambelii, however, that is so different as to require taxonomic recognition, as follows: Var. bon- ina Welsh var. nov. Persimilis Querco gambe- lii var. gambelii in foliis et habitu sed in glandibus (27-33 mm longis nec 8-18) et cupulis (20-25 mm latis nec 10-17) majoribus differt. TypE: USA: Utah. San Juan County, T36S, RI3E, $28, GCNRA, Lake Powell, Goodhope Bay springs, Cottonwood, oak, willow community, sandy alluvium over Chinle Formation, 21 Sept. 1983, S. L. Welsh & E. Neese 22575 (Holotype BRY; 6 isotypes distributed previously as Quercus). The Goodhope oak clusters about several springs on the east side of Goodhope Bay in the Glen Canyon National Recreation area. One of the stands has been burned by negligent recre- ationists, but all bear the large acorns atypical of Q. gambelii from other sites. Seeds taken at the time of the original collection were grown in the greenhouse at Brigham Young Univer- sity by Mr. Tom Black, who planted them simultaneously with others of Q. gambelii from Utah County. By May of the year follow- ing, the seedlings of the Goodhope oak were | twice as large as those from Utah County. The | plants at Goodhope Bay tend to average larger than those from other localities along Glen Canyon, but they do not appear to differ oth- erwise. The size might be a function of the continuous water supply available in the | spring and seep areas. The spring area seems to be associated with joint systems in the Wingate Formation to the east, these allowing water to penetrate to the impervious Chinle Formation, where the water surfaces. Quercus havardii Rydberg. Shinnery Oak. | [Q. undulata authors, not Torr. ]. Clonal, de-. ciduous, sand-binding shrubs, or, less com- monly, small trees to 2 m or more; leaves) densely grayish to yellowish stellate-hairy on both sides when young, less densely so in age, . but only slightly, if at all, paler beneath than) above, even in age, 1.5—5.5 cm long, 0.9-3.3) cm wide, oblanceolate to elliptic in outline, | with usually 6-10 toothlike lateral lobes, these typically apiculate-acuminate and sometimes further notched or toothed api-| cally; catkins 1-2.5 cm long; involucral cups’ 7-10 mm long, 14—18 mm wide, subsessile, ca f | | | January 1986 1/4—1/3 the length of the acorn, clothed with imbricate, densely hairy scales; acorns 15—23 mm long, 14-18 mm thick. Blackbrush, ephe- dra, vanclevea, purple sage, and pinyon-ju- niper communities, usually in sand, at 1,125 to 2,135 m in Emery, Garfield, Grand, Kane, San Juan, and Wayne counties; Arizona, New Mexico, Oklahoma, and Texas; 54 (x). The _ shinnery oak, as it occurs in Utah and adjacent Arizona, is more or less influenced by in- tergradation with the partially sympatric Q. gambelii and Q. turbinella (Tucker 1970). In- termediates between both of those parental types and Q. havardii are known. However, in the sandy footslope of the San Rafael Swell -in Emery and Wayne counties and adjacent portions of the Navajo Basin of Utah and Ari- zona, the species is more or less stable and tends to be habitat specific. Possibly because of the contribution of both |Q. gambelii and Q. turbinella, even the more uniform portions of the species differ from the body of the taxon lying far to the east in New Mexico, Texas, and Oklahoma. The Navajo Basin material differs from the parental type in smaller and more densely hairy leaves that tend to be sharply toothed. Because of the differences noted and because of the isolation of the Navajo Basin material from its eastern counterpart, our material is regarded as Var. | tuckeri Welsh var. nov. Planta inter Querco gambelii Nutt. et Q. turbinella Greene et Q. havardii Rydb. var. havardii sed in pro max _parte Q. harvardii Rydb. var. harvardii maxime affini, sed in foliis minoribus et pilis plus dense, et dentibus acutis differt. TyPE: USA: Utah. San Juan Co., Low Pass, ca 8 mi _ SE of Moab, T27S, R23E, S.5, mixed desert -shrub community, Entrada Sandstone For- - mation, at 1,635 m, 10 July 1985, S.L. Welsh and L. C. Higgins 23630 (Holotype BRY; Iso- types to be distributed). Additional speci- mens: Utah, San Juan County, Elephant Hill vicinity, im pinyon-juniper community, - Canyonlands National Park, 6 May 1969, S. L. Welsh, D. Atwood, & L. Higgins 8878 (stami- nate, BRY); Garfield Co., Henry Mts., 5 May 1977, E. Neese & S. White 2784. Grand Co., Moab, 8 June 1927, W. P. Cottam, 2139; ‘ Courthouse Towers, 25 Apr 1947, B. F. Har- rrison 11124; Arches National Park, 2 ,May | 1963, L. B. Barnett et al. 56, 66; 13 July 1972, |J.S. Allan 128; 19 July 1972, J. S. Alan 133; 5 WELSH: UTAH FLORA, QUERCUS 109 mi NW of Moab, 21 May 1984, D. Atwood et al. 9699. Kane Co., 28 mi e of Kanab, 11 May 1953, B. F. Harrison 12049: 60 mi SE of Es- calante, 7 Aug. 1957, B. F. Harrison 12723: Dance Hall Rock, 2 May 1962, J. R. Murdock 381; Hole-in-the-Rock, 4 May 1962, D. A. White 125, 27 Apr 1977, R. Foster 3602; Es- calante Arm of Lake Powell, 5 June 1927, S. L. Welsh & G. Moore 11825, 11826. San Juan Co., Bluff, 2 July 1927, W. P. Cottam 2529, 2532; Monument Valley, 4 July 1927, W. P. Cottam & S. Hutchings 2585; Cow Canyon, 29 Apr 1961, C. A. Hansen 97; Squaw Flat, 15 May 1965, G. Moore 375. All are deposited at BRY. Quercus pauciloba Rydb. (hybrid) Clonal, semievergreen shrubs or small trees mainly 2—4 m tall and with trunks 4-15 cm in diame- ter; leaves stellate-hairy on both surfaces when young, becoming sparingly so to glabrate on one or both sides in age, bicolored (more or less), typically green to dark green above and paler beneath, 2-10 cm long, 1-7 cm wide, usually with (0) 4—8 toothlike lateral lobes, these typically apiculate and some- times apiculate-acuminate, rarely some of them again notched or toothed; staminate catkins 3-4 cm long; pistillate catkins, mat- ure involucral cups, and mature acorns not present in specimens examined. Sagebrush, mountain brush, pinyon-juniper, and pon- derosa pine communities at 1,220 to 2,045 m in Beaver, Iron, Juab, Kane, Millard, Salt Lake, Tooele, Utah, Washington, and Weber counties; Colorado, Arizona, and Nevada; 11 (0). Specimens designated as Q. pauciloba consist of an aggregation of hybrids and pre- sumed introgressants involving Q. gambelii and Q. turbinella as parental types. They oc- cur relatively commonly in areas where the two parental species coexist in Washington and Kane counties, but they occur also along the western margin of the plateaus and moun- tains north to Weber County, far removed from the body of Q. turbinella in Washington, Kane, and San Juan counties. The hybrids and introgressants are about on a line marking the edge of the major overthrust fault that bisects Utah. Cottam et al. (1959) and Tucker (1961a, 1961b, 1970) have postulated an interglacial advance of Q. turbinella into the Great Basin, followed by a retreat during return of harsh conditions. The hybrids were presumed to 110 represent first generation only, were postu- lated to have resulted during the incursion, and were judged to have persisted following the retreat of the one parental type. Ruled out are other possibilities such as long-distance pollination, suggested as a probability by Harper et al. (1985) and concurred in by me, because of lack of coincidence of flowering times, primarily. Examination of specimens of the hybrid complex demonstrates several problems. First, the variation among the intermediates is greater than would be expected from first generation hybrids, suggesting introgression as well as hybridity. Indeed, a specimen from the mouth of American Fork Canyon (Lind- quist s.n. 20 Jan 1981 BRY) is strictly ever- green, has staminate catkins and abortive acorns, and, if placed within specimens of Q. turbinella from Washington County, would be identified as that species. Second, al- though mature acorns and caps were not noted on the specimens examined, staminate catkins were present in some, suggesting the possibility of introgression occurring far north of the primary pollen source. And, the follow- ing questions are unanswered by the flow and ebb hypothesis: (1) Did a migration route oc- cur during the time of the thermal maximum that is not now in evidence? (2) Was not a simpler and as equally accessible a route avail- able along the canyons of the Colorado (where Q. turbinella exists in some small part, even now) and, if so, where is the matching set of hybrids in the Colorado Basin? (3) Could not long-distance pollination, even though the juxtaposition of flowering time occurs only irregularly, be sufficient to account for some, if not all, of the occurrences of the hybridiza- tion northward? And does not the pattern of distribution of hybrids along the windward side of the Wasatch frontal ranges suggest long-distance pollination? Additional studies are indicated. Quercus eastwoodiae Rydb. (hybrid) In the Navajo Basin of Utah, along the Colorado River and its canyons, another set of hybrids and presumed introgressants are known. In general aspect and leaf morphology they re- semble Q. pauciloba, but their origin seems to be different. Gambel oak grows at even the lowermost elevations in mesic canyons, on stream terraces, and around seeps, springs, GREAT BASIN NATURALIST Vol. 46, No. 1 and hanging gardens. Although Q. turbinella is present also, the extent and total numbers appear to be limited now, and there does not seem to be evidence of a much greater inci- dence in the past. The turbinella live-oak is present at the confluence of Glen Canyon and the San Juan River, and also occurs as a nar- row tongue along the Cockscomb in central Kane County, where it is confluent southward into the Houserock Valley area of Arizona. Intermediates between turbinella live-oak and Gambel oak are also known from this latter region. In the remainder of the Navajo Basin the picture is complicated by still another Quer- cus species, i.e., the shinnery oak, Q. havar- dii. That species occurs as a moderately stable _ entity on the sandy plateaus and slopes away from the canyons proper but reaches to their margins, where the stream courses are deeply entrenched. They, too, hybridize with Gam- bel oak, and the hybrids and presumed intro- gressants survive in usually sheltered posi- tions adjacent to or intermixed with Gambel Oak. Such intermediates are known also from the islandlike mountains that protrude from the floor of the Navajo Basin. Intermediate specimens appear superficially like those of | Q. pauciloba, but they tend to be more | densely hairy, to have hairs persistent on the : upper surface, even in age, and to be less distinctly green, even if bicolored. These in- termediates are here designated as Q. x east- | woodiae Rydb. (Bull. New York Bot. Gard. 2: 210. 1901; based on, Eastwood 141, from But- — ler Wash, San Juan County, Utah in 1895), — although the description of the type specimen _ indicates it to be a portion of the variation nearer to Q. gambelii. The plants are present | in blackbrush, other warm-desert shrub, | mountain brosh: pinyon-juniper, and Doug- las-fir communities at 1,130 to 1,830 m in co) field, Grand, Kane, and San Juan counties; Arizona; 13 (ii). Quercus turbinella Greene Turbinella | Live-oak. Clump-forming (clonal?) evergreen | | shrubs or, less commonly, small trees, mainly | 1—4 dm tall and with stem diameters to 2 dm; _ leaves yellowish stellate-hairy on both sur-_ faces when young, finally glabrate and glau-. cous above, not especially bicolored, typically 1.3-4cm long, ().7—2.4 cm wide, lanceolate to. oblong or suborbicular in outline, with 2-6 January 1986 WELSH: UTAH FLORA, QUERCUS alt pairs of lateral, spine-tipped teeth or entire; staminate catkins 1-3 cm long; involucral cup 6—8 mm long, 10-14 mm wide, ca 1/4 as long as the acorn; acorns 12-24 mm long, 7-10 mm thick. Chaparral (oak, manzanita, ceanothus), pinyon-juniper, and riparian communities at 820 to 1,710 m in Kane, San Juan, and Washington counties; Nevada and Arizona; 39 (iv). LITERATURE CITED Cottam, W. P., J. M. TUCKER, AND R. DRosNIck. 1959. Some clues to Great Basin postpluvial climates provided by oak distributions. Ecology 40: 361-377. HARPER, K. T., F. J. WAGSTAFF, AND L. M. KUNZLER. 1985. Biology and management of the Gambel Oak veg- etative type: A literature review. Gen. Tech. Rep. INT-179. 31 pp. TUCKER, J. M. 196la. Studies in the Quercus undulata complex. I. A preliminary statement. Amer. J. Bot. 48:202—208. . 1961b. Studies in the Quercus undulata complex. II. The contribution of Quercus turbinella. Amer. J. Bot. 48:329-339. - 1970. Studies in the Quercus undulata complex. IV. The contribution of Quercus harvardii. Amer. J. Bot. 57:71-84. . 1971. Studies in the Quercus undulata complex. V. The type of Quercus undulata. Amer. J. Bot. 58:329-341. SEASONAL PHENOLOGY AND POSSIBLE MIGRATION OF THE MOURNING CLOAK BUTTERFLY NYMPHALIS ANTIOPA (LEPIDOPTERA: NYMPHALIDAE) IN CALIFORNIA Arthur M. Shapiro! ABSTRACT. —Circumstantial evidence is presented that suggests the Mourning Cloak undergoes regular seasonal up- and downslope movements in northern California. The species breeds at low elevations in spring and then disappears until autumn; its disappearance coincides with the appearance of fresh individuals in the Sierra Nevada alongside obvious hibernators. The Mourning Cloak or Camberwell Beauty, Nymphalis antiopa L., is one of the most characteristic Holarctic butterflies; it ranges from the subarctic to the subtropics. Nonetheless, its basic biology is poorly known. There have been many discussions of its rare and intermittent occurrence in the British Isles (Williams et al. 1942, Williams 1958, Chalmers-Hunt 1977), but only one de- tailed description of its natural history—that by Young (1980), who narrated the situation in Wisconsin, USA, in the hope that it would help Palearctic workers understand the dy- namics of their own populations. The biology of N. antiopa in California, USA, is quite dif- ferent; specifically, it seems to involve either estivation or altitudinal migration or both. Al- titudinal migration appears to occur in Cali- fornia populations of N. (Aglais) milberti Latr. and N. californica Bdv. (Shapiro 1973, 1974a, 1974b, 1975, 1979, 1980). Since 1972 phenological data have been taken for all butterflies at a series of stations forming a transect parallel to Interstate High- way 80 from sea level at the Suisun Marsh, Solano County, to tree-line at Castle Peak, Nevada County (2750m). Each station is vis- ited at roughly two-week intervals throughout the butterfly season, and all species flying are recorded. Figures 1 and 2 represent the N. antiopa data from this transect for 1983 and 1984. These two years were extremely differ- ent meteorologically and essentially embrace the range of variation observed during the 13 years of the study. The year 1983 was one of ‘Department of Zoology, University of California, Davis, California 95616. record high precipitation, with both rainfall _ and snowpack greater than 200% of the 30- — year means. Summer was cool and unusually moist, after a very late and cloudy spring. Precipitation in the 1983-84 season was | slightly below normal. Rain- and snowfall | were heavy before Christmas and nearly | nonexistent thereafter. Spring was early and hot, and summer 1984 was the hottest of record (over 125 years) at low elevations and much warmer than normal in the mountains, with an unusual frequency of thunderstorms. The fact that the seasonal patterns of N. an- | | | tiopa are consistent in two such different years _ suggests that they accurately represent the — seasonal dynamics of the animal. Shapiro (1974c) reported that N. antiopa was univoltine in the Sacramento Valley de- spite the very long growing season. Nearer the coast, at the Suisun Marsh, Shapiro | (1974d) reported essentially the same phenol- | ogy. The same pattern was again reported for | suburban Sacramento by Smith (1983). Smith provides counts of sightings at one specific ! locality for the years 1970 through 1982. His | pattern is quite consistent, with no animals | seen after 8 July in 50% or more of years, after} a very dramatic peak between 20 May and 1 July. It was initially assumed by Shapiro that the single spring brood in these areas entered estivation, followed by a brief period of activ- ity in autumn, followed by hibernation, such. that Sacramento Valley animals lived a full year as adults. Although estivation remains a) possibility, no estivating adult has been found — { i ) 112 | | | } | | January 1986 SHAPIRO: MOURNING CLOAK 113 Castle Peak NONE SEEN IN 1983 (2750 m) Donner Pass (2100 m) Lang Crossing (1500 m) Sierra Nevada W slope Coast Range (250m) Sacramento Valley (Om) Jim RoaM) oAc Maids. VIRy AL S230 4N:; iD Fig. 1. Phenology of Nymphalis antiopa along a transect across northern California in 1983. Intensity of stippling indicates degree of wing wear. Dots are individual occurrences. Triangles are observations of larval colonies: L;=third instar, etc. | 3 ro - | 6 Castie Peak NO HIBERNATORS pa & (2750m) s | 3 Donner Pass 1! 3 (2100 m) > 0 HH < Lang Crossing i = (1500 m) | d Coast Range . i (250m) h Sacramento Valiey (Om) | | Fig. 2. Same for 1984. 114 in 13 years, the condition of the adults observed in autumn and early winter argues against it, and, most importantly, there is a persuasive cir- cumstantial case implicit in Figures 1 and 2 for altitudinal migration on a regular seasonal basis. Overwintered adults in typical posthiberna- tion condition (borders nearly or totally white, much wing wear) were observed near Sacra- mento and in the Inner Coast Range beginning in late winter in both years. Similar hibernators appear at montane sites shortly after snowmelt—the timing of which varied by sev- eral weeks between the two years. The single flight of fresh animals was observed at low eleva- tions in late May—early July, lasting from 10 to 41 days in different locations. Smith (personal com- munication) saw singletons in Sacramento dur- ing the second, third, and fourth (three records) weeks of July and the second week of August, 1983, and on 9 July and 25 July 1984. As can be seen from Figures.1 and 2, at moun- tain stations more or less fresh animals can also be found in spring, flying with the hibernators. At Lang Crossing (1500 m), for example, the first 1984 hibernators appeared on 10 March and fresh animals followed on 6 May while hiberna- tors were still numerous. At Donner Pass (2100 m), both fresh and worn animals were already flying 27 May and continued to be distinguish- able through 20 June. Thereafter, worn animals probably derived from both groups flew until 20 July. On that date two colonies each of 3rd- and Sth-instar larvae were censused, presumably representing a differential in the timing of repro- duction by the two groups of adults. On 15 Au- gust fresh adults were very numerous in the vicinity of the previous L; colonies. By 6 Sep- tember only a few were present and these were moderately worn, but a second group of fresh animals appeared 19 September and flew for a week. At the highest station, Castle Peak (2750 m), there is no permanent population, and the spe- cies is present in some years (generally those with early snowmelt) and absent in others (gen- erally late years). It was absent in 1983. In 1984 no hibernators were seen, but fresh animals flew 27 July— 17 August, and on the latter date one colony of L, was found. Given that reproduction by N. antiopa was late at low elevations in 1983 and that the timing of snowmelt was even more distorted, it is striking that in both years figured (and in GREAT BASIN NATURALIST Vol. 46, No. 1 general) the appearance of fresh animals at mountain stations commences about a week. after the valley brood appears. | Similarly, the widely scattered autumn sightings are properly timed to represent downslope dispersal by the midsummer brood reared in the mountains. Several appar- ently fresh antiopa were seen at Lang Cross- | ing 30 August (two weeks after the flight started at Donner; no larval colonies were | seen at Lang in 1984). One apparently fresh | specimen was seen in Gates Canyon in the| Inner Coast Range on 26 August, and Smith’ | saw one in Sacramento on 4 September. | Nothing in these data either requires altitu- | dinal migration or precludes estivation, but | they are clearly suggestive. How plausible is. the hypothesis of altitudinal migration? One| alternative explanation is that antiopa may over-winter in both the adult and pupal stages | in the Sierra Nevada. Klots (1951) raised this | possibility for the eastern United States. | Shapiro (1969) recorded a single case in cen- | tral New York in which overwintering of the | pupa seems inescapable; the animal, when: captured, voided meconium. I have tried re-| peatedly to maintain laboratory-reared low- | land California pupae at 2—3 C for extended | periods, but with no success after eight! weeks. True pupal diapause appears to be| unknown in the Nymphalini. Another alternative explanation for the low- | altitude phenomena is a low level of reproduc- | tion in summer. In 13 years of extensive field | work in the Sacramento Valley and Inner. Coast Range, I have never found a single lar- | val colony after 1 June, nor has Smith in 14, years at Sacramento. On the other hand, sec- | ond broods occur occasionally beginning at Fairfield, Solano County, nearer the coast, © and even a third brood has been recorded (Lz, ' 20 October 1980). This is apparently standard | on the immediate coast, for Tilden (1965) re- ports “two to three broods a year” in the San. Francisco Bay area, and Emmel and Emmel (1973) report “multiple-brooded at lower ele- vations and single- or double- brooded in ; higher zones” in southern California—the “lower elevations” referring almost entirely to ‘ urban and suburban areas, not to the interior — deserts. Nymphalis antiopa is a common ur- ban species in Mexico City, where it breeds | all year. | i \ January 1986 Some light can be shed on the annual cycle by _ examining the reproductive status of the ani- mals. Although Young (1980) claims that spring populations consist solely of females—the males presumably having died overwinter—this is not true in California (or southeastern Pennsylvania or New York). In the Sacramento Valley and Coast Range, courtships and matings are ob- served both in autumn (occasionally) and spring (frequently). At Donner Pass they have been observed in September but not in spring. In 1984 I dissected eight August and three Septem- ber Donner females. All had large amounts of fat and no well-developed oocytes, but all the Au- gust and none of the September females were virgins. All the fresh-looking females collected in the mountains in June and July have been fully reproductive (N= 15). Herman and Bennett (1975) reported that summer females (source population and rearing regime unspecified) eclose with large fat bodies and no oocytes, and subsequently mature as a function of environment. Matura- tion was essentially completed within 10 days at 25 C on LD 16:8 but did not occur after 14 days at 10 C, LD 8:16. This experiment does not separate the effects of photoperiod and temperature. Photoperiods at Donner Pass in late August are ca LD 13.5:10.5, mean tem- perature 16 C. Donner Pass antiopa reared on LD 14:10 at 25 C failed to mature after 15 weeks in the dark at 2 C. For adult antiopa emerging in the Sacra- mento Valley in late May, photoperiods are ca LD 15:9 and mean temperature 18.9 C, con- ditions that should permit rapid gonadal mat- uration, but no reproduction is seen. Of three late May—early June Sacramento females, two had mated and one of these showed early oocytes. If altitudinal migration is real, ovarian maturation may occur during the up- slope flight, its termination coinciding with the beginning of oviposition. The distances — involved (125-150 km from our Sacramento Valley sites to Donner Pass) could be tra- ’ versed in a week or so, based on the progress of N. californica migrations I have tracked. Shannon (1917) believed eastern U.S. pop- ulations of N. antiopa were at least somewhat migratory. Gibo (1981) records the species riding thermals in east central Canada; such passive soaring is associated in many insects with the initiation of long-range dispersal. SHAPIRO: MOURNING CLOAK 115 One additional aspect of the problem de- serves mention. Shapiro (1981a,b) studied the canalization of the wing pattern as a trait adap- tive to climate. He found that Alaskan animals are more strongly buffered physiologically against cold shock than either lowland or Sier- ran montane N. antiopa. The similarity of the physiological responses of the Californian broods, from radically different climates, could imply gene flow over the fairly short distances separating them—an interpretation consistent with the notion of altitudinal migra- tion. Since the 1981 papers were published, the California experiments have been repli- cated three times with the same results for the major aberration “hygiaea.” The minor pat- tern differences reported in Shapiro (1981b) have been inconsistent among broods. Granted that at least part of the population of N. antiopa overwinters at 2100 m each year, gene flow between these and animals dispers- ing from the lowlands could be very substan- tial in some years. Such gene flow—depend- ing on the timing of spring at both elevations—would be expected to inhibit if not prevent genetic differentiation along our altitudinal transect. Numbers of this species are consistently too low for mark-recapture experiments to hold much promise as a test of the altitudinal-mi- gration hypothesis. If genetic markers can be found to facilitate identification of low-eleva- tion animals, they would be useful for docu- menting movements—but if gene flow is fre- quent, such markers are unlikely to be found. In the meantime, detailed seasonal data for a variety of localities are very desirable. ACKNOWLEDGMENTS This research forms part of the California Agricultural Experiment Station Project CA- D*-AZO-3994-H, “Climatic Range Limita- tion in Phytophagous Lepidopterans.” I thank Leslie V. Smith for making his Sacramento data available for use in this paper. LITERATURE CITED CHALMERS-HUNT, J. M. 1977. The Camberwell Beauty in Yorkshire in 1977. Ent. Rec. J. Var. 89:348. EMMEL, T. C., AND J. F. EMMEL. 1973. The butterflies of southern California. Natural History Museum of Los Angeles County 148 pp. 116 G1po, D. L. 1981. Some observations on soaring flight in the Mourning Cloak butterfly (Nymphalis antiopa L.) in southern Ontario. J. New York Ent. Soc. 89:98—-101. HERMAN, W. S., AND D. C. BENNETT. 1975. Regulation of oogenesis, female specific protein production, and male and female reproductive gland development by juvenile hormone in the butterfly Nymphalis antiopa. J. Comp. Physiol. 99:331—338. Kxots, A. B. 1951. A field guide to the butterflies of North America, east of the Great Plains. Houghton Mif- flin, Boston. 349 pp. SHANNON, H. J. 1917. Autumn migration of butterflies. Amer. Mus. J. 17:32—40. SHapiro, A. M. 1969. Overwintering by pupal Nympha- lids in New York? Ent. News 80:130. . 1973. Altitudinal migration of butterflies in the central Sierra Nevada. J. Res. Lepidopt. 12: 231-235. . 1974a. Movements of Nymphalis californica in 1972 (Nymphalidae). J. Lep. Soc. 28:75—78. . 1974b. Altitudinal migration of central California butterflies. J. Res. Lepidopt. 13:157—161. . 1974c. The butterfly fauna of the Sacramento Val- ley, California. J. Res. Lepidopt. 13:73-82, 115-122, 137-148. . 1974d. Butterflies of the Suisun Marsh, California. J. Res. Lepidopt. 13:191—206. GREAT BASIN NATURALIST Vol. 46, No. 1 ———. 1975. Why do California tortoiseshells migrate? J. Res. Lepidopt. 14:93-97. ——__—.. 1979. Nymphalis milberti (Nymphalidae) near sea level in California. J. Lepidopt. Soc. 33:200—201. —___. 1980. Mediterranean climate and butterfly migra- tion: an overview of the California fauna. Atalanta 11:181-188. .198la. Canalization of the phenotype of Nymphalis antiopa (Nymphalidae) from subarctic and montane climates. J. Res. Lepidopt. 19: 82-87. . 1981b. Phenotypic plasticity in temperate and subarctic Nymphalis antiopa (Nymphalidae): evi- dence for adaptive canalization. J. Lepidopt. Soc. 35:124-131. SMITH, L. V. 1983. A twelve-year count of three California butterflies. J. Lepidopt. Soc. 37:275—280. TILDEN, J. W. 1965. Butterflies of the San Francisco Bay Area. University of California Press, Berkeley. 88 pp. WILLIAMS, C. B. 1958. Insect migration. Wilmer Bros., Harram, London. 235 pp. WILLIAMS, C. B., G. F. COCKBILL, M. E. GIBBS, AND J. A. Downes. 1942. Studies in the migration of Lepi- doptera. Trans. R. Ent. Soc. London 92:101—283. YouNG, A. M. 1980. Some observations on the natural history and behaviour of the Camberwell Beauty (Mourning Cloak) Butterfly, Nymphalis antiopa (Linnaeus) in the United States. Ent. Gaz. SEASONAL MICROHABITAT RELATIONSHIPS OF BLUE GROUSE IN SOUTHEASTERN IDAHO Dean F. Stauffer’? and Steven R. Peterson!® ABSTRACT. —Microhabitat characteristics of blue grouse (Dendragapus obscurus) were analyzed in breeding and wintering habitats in southeastern Idaho. Breeding habitats typically were open sagebrush (Artemisia spp.), mixed shrub, mountain mahogany (Cercocarpus ledifolius ), and maple (Acer grandidentatum) stands on east to south facing aspects of slopes below 2100 m elevation. Breeding blue grouse selected areas with approximately a 50:50 or greater open to cover ratio. Blue grouse selected areas with higher tree coverage than available on average within the mixed shrub vegetation type. Hens with broods preferred sites with relatively tall (>50 cm) herbaceous vegetation. During autumn and winter, blue grouse preferred high elevation (>2285 m) stands of open (50% tree cover) conifer. Douglas-fir (Pseudotsuga menziesii) were preferred as winter roost trees. Sites selected in winter had significantly more Douglas-fir than those selected in autumn. Blue grouse occur throughout western North America. Substantial work on this spe- cies has been conducted on Vancouver Island, British Columbia (e.g., Bendell and Elliott 1966, 1967, Fowle 1960, Zwickel and Bendell 1967, Lewis and Zwickel 1980). Blue grouse also have been studied throughout the Rocky Mountains (Marshall 1946, Caswell 1954, Heebner 1956, Blackford 1958, Boag 1966, Maestro 1971, Harju 1974, Weber 1975). Most reports have concerned breeding be- havior, with relatively little work being done on habitat requirements. Studies on blue grouse habitat typically have been qualitative in nature, relating grouse to general habitat categories (e.g., Marshall 1946, Caswell 1954, Heebner 1956, Bendell and Elliott 1966) or breeding habitat (Mussehl 1960, 1963, Maestro 1971, Martinka 1972, Weber 1975, Lewis 1981). Except for some analyses of male hooting sites (Martinka 1972, Lewis 1981), little quantitative information on blue grouse has been reported. To adequately manage habitat for blue grouse, we must know their relationship to patterns of macro- and microhabitat charac- teristics. We previously described the macro- habitat relationships of blue grouse in south- eastern Idaho (Stauffer and Peterson 1985). Here we address the microhabitat character- istics of blue grouse. Our objectives are to quantitatively describe habitats used by blue grouse for breeding and wintering and to com- pare characteristics of used habitats to avail- able habitats. Funding was provided by the USDA Forest Service. All personnel of the Montpelier Ranger District are gratefully acknowledged for their assistance throughout the fieldwork. B. S. Cade and B. A. Schrader were field assistants. Reviews of the manuscript by J. J. Spillett, E. O. Garton, M. Hironaka, E. G. Bizeau, R. K. Steinhorst, B. R. Noon, P. G. Stettenheim, and G. H. Cross are appreci- ated. STUDY AREA AND METHODS We examined blue grouse habitat relation- ships on the western portion (108,000 ha) of the Montpelier District of the Caribou Na- tional Forest, Bear River Range of the Wasatch Mountains in southeastern Idaho. We classified the study area into eight rela- tively discrete vegetation types based on the dominant (according to density) tree and shrub species. Four open vegetation types (44% of the area) were most common at lower (<2130) elevations: sagebrush, mixed shrub, mountain mahogany, and bigtooth maple. Four forested vegetation types (56% of the area) were most common at mid and high ‘Department of Fisheries and Wildlife, University of Idaho, Moscow, Idaho 83843. “Present address: School of Forestry and Wildlife, VPI & SU, Blacksburg, Virginia 24061. 3Present address: Game Research, Game Division, Department of Fish and Game, Box 3-2000, Juneau, Alaska 99801. 117 118 elevation (>2130 m): aspen (Populus tremu- loides), aspen/conifer mixed; dense conifer; and open conifer. We have described the floristic character of each vegetation type else- where (Stauffer and Peterson 1985). We spent 1593 h (spring, 322 h; summer, 543 h: autumn, 296 h; winter, 432 h) searching for grouse from May 1979 through May 1981. Searching effort was distributed among the vegetation types in approximate proportion to their occurrence on the study area. Each time a grouse (or group) was flushed, we used the location as the center of a 0.01 ha circular plot for which we recorded: percent of area within 40 m composed of coniferous or deciduous cover or open; canopy height and average height of herbaceous vegetation; number of woody stems <7 cm dbh in 2 per- pendicular arm-width transects across the plot; number of trees by species within 6 dbh categories (7.0-15.0 cm, 15.1—23.0 cm, 23.1—38.0 cm, 38.1—53.0 cm, 53.1—-69.0 cm, >69 cm dbh); and vegetation type. We recorded 120 sets of plot data at random locations, 60 in the mixed shrub type and 60 in the maple vegetation type, to sample breed- ing habitat characteristics available to blue grouse. We calculated means for data recorded in the 0.01 ha plots at grouse locations for various combinations of vegetation types and season to describe the characteristics of sites se- lected. For 38 winter roost trees, we mea- sured a second set of plot data at the nearest potential roost tree that had no evidence of use. Additional data recorded at roost trees included tree species, diameter, and pres- ence or absence of dwarf mistletoe (Arceutho- bium spp.) infestation. We evaluated differ- ences between used and unused sites with a paired t-test. Prior to statistical analysés, all data were checked for normality, and those variables found to be nonnormal were transformed (log, square-root, or arc-sine) to achieve a more normal distribution. RESULTS AND DISCUSSION Blue grouse used a variety of vegetation types (Table 1). The open vegetation types agebrush, mixed shrub, mountain ma- gany, and maple) were used primarily dur- GREAT BASIN NATURALIST Vol. 46, No. 1 ing spring and summer. These types consti- tute breeding habitat. The dense and open conifer types were used most heavily in fall and winter, although the open conifer type also was used in spring and summer. These use patterns are similar to those recorded elsewhere for the intermountain region (Mar- shall 1946, Caswell 1954, Heebner 1956, Mussehl 1960, 1963, Boag 1966, Zwickel et al. 1968, Maestro 1971, Harju 1974, Weber 1975). In spring and summer, junipers (Juniperus spp.) and bigtooth maple were most com- monly associated with blue grouse (32% and 52% of 227 observations, respectively). We found Douglas-fir and subalpine fir (Abies la- siocarpa) at 73% and 53% of 191 fall and win- | ter observations, respectively. Additionally, | limber pine (Pinus flexilis ) was noted at 47% of — 57 winter Blue Grouse observations (Stauffer | 1983). Common shrubs at 227 blue grouse — locations in spring and summer were sage- | brush (72% occurrence), snowberry (Sym- | phoricarpos spp., 54%), bitterbrush (Purshia | tridentata, 27%). Snowberry (72%), sage- | brush (57%), chokecherry (Prunus virgini- ana, 28%), and snowbrush (Ceanothus veluti- | nus , 19%) most commonly occurred at 134 fall - observations (Stauffer 1983). CHARACTERISTICS OF BREEDING HABITAT Sites used by blue grouse (Table 2) differed | among the four open vegetation types, based | on 10 microhabitat characteristics | Multivari- | ate Analysis of Variance (MANOVA, P<001)]. — Although habitat characteristics differed | among the vegetation types, used sites were | all relatively open, with the highest cover of | about 60% conifer and deciduous cover occur- | ring in the mountain mahogany vegetation — type. Use of open areas by blue grouse during | spring and summer has been documented | : throughout their range (Marshall 1946, © Caswell 1954, Mussehl 1960, 1963, Boag! 1966, Zwickel et al. 1968, Martinka 1972, | Harju 1974, Weber 1975, and Lewis 1981). _ We found no difference between spring and | summer sites used by blue grouse in sage-| | brush (Hotelling’s T*, P>0.05). The data re-| flected the openness of this habitat (Table 2), | but the presence of some trees indicated that’ areas with at least some taller cover were pre-. i January 1986 STAUFFER, PETERSON: BLUE GROUSE ECOLOGY 119 TABLE 1. Distribution of blue grouse plot observations among the vegetation types studied; southeastern Idaho, 1979-1981. Percent of observations Vegetation type Brood Hooting Spring Summer Autumn Winter | Sagebrush 12 W 25 14 2 0 Mixed shrub 9 25 17 14 6 0 ' Mountain mahogany 0 21 Hi 9 11 0 _ Maple 36 45 34 35 2 0 , Aspen 9 0 0 3 4 0 _ Aspen/conifer 3 0 4 ] 8 0 | Dense conifer 3 0 0 3 14 18 ( Open conifer 27 2 13 23 54 82 | Number of observations 33 44 71 80 134 56 Spring and summer data exclude brood and hooting observations. TABLE 2. Means of habitat characteristics recorded for 0.01 ha circular plots at blue grouse locations and random locations in open habitats in southeastern Idaho, 1979-1981. Mountain Sagebrush mahogany Mixed shrub Maple Spring-autumn Random obs. Spring-summer Random obs. Variable n= 33 n =36 n= 45 n= 60 n= 84 n= 60 Coniferous cover (%) 2.6(0.6)* 4.0(1.1) 3.1(0.9) * 0.2(0.1) 5.0(0.6) 5.8(0.8) Deciduous cover (%) 8.4(1.7) 56.0(2.5) 26.4(2.8) *15.4(2.0) 43.6(1.9) 49.4(3.1) \ Open (%) 88.8(1.8) 40.0(2.4) —-70.5(2.7) —-*84.0(2.0) 51.4(1.8) 44.8(3.1) | Tree canopy cover (%) 8.2(3.3) 36.9(4.7) 8.3(2.4) * 4.1(1.4) 23.8(2.5) 24.6(4.0) Ground cover (%) 48.0(4.4) 42.6(3.9) 49.0(3.2) *57,2(2.1) 56.5(2.5) *78.6(2.1) Canopy height (m) 1.3(0.2) 3.4(0.1) 2.0(0.2) * 1.5(0.1) 3.5(0.2) 3.6(0.3) Stems <7cmdbh/ha _ 847(372) 2428(406) 7400(1590) *4056(460) 4395(435) 4092(462) Trees/ha 15(8) 352(50) (19) 40(19) 249(36) 305(60) - Coniferous trees/ha 12(6) 22(10) (2) (0) 27(9) 18(7) | Deciduous trees/ha 3(3) 330(49) (19) 40(19) 222(35) 287(60) ~ Standard error. *Indicates a significant (p<0.05) difference based upon a t-test between the grouse observations and random observations in the vegetation type. ferred. Mussehl (1963) and Weber (1975) noted that blue grouse often were found near clumps of trees in sagebrush stands. Mountain mahogany was used _ spring through autumn and had the highest tree _ cover of the four open vegetation types (Table 2). Percent ground cover was the only charac- ' teristic that differed among spring, summer, and autumn observations and was highest in summer and autumn (47.8% and 56.0%, re- spectively) and lowest in spring (22.1%). Twenty-one percent of the hooting observa- tions were in mountain mahogany, but no broods were found here. Sites selected by blue grouse in the mixed shrub vegetation type did not differ among spring, summer, and autumn observations /(MANOVA, P>0.05). However, microhabi- «tat characteristics of sites used were different i from a random sample of 60 sites in this type \(Hotelling’s T’, P<0.001, Table 2). Percent coniferous and deciduous cover, percent tree canopy cover, and density of small stems were higher and percent open area and ground cover were lower at sites used by grouse than at random sites. Thus, blue grouse are select- ing areas within the mixed shrub vegetation type with higher than average woody cover (see also Weber 1975). We found differences in sites used between spring and summer by blue grouse in the maple vegetation type (Hotelling's 1 Pix 0.01). Percent coniferous cover and density of coniferous trees were higher and percent de- ciduous cover was lower at sites used in spring (spring x = 7.1%, 59/ha, and 37.6%, respec- tively; summer < = 3.4%, 2/ha, and 48.3%, respectively; df = 79 and t = 3.2, 4.1, and 2.3, respectively.) During spring, grouse often were associated with junipers in the maple type, which may provide cover prior to leaf- out of the deciduous trees. Weber (1975) 120 found that male blue grouse often were associ- ated with junipers on breeding areas in Utah. Except for percent ground cover, which was lower at used sites, (t = 6.1, P<0.01), habitat characteristics were not different between random sites and those used in maple vegeta- tion types (Table 2). Thus, blue grouse were not selecting for any particular characteristic of the maple vegetation type. These open types provide suitable habitat for hooting by male blue grouse. Lewis (1981) reported tree cover of 6.6% and canopy height of 3.3 m at hooting sites on Vancouver Island. In Montana, Martinka (1972) found a tree crown cover of 30% at male display sites and Maestro (1971) noted that breeding blue grouse preferred areas of 41%-50% tree cover in Utah. These values are comparable to the characteristics of habitats where we found blue grouse breeding in southeastern Idaho (Table 2). The primary characteristic of hoot- ing habitat is an interspersion of open areas with taller woody cover (Weber 1975). Blue grouse broods selected areas with rel- atively high herbaceous cover. Within the maple vegetation type, mean height of herba- ceous vegetation at 12 brood locations was 50.8 cm (SE = 4.0), which was higher than that of 35 other summer observations in maple @=83,0cm, Sh = o.45 b= Q1G, chi 25), lin the open conifer vegetation type, mean herbaceous vegetation height at nine brood locations was 63.3 cm (SE = 14.8), whereas that for 19 other summer observations in open conifer was 31.0 cm (SE = 5.0, t = 2.61, df = 26). Sample sizes of broods in sagebrush (n = 4) and mixed shrub (n = 3) were not adequate for testing. Mussehl (1963) felt that herba- ceous cover at least 50 cm tall, interspersed with bare ground to provide travel lanes, was the most important aspect of good blue grouse brood cover. Additionally, clumps of small trees and shrubs may enhance brood habitat by providing nesting sites and protection from predators (Mussehl 1960, 1963, Weber 1975) and may be particularly important in late sum- mer when herbaceous cover becomes dessi- cated or is heavily grazed (Zwickel 1973). These results indicate that a variety of vege- tation types can be managed as blue grouse breeding habitat. No major differences be- tween seasons within each type implies that maintaining habitat characteristics for each GREAT BASIN NATURALIST Vol. 46, No. 1 | type within the levels reported in Table 2 should provide adequate conditions for | breeding and brood rearing. Except for tall _ herbaceous cover for broods, different charac- | teristics need not be provided for different | stages of the breeding season. Although we — did find some blue grouse breeding at high | elevation, these areas probably are not as im- | portant for breeding as low elevation open | habitats (Stauffer and Peterson 1985). | CHARACTERISTICS OF CONIFEROUS HABITATS Blue grouse selected sites in dense conifer | stands during fall and winter with about 65%-—69% coniferous tree cover (Table 3). Since | mean percent tree canopy cover at blue grouse locations in autumn was 45%, blue grouse se- lected the more open areas within dense conifer. | Although tree density in dense conifer was simi-_ lar at autumn and winter locations, significantly | (t = 2.15, df = 27) more Douglas- re foun at winter locations of blue grouse. Blue grouse selected open conifer stands hall | had approximately a 50:50 conifer cover to open | ratio. Caswell (1954) found that blue grouse se- q lected open conifer slopes in winter with islands. of subalpine and Douglas-fir. Percent tree canopy cover was relatively low in all seasons, _ averaging 32%—44% (Table 3). Density of small stems at blue grouse locations _ in open conifer was lower for all seasons com- | pared to those of other vegetation types (Tables 2 ~ and 3). Densities of subalpine fir at grouse loca- | tions did not vary significantly among seasons. but Douglas-fir densities were higher (t =5.84.. df = 115) at winter locations than those for au: tumn (Table 3). In winter, this species is used for. food and as roost sites (Marshall 1946). Winter Roost Trees 1} We compared trees used as winter roost: and for feeding with those not used. Of 3& roost trees, 36 (95%) were Douglas-fir and one) each (2.5%) were subalpine fir and Engel: mann spruce (Picea engelmannii). Of 34. conifers recorded along randomly locatec. transects in three wintering areas, 57% were | Douglas-fir, 33% were subalpine fir, 5% were | limber pine, 2% were Engelmann spruce, anc | 3% were lodgepole pine (Pinus contorta) | Thus, Douglas-fir were preferred as roos) | tmees: | . waren as ‘sae January 1986 STAUFFER, PETERSON: BLUE GROUSE ECOLOGY 2h TABLE 3. Mean vegetation characteristics recorded for 0.01 ha circular plots at blue grouse locations in the dense and open conifer vegetation types in southeastern Idaho, 1979-1981. Dense conifer Open conifer Fall Winter Spring Summer Autumn Winter Variable n=19 = 10 p= n = 28 = 7 n= 46 Coniferous cover (%) 68.7(2.7)* —-65.5(2.7) 45.0(2.9) 48.9(2.2) 47.7(1.8) . 2(1.8) Deciduous cover (%) TRERGES) 1.7(1.0) 1.1(1.1) 3.0(1.3) 2.5(0.5) 4.8(1.3) Open (%) 28.2.8) 33.6(3.1) —_-53.9(2.3) 48.0(2.1) 49.7(1.8) 50.9(2.0) Tree canopy cover (%) 45.1(7.1) — 33.7(9.6)(n=3)? 38.5(6.3) 31.6(4. 1)(n=65) = 5(8.5)(n=11) Ground cover (%) 28.4(5.2) — 14.4(8.3) 41.8(3.8) — 32.4(2.5) .8(2.2) Canopy height (m) 19.8(2.2) 22.8(1.2) —15.7(2.0) 13.8(1.4) 14.6(0.9) 8 8(1.1) Stems <7cmdbh/ha 1026(247) —391(157) 134(71) 495(163) — 1275(332) 291(77) Trees/ha 426(126) — 350(72) 256(67) 275(40) 294(40) 361(68) Deciduous trees/ha 16(11) 0 0 18(9) 17(6) 15(10) Subalpine fir/ha 195(72) 60(34) 33(33) 61(21) 140(33) 143(54) Douglas-fir/ha 63(25) 180(57) 189(65) 154(38) 110(25) 193(30) Lodgepole pine/ha 63(41) 50(40) 0 11(11) 7(4) 0 “Standard error. Where noted, sample size is smaller because the variable was not recorded for roost tree observations. TABLE 4. Mean vegetation characteristics at 38 roost tree sites used by blue grouse and nearby unused sites; southeastern Idaho, 1979-1981. Variable Used Coniferous cover (%) 50.3(1.9) Open (%) 45. 2(1.9) Canopy height (m) 21.5(0.9) Trees/ha 321(32) Trees 7-15 cm dbh/ha 100(24) Trees 15-33 cm dbh/ha 97(23) Trees > 33 cm dbh/ha 124(13) Subalpine fir/ha 68(21) Douglas-fir/ha 218(25) Limber pine/ha 24(10) Roost tree dbh (cm) 49.2(3.1) Mean (SE) Significance* Not used 44.5(2.1) p<0.01 50.2(1.9) p<0.01 20.5(1.0) p > 0.20 303(34) p > 0.20 129(27) p > 0.20 87(16) p > 0.20 87(10) p< 0.02 87(26) p > 0.20 179(22) p > 0.20 24(10) p > 0.20 42,8(2.4) p <0.05 “Represents the significance of a paired t-test for differences between used and unused trees for the variable. A paired t-test revealed significant differences for four variables measured at used and unused trees (Table 4.) Coniferous cover was greater and percent of area open was less at used trees. Used trees had a larger dbh and the density of large (> 33 cm dbh) trees was higher at used trees, as measured within circular plots (Table 4). Thus, within open conifer stands, blue grouse were selecting trees for roosting and feeding that were in denser clumps and were larger than trees not used. Roost trees often were in clumps rather than solitary. Although not investigated here, nutritional differences in the needles of different trees might influence roost and feeding tree se- lection, as has been found for spruce grouse (Dendragapus canadensis , Ellison 1976). Blue grouse appeared to prefer large Dou- « glas-fir that had dense foliage. The dense fo- | liage may provide protection from predators and weather, which often is harsh at high elevations in winter. Maintenance of open, high elevation stands of conifers, especially those containing Dou- glas-fir, should provide adequate winter habi- tat for blue grouse. Such stands have low com- mercial value; thus winter habitat of blue grouse probably is not threatened in south- eastern Idaho. LITERATURE CITED BENDELL, J. F., AND P. W. E.Liort. 1966. Habitat selec- tion in blue grouse. Condor 68:431—446. ___. 1967. Behavior and the regulation of numbers in blue grouse. Canadian Wildl. Ser. Rept. Ser. No. 4. BLACKFORD, J. L. 1958. Territoriality and breeding behav- ior of a population of blue grouse in Montana. Condor 60:145-158. 122 Boac, D. A. 1966. Population attributes of blue grouse in southwestern Alberta. Canadian J. Zool. 44:799-— 814. CASWELL, E. B. 1954. A preliminary study on the life history and ecology of the blue grouse in west central Idaho. Unpublished thesis, University of Idaho, Moscow. 105 pp. ELLISON, L. N. 1976. Winter food selection by Alaska spruce grouse. J. Wildl. Manage. 40:205—213. FowLe, C. D. 1960. A study of the blue grouse (Dendra- gapus obscurus Say) on Vancouver Island, British Columbia. Canadian J. Zool. 38:701—713. Harju, H. J. 1974. An analysis of some aspects of the ecology of dusky grouse. Unpublished disserta- tion, University of Wyoming, Laramie. 142 pp. HEEBNER, G. C. 1956. A study of the life history and ecology of the blue grouse in west central Idaho. Unpublished thesis, University of Idaho, Mos- cow. 51 pp. Lewis, R. A. 1981. Characteristics of persistent and tran- sient territorial sites of male blue grouse. J. Wildl. Manage. 45:1048-1051. Lewis, R. A., AND F. C. ZWICKEL. 1980. Removal and re- placement of male blue grouse on persistent and transient territorial sites. Canadian J. Zool. 58:1417-1423. Maestro, R. M. 1971. Spring and summer habitat prefer- ences of blue grouse on the Bear River Range, Utah. Unpublished thesis, Utah State University, Logan. 57 pp. GREAT BASIN NATURALIST Vol. 46, No. 1 MARSHALL, W. H. 1946. Cover preferences, seasonal movements, and food habits of Richardson’s grouse and ruffed grouse in southern Idaho. Wilson Bull. 58:42—52. MAaRrTINKA, R. R. 1972. Structural characteristics of blue grouse territories in southwestern Montana. J. Wildl. Manage. 36:498—510. MUSSEHL, T. W. 1960. Blue grouse production, move- ments, and populations in the Bridger Mountains, Montana. J. Wildl. Manage. 24:60-68. ____.. 1963. Blue grouse brood cover selection and land- use implications. J. Wild]. Manage. 27:547—555. STAUFFER, D. F. 1983. Seasonal habitat relationships of ruffed and blue grouse in southeastern Idaho. Un- published dissertation, University of Idaho, Moscow. 108 pp. STAUFFER, D. F., AND S. R. PETERSON. 1985. Ruffed and blue grouse habitat use in southeastern Idaho. J. Wildl. Manage. 49:459—466. WEBER, D. A. 1975. Blue grouse ecology, habitat require- ments, and response to habitat manipulation in north central Utah. Special Rept. No. 33. Utah Coop. Wildl. Res. Unit. ZWICKEL, F. C. 1973. Dispersion of female blue grouse during the brood season. Condor 75:114—119. ZWICKEL, F. C., AND J. F. BENDELL. 1967. Early mortality and the regulation of numbers in blue grouse. Canadian J. Zool. 45:817—851. ZWICKEL, F. C., I. O. Buss, AND J. H. BRIGHAM. 1968. Au- tumn movements of blue grouse and their rele- vance to populations and management. J. Wildl. Manage. 32:456—468. _ FALL DIET OF BLUE GROUSE IN OREGON! John A. Crawford,” Walt Van Dyke,’, S. Mark Meyers,” and Thomas F. Haensly” ABSTRACT. —The early fall diet of Oregon blue grouse (Dendragapus obscurus pallidus) from Wallowa County, ( Oregon, was determined from 145 crops obtained during 1981 and 1982. Of more than 50 plant and animal foods in the ‘diet, short-horned grasshoppers (Acrididae), prickly lettuce (Lactuca serriola), yellow salsify (Tragopogon dubius), \ wild buckwheat (Eriogonum spp.), and snowberry (Symphoricarpos albus ) occurred in 30% or more of the crops and ‘ collectively amounted to 68% of the diet by weight. Seven of the 12 most common foods were consumed differentially ' by the four sex and age classes of birds. Results indicated that blue grouse foraged in forest and grassland habitats. In 1981, a study was established to deter- ‘mine the early fall diet of the Oregon blue grouse in Wallowa County, Oregon. Our ob- _jectives were to compare the diet of grouse in this area with diets from other locations, espe- -cially within the range of this subspecies, and ‘to determine if dietary differences existed among the sex and age classes. Previous work indicated that western larch (Larix occidentalis) needles were one of the ‘most important foods from August through ‘October for blue grouse in northcentral ‘Washington and Idaho (Beer 1943, Boag /1963). Fir (Abies spp.) and Douglas- Se (Pseu- dotsuga menziesii) needles, staple winter ‘foods, also composed a major portion of the ‘fall diet (Beer 1943, Stewart 1944, Marshall | 1946, Boag 1963). Bearberries Ugciosteralin - los uva-ursi) likewise were noted as common foods by Beer (1943) and Boag (1963). Grasshoppers and ants composed most of the ‘animal foods in the fall diet of blue grouse (Beer 1943, Stewart 1944, Marshall 1946, Martin et al. 1951, Boag 1963, King and Ben- dell 1982). _ Few differences in the diets of adult and ‘immature blue grouse during late summer and fall have been noted. King and Bendell (1982) commented that from late July through September adults and juvenile blue grouse (D. o. fuliginosus) on Vancouver Island con- ‘sumed approximately the same types and amounts of foods. Beer (1943), in Idaho, re- ported that adults consumed more larch | Oregon Agricultural Experiment Station Publication 6978. needles than did immatures during August, but by September the diets were similar. A contrasting work by Boag (1963) revealed that greater amounts of larch needles were eaten by adults than by immatures during September and October, but no other differ- ences in the consumption of the major foods by the sex and age classes of blue grouse were found. STUDY AREA AND METHODS In Wallowa County blue grouse typically inhabit bunchgrass ridges, which are dis- sected by draws that are timbered on the north-facing aspect. Elevation ranges from 600 to 1500 m. Bunchgrass communities are dominated by bluebunch wheatgrass (Agro- pyron spicatum) and Idaho fescue (Festuca idahoensis ). Dominant forbs are wild buck- wheat (Eriogonum spp.) and arrow-leaf bal- samroot (Balsamorhiza sagittata). Timbered draws are dominated by ponderosa pine (Pi- nus ponderosa), Douglas-fir, true firs, and western larch. Shrub understories are com- posed of mallow ninebark (Physocarpus mal- vaceus), snowberry (Symphoricarpos albus), big huckleberry (Vaccinium membrana- ceum), creambush ocean-spray (Holodiscus discolor), currants (Ribes spp.), and shiny-leaf spiraea (Spiraea betulifolia). Idaho fescue and pinegrass (Calamagrostis rubescens) are the most common grasses in the understories. Grazing by domestic livestock occurs on most “Department of Fisheries and Wildlife, Oregon State University, Corvallis. Oregon 97331-3803. 3Oregon Department of Fish and Wildlife, Enterprise, Oregon 97828. 123 124 TABLE 1. Fall diet of blue grouse, Wallowa County, Oregon, 1981-1982. Food item PLANTS Prickly lettuce (Lactuca serriola) Yellow salsify (Tragopogon dubius ) Wild buckwheat (Eriogonum spp.) Snowberry (Symphoricarpos albus) Douglas-fir (Pseudotsuga menziesii ) Elkhorns clarkia (Clarkia pulchella) Unidentified Gramineae Western larch (Larix occidentalis ) Clover (Trifolium spp.) Everlasting (Antennaria spp.) Serviceberry (Amelanchier alnifolia ) Ponderosa pine (Pinus ponderosa) Dwarf mistletoe (Arceuthobium spp.) Smartweed (Polygonum spp.) Unidentified Compositae Lodgepole pine (Pinus contorta) Filaree (Erodium circutorium) Selaginella (Selaginella spp.) Hawthorne (Crataegus douglasii) Elderberry (Sambucus cerulea) Currant (Ribes spp.) Huckleberry (Vaccinium membranaceum) Unidentified Ericaceae Juniper (Juniperus occidentalis ) Yew (Taxus brevifolia) Engelmann spruce (Picea engelmannii) Fir (Abies spp.) Microseris (Microseris spp.) Dandelion (Taraxacum spp.) Sorrel (Rumex spp.) Bedstraw (Galium spp.) Strawberry (Fragaria virginiana) Unidentified Leguminosae Brome (Bromus spp.) Bentgrass (Agrostis spp.) Unidentified plant material ANIMALS Short-horned grasshoppers, Acrididae Ants, Formicidae Long-horned grasshoppers, Tettigoniidae Spittle bugs, Cercopidae Ground beetles, Carabidae Ladybird beetles, Coccinellidae Stink bugs, Pentatomidae Darkling beetles, Tenebrionidae Unidentified Insecta larvae Unidentified Insecta Chinch bugs, Lygaeidae Stink bugs, Scutelleridae Flies, Diptera Spiders, Araneida Sawflies, Tenthredinidae Yellow jackets, Vespidae Treehoppers, Membracidae Plant bugs, Miridae Stilt bugs, Berytidae Unidentified Hemiptera * Weight was obtained only for crops obtained during 1981. GREAT BASIN NATURALIST Parts consumed seed heads seed heads leaves berries needles, buds seed heads leaves, seeds needles leaves leaves berries seeds entire plant seeds seed heads seeds, needles seeds, leaves entire plant berries, stems berries berries berries berries berries needles needles needles seed heads seed heads leaves leaves, seeds berries seeds leaves stems leaves All items marked “tr” combined amounted to 1% of the diet by weight. Frequency (%) (n = 145) BE BE WWD WwWwWe KR OUUUO®D Nw & WA SorKD Re RRB BP RB eB Pe EP NOW Ww Pp Vol. 46, No. 1 Weight (%) (n = 83)" ) January 1986 sites. Many of the forest stands are managed for commercial timber production and some have extensive road systems. From 28 August through 29 September of 1981 and 1982, 145 crops containing food (39 adult males, 34 adult females, 39 immature males, and 33 immature females) were obtained ‘from hunter-killed blue grouse. The majority of crops (61%) were from birds taken between 28 _ August and 3 September, 33% came from the ‘second week, and the remainder from birds | killed from 11 to 29 September. The four sex age _classes were represented similarly through time; for example 59% of the crops from immature ) males and females and 63% from adult males and ‘females were collected during the first week. _ Ages of immatures in weeks was determined by «stage of molt sequence of primary feathers (Zwickel and Lance 1966) and ranged from 9 to 17 weeks (x = 13 weeks) (Redfield and Zwickel 1976). Contents of the crops were dried in an oven at 50 C for three days, identified and weighed. Contents of the 62 crops collected in 1982 were inadvertently destroyed before »weighing. A subjective evaluation of the relative amounts of foods in the 1982 sample revealed that they were essentially identical to the 1981 ssample. Frequencies of occurrence of the most -common foods were tested among the four sex and age classes with Chi-square analysis (Snedecor and Cochran 1967:20). Analysis of variance (Snedecor and Cochran 1967:258) was used to test for differences in weights of foods eaten by the sex and age classes. RESULTS AND DISCUSSION More than 50 plant and animal foods were consumed by blue grouse (Table 1). Of these, short-horned grasshoppers, prickly lettuce, yel- low salsify, wild buckwheat, and snowberry j were consumed at a frequency of 30% or more; collectively, these foods accounted for 68% of _the diet by weight. Five additional plant foods and two groups of insects were found in 10% to 29% of crops and contributed 11% of the diet by weight. Of the 12 most common foods, 7 were consumed differentially by the sex or age classes _ of blue grouse (Table 2). _ Short-horned grasshoppers (Acrididae), an important summer food of blue grouse (Stewart ~ 1944, Marshall 1946, Martin et al. 1951), were ‘found in 46% of the crops and contributed 32% CRAWFORD ET AL.: DIET OF BLUE GROUSE 125 by weight. The frequency of short-horned grasshoppers in the diet was higher (P < 0.001) for immatures (64%) than for adults (27%). Prickly lettuce was the second most frequent item in the fall diet and was con- sumed with similar frequency by all sex and age groups. Females, of both ages, consumed yellow salsify (P ~ 0.02) and wild buckwheat (P ~ 0.06) more commonly than did males. Previous work (Boag 1963) indicated that prickly lettuce, yellow salsify, and wild buck- oe were only minor components of the fall iee Snowberries were consumed equally by all sex and age classes and apparently are a more important fall food in Oregon than elsewhere within the range of the Oregon blue grouse (Beer 1943, Stewart 1944, Boag 1963). No differential use by the sex and age classes of blue grouse were found for ants, Douglas-fir needles, or unidentified grasses. Both ants and Douglas-fir needles were reported as common in the fall diet of blue grouse (Stew- art 1944, Boag 1963). Clarkia (Clarkia pulchella) and clover (Tri- folium spp.) were more frequently (P ~ 0.06) consumed by adults than by immatures. Clarkia was more common in the diet of blue grouse from Oregon (9% to 24% frequency) than from Washington (1% to.3% frequency) (Boag 1963), whereas the use of clover was similar. Clarkia was considered a food rejected by blue grouse on Vancouver Island (King and Bendell 1982). Long-horned grasshoppers (Tettigoniidae) were consumed more frequently (P ~ 0.05) by imma- tures than by adults. Needles of western larch were the most im- portant food in the diet of blue grouse during September and October in eastern Washington, where they were consumed with a frequency of 28% to 60% (Boag 1963). Beer (1943) found that larch needles composed 46.9% of the diet. by volume during August, but they dropped to 2.3% in September. In our study, larch needles were consumed in a moderate amount (10% fre- quency, 4% weight), but were used most com- monly by adult males (P < 0.001). Larch needles occurred in 28% of the crops from adult males and in only 3% to 6% of the other three groups. Boag (1963) found that adults utilized larch sig- nificantly more than did immatures, but he found no differences in use between adult males and adult females. GREAT BASIN NATURALIST Vol. 46, No. 1 TABLE 2. Frequency of the 12 most common foods by sex and age classes of blue grouse, Wallowa County, Oregon, 126 1981-1982. Adult male Food item (n = 39) Prickly lettuce* 43 Yellow salsify” 35 Wild buckwheat® 25 Snowberry* 35 Clarkia 20 Doulas fir’ 18 Unidentified Gramineae* 15 Western larch® 28 Clover‘ 15 Short-horned grasshoppers! 18 Ants* 25 Long-horned grasshoppers® 10 * No significant differences among sex and age classes. b Females > males, P ~ 0.02. © Females > males, P ~ 0.06. 4 Adults > immatures, P ~ 0.06. © Adults males > all other, P < 0.001. ! Immatures > adults, P < 0.001. ® Immatures > adults, P ~ 0.05. The average number of food items in a crop was 4.2. Average numbers of items ranged from 3.7 (adult females) to 4.6 (immature fe- males), but no difference (P > 0.25) existed in the number of items/crop among the sex and age groups. Our work revealed, in contrast to results of studies by Beer (1943), Stewart (1944), and Boag (1963), that most of the common early fall foods were consumed differentially by the sex and age groups of blue grouse. Greater consumption of short-horned and _long- horned grasshoppers likely reflected avail- ability of these groups during late August and September in Oregon and probably repre- sented a retention of the feeding patterns of immatures from summer. Stewart (1944) re- ported a fivefold greater consumption of ani- mal matter by immatures during summer. Likewise, the greater use of Elkhorns clarkia and clover by adults possibly represented a continuation of summer feeding habits. Beer (1943) found that adults ate more fruits, seeds, and green leafy material during summer than did immatures. By contrast, King and Bendell (1982) noted that the consumption of fruits, leaves, and flowers during late summer and early fall was similar between adult and imma- tures. Of the four foods with the greatest fre- quency of use, yellow salsify and wild buck- Frequency (%) ! Adult Immature Immature | female male female (n = 34) (n = 39) (n = 33) 38 45 47 50 28 53 35 28 AT 24 28 35 18 5 12 9 20 15 | 9 13 9 { 6 3 3 15 5 6 38 63 65 18 35 18 15 28 24 wheat were taken more commonly by females than males. The preponderance of prickly let- tuce and yellow salsify seed heads and wild | buckwheat leaves in the fall diet of blue’ grouse in Oregon was likely related to) availability. Adult males during August and September } are solitary and commonly at higher eleva- | tions than adult hens and immatures (Mar- | shall 1946, Boag 1963). In addition, our obser- — aise Ot heli: mse by adult males during | September indicated that they commonly are’ found in larch thickets, which may account for) the greater use of ledh meedlles by | | males. The results of previous work (Beer 1943, Boag 1963) indicated that Oregon ee grou foraged primarily in forested habitats during early fall, e.g., areas containing larch, Dou. | glas-fir, true firs, and bearberry. In contrast, : birds in this study made use ofa broad range 01) foods and, presumably, foraging habitats. The | two most important plant foods, prickly let: tuce and yellow salsify, are introduced “weed | species. Both are common in bunchgrass com: munities, especially where the soil has beer | disturbed by the burrowing activities of smal | mammals or by grazing and trampling by wile) and domestic ungulates. Clarkia is common ir disturbed areas (W. C. Krueger, persona communication). Wild buckwheat character. January 1986 istically is found in dry, open habitats (W. C. Krueger, personal communication). In addition, the importance of forested areas for foraging by blue grouse in Oregon is exemplified by the use of snowberry, western larch, and Douglas-fir. ACKNOWLEDGMENTS We wish to thank R. G. Anthony and R. L. Jarvis for their reviews of this manuscript. LITERATURE CITED BEER, J. 1943. Food habits of the blue grouse. J. Wildl. Manage. 7(1):32—44. Boac, D. A. 1963. Significance of location, year, sex, and age to the autumn diet of blue grouse. J. Wildl. Manage. 27(4):555—562. CRAWFORD ET AL.: DIET OF BLUE GROUSE 127 KING, R. D., AND J. F. BENDELL. 1982. Foods selected by blue grouse (Dendragaus obscurus fuliginosus). Canadian J. Zool. 60:3268—3281. MARSHALL, W. H. 1946. Cover preferences, seasonal movements, and food habits of Richardson’s grouse and ruffed grouse in southern Idaho. Wilson Bull. 58(1):42—52. MakrtTIn, A. C., H. S. ZIM, AND A. L. NELSON. 1951. Ameri- can wildlife and plants. McGraw-Hill Book Co., New York, NY. 500 pp. REDFIELD, J. A., AND F. C. ZwWICKEL. 1976. Determining the age of young blue grouse: a correction for bias. J. Wild]. Manage. 40(2):349-351. SNEDECOR, G. W., AND W. G. COCHRAN. 1967. Statistical methods. 6th ed. Iowa. State University Press, Ames, Iowa. 593 pp. STEWART, R. E. 1944. Food habits of blue grouse. Condor 46(3):112—120. ZWICKEL, F. C., AND A. N. LANCE. 1966. Determining the age of young blue grouse. J. Wildl. Manage. 30(4): 712-717. STATUS AND DISTRIBUTION OF CALIFORNIA GULL NESTING COLONIES IN WYOMING Scott L. Findholt! ABSTRACT. —Historically, two California Gull nesting colonies existed in Wyoming. In 1984 there were six breeding locations consisting of 10 different colonies that included approximately 7,273 nests. The increase in the California Gull nesting population in the state is probably a consequence of man-caused environmental changes that have resulted in the creation of additional breeding habitat and new food sources. Breeding populations of California Gulls (Larus californicus ) have increased in Wash- ington State and throughout much of the western United States since the 1920s (Conover et al. 1979, Conover 1983). Reasons for the proliferation of this species include the construction of large water impoundments with isolated islands for nesting, as well as the gulls’ exploitation of new man-made terres- trial food sources such as garbage dumps, other human refuse, and agricultural land (Conover 1983). Also, the California Gull has probably benefited from reduced human pre- dation by egg and plumage hunters. In this paper I present information on the distribution, population status, and habitat of recent California Gull nesting colonies in Wy- oming. I also provide details on the history of each colony and give reasons for the popula- tion increase of this species in the state. My purpose is to provide baseline data on each colony and to clarify the literature on the cur- rent number of California Gull nesting colonies in Wyoming. METHODS In 1983 and 1984 I conducted a survey for California Gull breeding locations to obtain recent information on the distribution of nest- ing colonies in Wyoming. Between 4 April and 31 May 1984 seven aerial searches in fixed-wing aircraft were made for nesting ar- eas of all colonial water birds, including Cali- fornia Gulls. Reservoirs, lakes, marshes or other potential breeding sites not observed during aerial flights were surveyed from the ground with binoculars or a spotting scope. Estimates of active nests were based on total ground counts except for one colony, where the number of nests was determined by a belt transect technique. All colonies were cen- sused in 1984 except for the Molly Islands colony, Yellowstone Lake, Yellowstone Na- tional Park; and Bamforth Lake colonies, Al- bany County. Colonies were censused when most birds were in the late incubation or early hatching stages and based on one visit to each colony. A colony was defined as a geographi- cally continuous group of breeding birds whose territorial boundaries are contiguous (Penney 1968). One exception was the Sand Mesa Wildlife Habitat Management Unit (WHMU), where several small breeding ag- gregations of California Gulls on man-made islands were treated as one colony. Historical sources of information on Califor- nia Gull nesting areas included a literature review, an examination of the files of the Wyo- | ming Game and Fish Department, and con- tact with biologists, naturalists, bird-watch- | ers, and others considered knowledgeable of | California Gull breeding locations in the state. COLONY DESCRIPTIONS AND HISTORIES Bamforth Lake McCreary (1939) considered the California Gull a common summer resident in Albany — County and indicated that a small colony had ~ existed since 1934 on an island in Bamforth Lake, 15 km northwest of Laramie (Fig. 1). In 1937 this nesting colony was about one-fourth | as large as the Molly Islands, Yellowstone | ‘Wyoming Game and Fish Department, 260 Buena Vista Drive, Lander, Wyoming 82520. 128 January 1986 FINDHOLT: CALIFORNIA GULL 129 —— ee ee ee eee CRBIONE C | \ SHERIDAN | | i NATIONAL _. ! ~s | " @ROOK | te jPARK OG PARK 4 Ne ee = I C | BIGHORN 1 | | | I > | ‘ ' CAMPBELL | oo | seca i l ; ' ne ah eee | JOHNSON | | ee aera a eae | | \taans | TETON ! \ HOT ae | | | | | Hite a ee ok Sata oe Es apogee [ra oe Seda ge L. oe 2 CL Ee RAN eR | ae, a a @ | | | 1 eal - \ ' | 1 | ' NIOBRARA | | << | NATRONA Con | SUB. ' sie aaa! eo ! ee Pe | hae ee | Pee hak eit 22 is ae ese | | | PLATTE | | i | t \ | LINCOLN | | | | GosEEN| ! : ' 1 | 1 | : \ SWEETWATER CARBON | i= a eee | ! Blign cs | oth In | LARAMIE ! ! { | Peirce tele teA oes aoa | sie swe nea : See weet ee ee 5 Fig. 1. Locations of California Gull nesting colonies in Wyoming, 1984. Lake, colony in 1932 (McCreary 1939). Total numbers of California Gull nests in the Bam- forth Lake colony ranged from a low of 1,364 in 1967 to 2,003 in 1968, with a mean of 1,842 between 1967 and 1972 (Kennedy 1973). In 1974 this colony contained 2,069 nests and increased to 2,470 nests in 1975 (Raper 1975). Approximately 5,000 gulls were estimated to be breeding at the Bamforth Lake colony in 1979 and 1980 (Pugesek 1983). In 1984 from 6,000 to 7,000 California Gulls appeared to be nesting at Bamforth Lake. In addition to breeding on Bamforth Island, the traditional nesting site, a portion of the California Gulls bred on a 0.4-ha natural island, about 0.5 km southwest of the main colony. Establishment of the new nesting colony in 1984 was proba- bly in response to reduced nesting habitat. On 14 July 1983 Bamforth Island was 1.2 ha, which was 45% smaller than it was in 1975 (Raper 1975). The nesting island continued to decline in size throughout 1984 because of rising water levels caused by above normal precipitation and increased irrigation runoff. By 1 August Bamforth Island was completely inundated except for the tops of black grease- woods (Sarcobatus vermiculatus ). Bamforth Island is primarily composed of bare ground interspersed with dense patches of black greasewood. The new nesting island, Peninsula Island, had more vegetative cover because the colony was recently established. Predominant plants included black grease- wood and prickly-pear cactus (Opuntia sp.) interspersed with grasses and forbs. Nesting associates at Bamforth Lake in 1984 included the Double-crested Cormorant (Phalacroco- rax auritus, 72 active nests), Snowy Egret (Egretta thula, 7 active nests), Black-crowned Night-Heron (Nycticorax nycticorax, 1 active nest), and Herring Gull (Larus argentatus, 3 active nests). Bamforth Lake is a naturally occurring lake, almost entirely in private ownership, except for small portions under jurisdiction of the U.S. Fish and Wildlife Service and set aside as 130 Bamforth National Wildlife Refuge. Because access is restricted, Bamforth Lake receives minimal use from the public. Pathfinder Reservoir The nesting colony at Pathfinder Reservoir, an irrigation impoundment on the North Platte River, is about 70 km northeast of Raw- lins, Carbon County (Fig. 1). Although Cali- fornia Gulls were first reported to be breeding here on 7 May 1982 by T. Varcelli, it is un- known when the colony was initiated. On 15 June 1984, I counted 683 California Gull nests. Birds were nesting on a 5.2-ha island, hereafter called Bird Island, about 0.8 km from the mainland and near the mouth of Sand Creek. The island is well vegetated, with sil- ver sagebrush (Artemisia cana), gray rabbit- brush (Chrysothamnus nauseosus ), sand dock (Rumex venosus ), plains cottonwood (Populus sargentii), and willow (Salix sp.) being most common. Other colonial waterbirds on Bird Island were the American White Pelican (Pelecanus erythrorhynchos, 245 active nests), Double-crested Cormorant, 126 active nests; Great Blue Heron (Ardea herodias , 38 active nests), and Caspian Tern (Sterna caspia, 15 to 20 active nests). Although portions of Pathfinder Reservoir receive considerable recreational use, presently human disturbance in the vicinity of Bird Island appears minimal. Ocean Lake WHMU Ocean Lake, about 24 km northwest of Riv- erton, Fremont County (Fig. 1), is a man- made lake that was formed in 1926 from irriga- tion return flow and seepage from Pilot Butte Reservoir (Serdiuk 1965). California Gulls started breeding here in small numbers dur- ing the early 1950s (W. Higby, personal com- munication). R. Klataske (1970) counted 550 nests on Third Point Island and 110 nesting pairs on Gull Island, and he estimated that about 2,000 California Gulls nested at Ocean Lake in 1970. The one colony that I discov- ered at Ocean Lake in 1981 was deliberately destroyed by humans. There were 609 Cali- fornia Gull nests at Ocean Lake on 14 May 1983. All gulls were breeding on Peninsula Island in the southeastern portion of the lake. This nesting colony increased to 775 nests on 19 May 1984. GREAT BASIN NATURALIST Vol. 46, No. 1 The 0.4-ha nesting island was almost void of vegetation except for sparse black greasewood on its southern portion and cattails (Typha latifolia) along the southwestern and eastern shorelines. The only nesting associate in 1984 was the Double-crested Cormorant (205 ac- tive nests). Ocean Lake is included in Ocean Lake WHMU and managed by the Wyoming Game and Fish Department. It is heavily used for _ recreation during the nesting season. Except — as noted in 1981, human disturbance of breed- ing California Gulls has not been a serious | problem because shallow water in the vicinity of the nesting island discourages boating ac- tivity. Sand Mesa WHMU | | Sand Mesa WHMU, managed by the Wyo- | ming Game and Fish Department, is approxi- | mately 32 km northeast of Riverton, Fremont | County (Fig. 1). It consists of several small reser- | voirs designed primarily for waterfowl produc- | tion. I counted 181 California Gull nests at Pond No. 1 on 15 May 1983. This was the first year that gulls bred here (K. Asay, personal communica- tion). California Gulls were nesting on six man- made islands that averaged about 0.013 ha. Is- lands were composed of cobble and were / generally void of vegetation. The nesting colony — contained 162 nests on 12 May 1984, which is 19. fewer nests than in 1983. This decline was most | | likely caused by the addition of more cobble, | making islands dome shaped and less suitable as. nesting substrate. Nesting associates in 1984 in- I, cluded the Double-crested Cormorant (21 active | nests) and Snowy Egret (from | to 2 active nests). | When I rechecked the colony 22 June 1984, all California Gull nests had been destroyed, most likely from human intervention. Soda Lake | Soda Lake, located 3 km north of Casper, Na- | trona County (Fig. 1) occurs naturally. How- ever, the lake has increased in size because ' waste water is added to it by the Amoco Oil, ' Refinery. California Gulls have nested here since the late 1950s (O. K. Scott, personal com- ‘ munication). In 1970 G. Dern estimated that approximately 400 breeding gulls were present. There were four California Gull nesting colonies at Soda Lake when I surveyed it in 1984. ! January 1986 MANMADE ISLAND.—This 1.3-ha island on the southeast portion of Soda Lake was devel- oped after the 1983 nesting season. Thus, 1984 was the first year that California Gulls bred here. On 22 May 1984, I estimated that 1,907 +204 (SE) active California Gull nests were present. Nesting associates included the Double-crested Cormorant (58 active nests), Snowy Egret (1 active nest), and Ring-billed Gull (Larus delawarensis, 70 active nests). There was dense vegetative cover on this is- land. Predominant species were silver sage- brush and cheat grass (Bromus tectorum). Other common plants included green rabbit- brush (Chrysothamnus _ viscidiflorus), prickly-pear cactus (Opuntia polyacantha), and other grasses. RATTLESNAKE ISLAND. — This 0.2-ha natural is- land near the east end of Soda Lake was first utilized for nesting by California Gulls in 1984. On 21 May 1984 I counted 560 active gull nests. One pair of Caspian Terns also bred here. Rat- tlesnake Island was covered with grasses inter- spersed with black greasewood and prickly-pear cactus when censused for gulls. East IsLaND.—On 22 May 1984 I found 24 active California Gull nests on this 0.08-ha natu- ral island. Nesting associates included the Dou- ble-crested Cormorant (10 active nests) and Black-crowned Night-Heron (1 active nest). This island.is disappearing because of high water lev- els at Soda Lake. It is void of vegetation except for sparse cheat grass, other grasses, black greasewood, and prickly-pear cactus. WEST ISLAND. —I counted 93 active California Gull nests on this 0.12-ha natural island 22 May 1984. Other colonial waterbirds nesting here were the Double-crested Cormorant (290 active nests), Snowy Egret (2 active nests), and Black- _ crowned Night-Heron (2 active nests). West Is- land is mostly bare except for scattered black greasewood and an unidentified forb. Like East Island, this island has decreased in size because of high lake water levels. California Gulls nesting at Soda Lake are se- cure from human intervention because this area is “off limits” to the public. Yellowstone Lake ___Linton (1891) indicated there were many + adult California Gulls present at Yellowstone | Lake, Yellowstone National Park, 10 July FINDHOLT: CALIFORNIA GULL 131 1890 but did not provide substantial evidence of nesting. Breeding was first reported by Skinner (1917), who estimated that 1,000 gulls were present in Yellowstone National Park and practically all nested on the Molly Is- lands, Yellowstone Lake, in 1898. The Cali- fornia Gull breeding population appeared to have varied considerably between 1917 and 1966 (Diem and Condon 1967). On 6 July 1962 Schaller (1964) estimated there were 600 ac- tive nests. Recently, numbers of California Gulls breeding on the Molly Islands have re- mained relatively unchanged since Schaller’s census (K. L. Diem, personal communica- tion). The Molly Islands have been previously described (Schaller 1964, Diem and Condon 1967, Diem 1979). The two Molly Islands, Rocky Island and Sandy Island, are about 0.4 km apart and 0.8 km from the southern shore of the Southeast Arm of Yellowstone Lake (Fig. 1). The combined surface area of the islands varies between 0.3 ha and 0.5 ha de- pending on water levels. Both islands are sparsely vegetated. Nesting associates in 1981 included the American White Pelican (290 active nests), Double-crested Cormorant (17 active nests), and Caspian Tern (14 active nests) (K. L. Diem, personal communication). Because of National Park Service concern for the welfare of white pelicans and other colonial waterbirds breeding on the Molly Is- lands, scientific investigations have been re- stricted and the nesting colonies are closed to the public. DISCUSSION According to Conover (1983), two Califor- nia Gull nesting colonies existed in Wyoming in 1980; one was at Bamforth Lake, Albany County, and the other at Yellowstone Lake, Yellowstone National Park. Nesting colonies were also present at Ocean and Soda lakes but were overlooked when Conover conducted his survey. In addition, several new California Gull colonies have become established else- where in Wyoming in recent years (Table 1). Table 1 indicates the occurrence of approxi- mately 7,273 nests in the state. The construction of large reservoirs with isolated islands has created substantial nest- ing habitat for California Gulls in Wyoming. 132 GREAT BASIN NATURALIST TABLE 1. Location, number of nests, year of establishment, and habitat of California Gull colonies in Wyoming, 1984. Name Location ALBANY COUNTY Bamforth Lake 1. Bamforth Island 2. Peninsula Island CARBON COUNTY Pathfinder Reservoir 3. Bird Island FREMONT COUNTY Ocean Lake WHMU 4. Peninsula Island Sand Mesa WHMU 5. Pond No. 1 NATRONA COUNTY Soda Lake 6. Man-made Island 7. Rattlesnake Island 8. East Island 9. West Island YELLOWSTONE NATIONAL PARK Yellowstone Lake 10. Molly Islands 44°19'N,110°16’W “Population estimate is from 1975 (Raper 1975). Mean + SE. “Population estimate is from 1962 (Schaller 1964). 41°24'N, 105°44’W 41°24'N, 105°44’W 42°23'N, 106°56’W 43°07'N, 108°35’W 43°19'N, 108°20’W 42°54'N, 106°18’W 42°54'N, 106°18’W 42°54'N, 106°18"W 42°54'N, 106°19’W Seven of eight California Gull nesting colonies (87%) that have become established since the 1960s are on man-made water impound- ments, and two of these colonies are on man- made islands. Although quantitative data are lacking on the diet of California Gulls in most colonies, food resources appear to have been influ- enced by man’s activities in Wyoming. Ex- panded agriculture may have provided con- siderable food for California Gulls, especially in colonies at Ocean Lake and Sand Mesa WHMUs. Gulls from these colonies com- monly forage in the extensive cultivated and irrigated fields nearby. Although gulls at Ocean Lake and Sand Mesa WHMUs also exploit refuse left by fishermen and other recreational users, minimal sign of garbage occurs in both colonies. In contrast, California Gulls at Soda Lake seem to rely heavily on food resources available at the nearby Casper garbage dump. Gulls nesting at Bamiorth Lake, Pathfinder Reservoir, and Yellowstone Lake appear to have a more natural diet. Pugesek (1983) found that most California Gulls from Bamforth Lake foraged at freshwa- ter sources within 20 km of the colony, and he Vol. 46, No. 1 Year colony Number of established nests Habitat 1934 2, 470° Lake 1984 — Before 1982 683 Reservoir 1983 775 Reservoir 1983 162 Reservoir 1984 1,906 + 204° Reservoir 1984 560 Early 1960s 24 Early 1960s 93 Before 1898 600° Lake observed that the diet of offspring mainly con-_ sisted of aquatic insects, salamanders, and — fish. Although California Gulls from colonies — in Yellowstone National Park and Pathfinder — Reservoir consume refuse left by fishermen — and other recreational users, their diet proba- — bly consists mostly of natural food resources | since man-caused environmental changes, which would create food, have not taken place — in either area. Conover et al. (1979) indicated that de-— creased human predation on California Gulls — and their eggs was partially responsible for the | population increase in Washington State. In Wyoming, California Gulls have continued to. increase despite continuous human persecu- tion in the state. ACKNOWLEDGMENTS I thank Larry Pate and Lynn Jahnke for. q assistance with censusing gull colonies in the ~ Casper area. Kenneth L. Diem provided re- | | cent information on the Yellowstone Lake colony. Nancy Findholt, Joseph Jehl, and Bob. Oakleaf made many helpful suggestions on the manuscript. All information gathered on January 1986 California Gull colonies was made possible through funding by the Wyoming Game and Fish Department, Nongame Project. LITERATURE CITED CONOVER, M. R. 1983. Recent changes in Ring-billed and California Gull populations in the western United States. Wilson Bull. 95:362—383. Conover, M. R., B. C. THOMPSON, R. E. FITZNER, AND D. E. MILLER. 1979. Increasing populations of Ring- billed and California Gulls in Washington State. Western Birds 10:31—36. Diem, K. L. 1979. White Pelican reproductive failures in the Molly Islands breeding colony in Yellowstone National Park. Proc. Research in National Parks Symposium. U.S. National Park Service Transac- tions and Proceedings. Sec. No. 5:489—496. Diem, K. L., AND D. D. CONDON. 1967. Banding studies of waterbirds on the Molly Islands, Yellowstone Lake, Wyoming. Yellowstone Library and Mu- seum Assoc., Yellowstone National Park, Wyo. 41 pp. KENNEDY, J. R. 1973. A study of a breeding colony of California Gulls (Larus californicus), Bamforth Lake, Albany County, Wyoming. Unpublished thesis, University of Wyoming, Laramie. FINDHOLT: CALIFORNIA GULL 133 KLaTAsKE, R. 1970. Island-nesting waterbirds. Wyoming Wildl. 34(6):22—927. Linton, E. 1891. A contribution to the life history of Dibothrium cordiceps Leidy, a parasite infesting the trout of Yellowstone Lake. Bull. U. S. Fish Comm. 9 (1889):337—358. McCreary, O. 1939. Wyoming bird life. Burgess Publ. Co., Minneapolis, Minnesota. PENNEY, R. L. 1968. Territorial and social behavior in the Adelie Penguin. Pages 83-131 in O. L. Austin, ed., Antarctic bird studies. Amer. Geophys. Union Publ. 1686. Washington, D.C. PUGESEK, B. H. 1983. The relationship between parental age and reproductive effort in the California Gull (Larus californicus). Behav. Ecol. Socio-biol. 13:161-171. RaPER, E. L. 1975. Influence of the nesting habitat on the breeding success of California Gulls (Larus cali- fornicus), Bamforth Lake, Albany County, Wyo- ming. Unpublished thesis, University of Wyo- ming, Laramie. SCHALLER, G. B. 1964. Breeding behavior of the white pelican at Yellowstone Lake, Wyoming. Condor 66:3-23. SERDIUK, L. J. 1965. An evaluation of waterfowl habitat at Ocean Lake, Wyoming. Unpublished thesis, Uni- versity of Wyoming, Laramie. SKINNER, M. P. 1917. The birds of Molly Islands, Yellow- stone National Park. Condor 19:177—182. NEW GENUS AND SPECIES OF LEAFHOPPER IN THE TRIBE TINOBREGMINI (HOMOPTERA: CICADELLIDAE: COELIDIINAE) M. W. Nielson! ABSTRACT— A new genus, Stenolidia, and new species, S. magna (type species), are described and illustrated. Stenolidia from Guyana is the fifth genus in the tribe Tinobregmini and the third that has come to light since the tribe was revised in 1975. Since the revision of the tribe Tinobregmini (Nielson 1975), two new genera, Tantulidia Nielson and Corilidia Nielson (Nielson 1979, 1982), have been described and assigned to the tribe. At this time I describe a new genus, Stenolidia , from Guyana and add a fifth mem- ber to the tribe that further broadens the tribal concept and distribution of the group. The tribe Tinobregmini, sensu stricto, originally included only Tinobregmus Van Duzee from the southern United States. De- Long (1945) described additional species of Tinobregmus from Mexico; then he later (1969) described Chilelana from Chile and assigned it to the Tinobregmini. Narrow crowns, brachytery, and pronotal conceal- ment of forewing bases were the primary characters that united these members of the tribe. The recent addition of Tantulidia and Co- rilidia not only broadened the tribal concept but also coalesced the distributional gap be- tween Tinobregmus and Chilelana. Stenolidia is placed in the tribe Tinobregmini with some reservations. Its external features are nearest to Tantulidia, but its aedeagal characters are closest to Corilidia. Therefore, it seems more appropriate to place Stenolidia in Tinobreg- mini than to erect a new tribe or relegate it to the tribe Coelidiini, to which it clearly does not belong. Stenolidia keys nearest to Tan- tulidia in my 1982 paper. In addition to the aforementioned charac- ters, the normal pronotum, scutellum, and forewings (not reduced) are now adjunct char- acters for the tribe. Except in Corilidia and Stenolidia, the aedeagal characters in the re- maining described genera are quite diverse. Stenolidia, n. gen. Type-species: Stenolidia magna, n. sp. | Medium-sized, slender species. Similar to — Tantulidia Nielson in general habitus but with — distinctive male genital characters. Color — black except for eyes and face. Head much narrower than pronotum; — crown elevated, very narrow and produced distally; eyes very large, elongate-ovoid; — pronotum large, scutellum moderately large; forewing elongate, 5 apical and 3 anteapical _ cells present, outer one closed; hind wing well developed; clypeus long, narrow, without — median longitudinal carina; clypellus long, © broad at distal 1/3. MALE. Genitalia partially asymmetrical; pygofer broad with short caudodorsal process; aedeagus asymmetrical, long with dorsal pro- | cesses near middle of shaft; connective Y-— shaped, stout; style moderately long; plate | | long, narrow with subapical constriction. | Stenolidia is known only by a single species | from Guyana. It is nearest to Tantulidia in general habitus by not having the bases of the forewings concealed by the posterior margin of the pronotum and to Corilidia Nielson in ~ certain aedeagal characters. The genus can be. readily separated from these genera by the | long, narrow elevated crown with parallel lat- eral margins and by the subapical constricted plate. | Stenolidia magna, n. sp. Length: male 7.10 mm. , General color black except for light reddish| brown eyes and deep tannish face, dark ochra-| ceous areas in costa of forewings. | | ‘Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah 84602. } | 134 | January 1986 NIELSON: NEW CICADELLIDAE 8 Figs. 1-8. Stenolidia magna, n.sp.: 1, Head and pronotum, dorsal view. 2, Face. 3, Male pygofer, lateral view. 4, | Aedeagus, lateral view. 5, Connective, aedeagus and right style, dorsal view. 6, Aedeagus distal enlargement, dorsal _ view. 7, Style, lateral view. 8, Plate, ventral view. Head much narrower than pronotum (Fig. | 1); crown very narrow, distinctly produced distally beyond anterior margin of eyes, width nearly 1/2 width of eyes, prominently ele- _ vated above level of eyes, lateral margins par- allel, slightly carinate; eyes very large, elon- gate-ovoid; pronotum large (atypical of tribe), length nearly equal to length of crown; scutel- lum moderately long; forewing elongate, ap- pendix well developed; clypeus long and nar- row, narrower anteriorly than posteriorly, surface finely rugose anteriorly; clypellus long, lateral margins broad at distal 1/3 (Fig. 2). MALE. Pygofer broad with rectangular ovate lobe on caudodorsal margin (Fig. 3); aedeagus asymmetrical, long, broad at basal 2/3, with longitudinal transparent area along ‘middle in lateral view (Fig. 4), compressed idorso-ventrally at distal 1/3 with apex nar- «towed and curved dorsally in lateral view, two |stout spines arising from dorsal surface of shaft just distad of middle, the dorsal one slightly un- dulated with a secondary, short lateral process, ventral one single and slightly curved (Fig. 5, 6); gonopore subapical, opening near dorso-lateral margin of shaft; connective stout, Y-shape, with short stem; style short, broad basally and nar- rowed at distal half (Fig. 7); plate long and nar- row, distinctive constriction subapically, with few microsetae apically (Fig. 8). FEMALE. Unknown. Holotype (male), GUYANA: New River, 750 ft, 1-5.V.1938, C. A. Hudson (BMNH). REMARKS: Stenolidia magna is the only known representative of the genus and can be separated from species of other genera in the tribe by char- acters described above. It is the largest in size among all known species in the tribe. ACKNOWLEDGMENTS I express my sincere appreciation to Dr. W. J. Knight, British Museum (Natural History) 136 GREAT BASIN NATURALIST Vol. 46, No. 1 (BMNH), for loan of this specimen, to Jean Stanger for the excellent illustrations, and to Dr. James P. Kramer, U.S. National Mu- seum, VVashington, D.C., and Dr. Paul W. Oman, Oregon State University, Corvallis, for reading the manuscript. LITERATURE CITED DELonG, D. M. 1945, The genus Tinobregmus (Ho- moptera: Cicadellidae) in Mexico. Bull. Brooklyn Ent. Soc. 40:97-102, pl. 2. _____. 1969. New South American genera related to Tino- bregmus and Sandersellus. J. Kansas Ent. Soc. 49:469-466, figs. 1-5. NIELSON, M. W. 1975. A revision of the subfamily Coelidiinae (Homoptera: Cicadellidae). Tribes Tinobregmini, Sandersellini, and Tharrini. Bull. British Mus. Nat. Hist. Suppl. 24. 197 pp. 514 figs. . 1979. Anew genus, Tantulidia, in the tribe Tinobreg- mini with a review of the species and generic limita- tions of the tribe (Homoptera: Cicadellidae). J. Kansas Entomol. Soc. 52:653-661. . 1982. A new genus Corilidia and a new species of the | Tribe Trinobregmini with a revised key to the Genera (Homoptera: Cicadellidae: Coelidiinae). J. Kansas Entomol. Soc. 55:423-426. | ( NEW ORIENTAL GENUS OF LEAFHOPPERS IN THE TRIBE COELIDIINI WITH DESCRIPTIONS OF NEW SPECIES (HOMOPTERA: CICADELLIDAE: COELIDIINAE) M. W. Nielson! ABSTRACT—A new genus, Stylolidia, and two new species, Stylotidia pectinata n. sp. (type species) and Stylolidia cristata n. sp., from Malaysia are described and illustrated. A key to the males of the known species is also included. The Oriental leafhoppers of the tribe Coelidiini are composed of five rather distinct genera, three of which are fairly large and have speciated over the entire region (Nielson 1982). Two genera, Mahellus Nielson and Jenolidia Nielson, are small and geographi- cally restricted. The former occupies the western edge of the Oriental region, whereas the latter is known only from Borneo. The new genus Stylolidia is described from two new species, S. pectinata (type-species) and S. cristata, from Malaysia. Both species have unusual characters, i.e., tubular aedeagi without processes and styles with prominent spines, which distinguish the genus from other Oriental genera in the tribe. The styles of S. cristata are similar to Lodiana unica Nielson, but in the former each style has two rows of uniseriate spines whereas the latter has only one row. Key to males of Stylolidia 1. Style with long spines arising from apex ........ 05 Bab Dua te tag Bech Gas Da ae eee oie es ee ane pectinata, n. sp. Style with two rows of short spines arising from inner lateral margin of apical half.... cristata, n. sp. Stylolidia n. gen. Type-species: Stylolidia pectinata, n. sp. Medium-sized species. Length male 8.00-8.80 mm. Similar to Lodiana Nielson in _ general habitus and to Taharana Nielson in aedeagal characters. Color deep brown to _ blackish with mottled markings on forewings. Head narrower than pronotum, subconical; crown narrow, width less than width of eye, produced slightly beyond anterior margin of eyes, nearly rounded at apex, lateral margins converging basally; pronotum and scutellum large; forewing with 5 apical cells and 3 an- teapical cells, outer one closed; hind wing well developed; clypeus narrow, without me- dian longitudinal carina; clypellus narrow at basal 2/3, expanded at apical 1/3. MALE. Genitalia partially asymmetrical. Pygofer with or without caudodorsal process, caudoventral process absent; 10th segment long and narrow; aedeagus tubular, without processes (similar to Taharana); dorsal apodeme long and narrow; connective broadly Y-shaped; style long with distinctive spines; plate long, narrow with few microse- tae on surface. The genus Stylolidia is known only from Malaysia and is represented by the two spe- cies described below. It can be distinguished from Lodiana by the absence of processes on the aedeagus and from Taharana by the long style with spines on its distal half. Stylolidia pectinata, n. sp. Length: Male 8.00 mm. Moderate-sized, slender species. General color dark brown with tannish mottled mark- ings on forewing. Head narrower than pronotum (Fig. 1); crown narrower than width of eye, slightly produced distally, slightly elevated above level of eyes; ocelli near anterior margin of crown; eyes large, semiglobular; pronotum, scutellum, and forewing as in description of genus; clypeus long, narrow, lateral margins broadly convex, flat medially (Fig. 2); clypel- lus as in description of genus. ‘Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah 84602. 137 138 GREAT BASIN NATURALIST Vol. 46, No. 1 Figs. 1-7. Stylolidia pectinata, n. sp.: 1, Head and pronotum, dorsal view. 2, Face. 3, Male pygofer, lateral view. 4, | Connective, aedeagus, and right style, dorsal view. 5, Aedeagus, lateral view. 6, Style, lateral view. 7, Plate, ventral | view. MALE. Pygofer in lateral view with hooked caudodorsal process (Fig. 3); aedeagus long, narrow, tubular, slightly curved in lateral view and extending beyond apex of style, without processes (Figs. 4, 5); gonopore dis- tad of middle of shaft on lateroventral margin; style short, not reaching apex of aedeagal shaft, with 4-5 very long nearly straight spines arising from apex, spines nearly as long as basal half of style (Figs. 4, 6); plate long and narrow (Fig. 7). FEMALE. Unknown. Holotype (male), MALAYSIA: Malay Pe- nins., Perak, 15-XI-1927, G. Kledang (BMNH). REMARKS: S. pectinata is easily separated from S. cristata by the hooked caudodorsal process on the pygofer and by the long spines that arise from the apex of the style. Additional data on the label attached to the holotype specimen include abbreviations “F.M.S.” and “2650” on the top side and “presented by Mr. E. S. Kimund, Sel: Mus:” — on the underside. Stylolidia cristata, n. sp. Length: Male 8.80 mm. Medium-sized, slender species. General | color blackish with numerous mottled tannish | markings on forewing. Head as in pectinata n. sp. but with larger | semiglobular eyes (Fig. 8); pronotum, scutel- | lum and forewing as in description of genus; — clypeus and clypellus as in pectinata (Fig. 9). | MALE. Pygofer in lateral view with small, | translucent, scalelike process on caudodorsal margin (Fig. 10); aedeagus long and tubular, without processes (Figs. 11, 12); gonopore as. in pectinata; style very long, extending dis- tally beyond apex of aedeagal shaft, with two | rows of uniseriate stout spines on distal half, one row on dorsal margin and the other on the | inner lateral margin (Figs. 11, 13); plate as in’ pectinata (Fig. 14). ] January 1986 NIELSON: NEW CICADELLIDAE 139 14 Figs. 8-14. Stylolidia cristata, n. sp.: 8, Head and pronotum, dorsal view. 9, Face. 10, Pygofer, lateral view. 11, FEMALE. Unknown. Holotype (male) MALAYSIA: Pahang Bukit | Ibam, 90 km WNW of Kuala Rompin, ca 50m 5-9. X. 1961, at night, K. J. Kuncheria (BPBM). REMARKS: This species can be _ distin- guished from S. pectinata by the long style that exceeds the length of aedeagus and by the two uniseriate rows of spines on the distal half of style. ACKNOWLEDGMENTS Dr. W. J. Knight, British Museum (Natural History), London (BMNH), and the late Dr. _ Connective, aedeagus, and right style, dorsal view. 12, Aedeagus, lateral view. 13, Style, lateral view. 14, Plate, | ventral view. J. Linsley Gressitt, Bernice P. Bishop Mu- seum, Honolulu (BPBM), graciously loaned the specimens on which this paper was based. Jean Stanger provided the illustrations and Dr. James P. Kramer, U. S. National Mu- seum, Washington, D.C., and Dr. Paul W. Oman, Oregon State University, Corvallis, gave of their time to review and improve the content of the paper. LITERATURE CITED NIELSON, M. W. 1982. A revision of the subfamily Coelidiinae (Homoptera: Cicadellidae). IV. Tribe Coelidiini. Pacific Insects Monogr. 38. 318 pp., 1,104 figs. ECOLOGICAL DIFFERENCES OF C, AND C, PLANT SPECIES FROM | CENTRAL UTAH IN HABITATS AND MINERAL COMPOSITION | C. Morden’, Jack D. Brotherson’, and Bruce N. Smith? ABSTRACT. —Six study sites were established in each of three community life form types (grass, forb, and shrub) | containing as dominants or subdominants either C, and/or C, plants. Soil and vegetation samples were analyzed for | total nitrogen, phosphorus, magnesium, calcium, potassium, sodium, zinc, iron, copper, and manganese. Discrimi- _ nant analysis and analysis of variance statistics were used to evaluate differences in mineral content of soils and plant tissues. C, plants in all study sites assimilated higher concentrations of potassium, iron, and calcium than did C; plants. Forbs in all sites contained the highest concentrations of minerals, followed by shrubs and grasses. if Studies have shown that 40%—50% of the soluble leaf protein in C; species consists of RUBPcase, whereas in C, species only about 5%-20% was RUBPcase (Blenkinsop and Dale 1974). Based on this evidence, Brown (1978) suggested that C, species should re- quire less nitrogen than do C; species. Several studies have supported this hypothesis (Christie 1979, Hallock et al. 1965, Wilson and Haydock 1971, Wilson 1975). It has also been shown that C, species require small amounts of sodium for growth (Brownell and Crossland 1972), although these studies were done on species specifically adapted to differ- ent environments (saline vs. nonsaline). It was the purpose of this study to investigate the mineral relationships of C; and C, plant spe- cies that grow in natural communities of com- parable environmental condition to assess whether ecological differences in mineral up- take do exist. MATERIALS AND METHODS Study Area Thirty-six study sites were established in plant communities bordering Utah Lake, Utah County, Utah, at approximately 40°10’ N, 11°50’ W (Fig. 1). Elevations ranged from 1,368 to 1,408 m above sea level, with a mean of 1,377 m. Six study sites were established in each of six community types. Communities were selected because of the presence of the species Sporobolus airoides, Puccinellia nut- @P on | @sP Ge | eK | UTAH LAKE 3 Provo Bay QD ‘ | @ Salt Lake City | Utah Lake [Provo @A HI Utah } | | | of | @A @SH | I @A ek i @SH | sce esP @SH esp | t I Fig. 1. A map of Utah Lake showing the locations ol) \ study sites around the lake. Communities shown corre- | f spond to P = Puccinellia, Sp = Sporobolus, A = Atriplex. \ K = Kochia, G = Greasewood and Sh = Shadscale. talliana, Atriplex patula, Kochia scoparia. | Sarcobatus vermiculatus, and Atriplex con- fertifolia. These species represent three life — forms (grass, forb, and shrub), with each life 'Tracy Herbarium, Department of Range Science, Texas A & M University, College Station, Texas 77843. 1 | "Department of Botany and Range Science, Brigham Young University, 435 WIDB, Provo, Utah 84602. 140 | | i} | | | | | January 1986 MORDEN ET AL.: PLANT ECOLOGY 141 TABLE 1. Species along with their mean cover values and life form designation. An asterisk (*) indicates C, species, and the letters indicate life form type (i.e., g = grass; f = forb; s = shrub). 1. Cy, grass sites % cover 2. C; grass sites % cover Allenrolfia occidentalis - s** 2.5 Bromus tectorum - g 2.6 Atriplex patula - f 4.3 Cardaria draba - £ 1.0 *Distichlis spicata - g 30.7 Circium undulatum - f 1.9 Hordeum jubatum - g 3.9 *Distichlis spicata - g 7.0 Juncus balticus - g 3.9) Hutchinsia procumbens - f 1.6 * *Kochia scoparia - f 1.8 Iva axillaris - f 5.8 Lepidium perfoliatium - £ 1.1 Juncus balticus - g 1.0 Polygonum ramosissimum - f 1.3 *Kochia scoparia - f 9.4 Puccinellia nuttalliana - g 34.1 Poa pratensis - g 9.4 Salicornia rubra - f 4.1 *Sporobolus airoides - g 55.5 __ Suaeda depressa - f 11.9 Suaeda depressa - f 1.5 3. C, forb sites % cover 4. C; forb sites % cover Ambrosia artimisiifolia - f 2.1 Bromus tectorum - g 8.3 Atriplex patula - f 44.9 Descurainia sophia - f 4.2 Bromus tectorum - g 2.2 *Distichlis spicata - g 1.3 *Distichlis spicata - g 5.1 Echinochloa crusgalli - g 2.6 Echinochloa crusgalli - ¢ 2.4 *Kochia scoparia - f 60.7 Eleocharis macrostachya - g 1.2 Lepidium perfoliatum - f 14.5 Hordeum jubatum - g 5.8 Puccinellia nuttalliana - g 1.2 ~ *Kochia scoparia - f 13 Ranunculus testiculatus - f 1S Lactuca scariola - £ 1.4 Populus alba - s 15.6 Salix amygdaloides - s 7.9 _ Xanthium strumarium - f 4.4 5. C, shrub sites % cover 6. C; shrub sites % cover Bromus tectorum - g 33.2 *Atriplex confertifolia - s 12.1 Cardaria draba - f ol Bromus tectorum - g 7.9 Descurainia sophia - f 1.9 Kochia americana - f 3.8 Erysimum repandum - f 3.5 Lepidium perfoliatum - f 8.2 Lepidium perfoliatum - f 3.4 Ranunculus testiculatus - f 57.4 Ranunculus testiculatus - f 16.2 Sarcobatus vermiculatus - s ise *Salsola iberica - f OX) Sitanion hystrix - g 1.5 Sarcobatus vermiculatus - s 23.5 Suaeda fruticosa - f ES Sitanion hystrix - g 1.0 Tetradymia spinosa - s 1.0 *Sporobolus airoides - g 2.6 i form represented by both C, and C, species (Table 1). Taxonomic references follow Cron- -quist et al. (1977) for the grasses and Welsh -and Moore (1973) for the dicots. Weather data for Provo, Utah, is represen- ‘tative of the study area. The average annual precipitation is 340 mm, with 60% of the total falling in the winter and spring months. The hottest month of the year is July, averaging 33 } C; the coldest month is January, averaging 3 C. Tributary streams from the Uintah and Wasatch mountain ranges directly east of Utah Lake provide the majority of its water. Precipitation in these mountains ranges from 760 to 1,270 mm annually (Swenson et al. 1972). Field Methods The study sites were selected to depict rep- ‘Tesentative samples of the six community types in the Utah Lake area (Fig. 1). A 10 x 10 m study plot (0.01 ha) was established at each site. Variation in slope, drainage, erosion, and exposure was kept to a minimum. Plots were delineated by a cord 40.0 m long with loops every 10 m for corners. The corners were secured by steel stakes. Twenty 0.25 m° quadrats were placed at regular intervals within the study plot. Density and frequency of all plant species encountered were deter- mined from the quadrat data. Cover values were estimated as suggested by Daubenmire (1959). Only those species showing 1% or more of the total cover are included in the analysis. Christie (1979) found that the top layer of soil is the region of most active mineral up- take. Therefore, soil samples (an 8 cm core) were taken from the top 20 cm of soil in each study plot from opposite corners and the cen- 142 ter. Samples were pooled and then analyzed for texture (Bouyoucos 1951), pH, soluble salts, and mineral content. The hydrogen ion concentration was measured with a glass elec- trode pH meter. Total soluble salts were de- termined with a Beckman electrical conduc- tivity bridge. A paste consisting ofa 1:1 g/v soil to water (distilled) mixture was used in deter- mining pH and soluble salts. Vegetation samples were obtained by tak- ing selected clippings of herbaceous material from the C, species (Sporobolus airoides, Kochia scoparia, and Atriplex confertifolia) and the C, species (Puccinellia nuttalliana, Atriplex patula, and Sarcobatus vermicula- tus) within the study plots. Soil and vegeta- tion samples (new growth leaves) were ana- lyzed for total nitrogen, magnesium, calcium, potassium, sodium (Hesse 1971), zinc, iron, copper, and manganese (Lindsay and Norvell, 1969). Discriminant analysis (Klecka 1975) and analysis of variance (Ott 1977) were used to statistically determine differences in min- eral content between C, and C, species and their habitats. Discriminant analyses were conducted us- ing the Statistical Package for the Social Sci- ences (SPSS) computer program (Klecka 1975). This technique distinguishes statisti- cally between two or more groups of stands on the basis of discriminating variables. The groups and variables are selected by the re- searcher. All variables measured can be used in the analysis (direct method), or a stepwise method can be used to reduce the number of variables to those that provide the best dis- criminating power among the groups. In this study both the direct and the Wilks stepwise methods were used. The Wilks method uses the overall multivariate F ratio to test for vari- able differences. It selects the variables inde- pendently for entry into the analysis based on the importance of their discriminating power. The analysis procedure combines the dis- criminating variables to create discriminant functions designed to provide maximum sepa- ration among the groups previously specified (life forms and C, and C, photosynthetic types). The discriminant program determines the relative percentage of the total variation in the discriminating variables that is accounted for in each function. It also determines the relative importance of each variable used to GREAT BASIN NATURALIST Vol. 46, No. 1 Discriminant Function 2 -6 4 -2 o 2 4 6 Discriminant Function 1 Fig. 2. Discriminant analysis for soil minerals. Of the groups, 83% were classified correctly. Numbers refer to grasses (1), forbs (2), and shrubs (3). 1 1 grasses Discriminant Function 2 Discriminant Function 1 Fig. 3. Discriminant analysis for soil chemistry and texture. Of the groups, 78% were classified correctly. Numbers refer to grasses (1), forbs (2) and shrubs(3). create the discriminant functions. This infor- mation can be used to identify the variables having the greatest influence on the outcome of the analysis. A graphic representation of the results of | discriminant analysis is possible if the discrim- inant functions are viewed as axes in geomet- ric space. A plot of stands based on the two most important functions locates the stands in | ! { | January 1986 alae ae ction 2 x Discriminant Fun Fig. 4. Discriminant analysis for vegetation (leaf) min- erals. Of the groups, 100% were classified correctly. Numbers refer to grasses (1), forbs (2), and shrubs (3). two-dimer sional space in such a way that the relationships among the groups can be visual- ized. Such a graphic representation is espe- cially important for assessing the amount of separation between one group and another as well as the degree of group overlap. RESULTS Results of cover analysis for our study sites are given in Table 1. Only those species show- ing 1% or more of the total cover are included. _In the grass- and forb-dominated communi- ‘ties, Sporobolus airoides, and Kochia sco- | paria provided over half the total cover. The shrub communities, however, were domi- /nated in their understory by the invader spe- cies Bromus tectorum L. and Ranunculus tes- ticulatus Cranz. Their presence in the understory is indicative of site disturbance as a result of grazing. __ Hydrogen ion concentration showed no sig- nificant differences between the communi- ties. All soils were basic, with a pH ranging from 8.1 to 8.7. Soil texture also showed no significant differences between communities, all of them being clay to silty clay loams. Solu- tble salts were highest in the grass communi- MORDEN ETAL.: PLANT ECOLOGY 143 and soil samples within each life form (grasses, forbs, and shrubs) were made. The percent of grouped cases classified correctly were 83% for the soil materials (Fig. 2) and 78% for soil chemistry and texture (Fig. 3), whereas the mineral content of the vegetation classified the groups 100% correctly (Fig. 4). This indi- cates that habitat differences in soil mineral chemistry and texture influence the life form type that dominates a site and that differential partitioning of the minerals by the plants oc- curs to a great extent. The soil, of course, may also be modified by the plants growing in it. Results of analysis of variance and New- man-Kuel tests for parameters of soil and veg- etation mineral content are given in Tables 2 and 3. Analysis of soil mineral content showed significant differences between means for manganese, sodium, and soluble salts. Analy- sis of vegetation mineral content showed dif- ferences between life forms in nitrogen, phos- phorus, zinc, manganese, copper, magne- sium, potassium, and sodium. C, and C, species were 78% correctly classi- fied by minerals for both soil and vegetation samples (Tables 2 and 3). Stem and leaf plots of the discriminant analyses based on soil and vegetation mineral content are given in Fig- ures 5 and 6, respectively. Results of the analysis of variance between C, and C, species are shown in Table 4. Iron and manganese are significantly (p <_ .05) higher in concentrations in the soils of C;- dominated species than of C,-dominated spe- cies. However, concentration of these miner- als within the plant tissue is not significantly different. Calcium and potassium showed no significant differences in concentration in soils but were significantly different in the plant tissues of C, and C, species. Plant:soil ratios were computed for each mineral. Mean differ- ences in iron and sodium assimilation exist between C; and C, species. Although trends existed for other elements, differences be- tween C, and C, species were not significant. DISCUSSION Brown (1978) suggested that differences in nitrogen use between photosynthetic types (C, vs. C,) would hold for grasses, but he was not sure of the results that might be obtained with respect to other life forms. It appears {ties and lowest in the shrub communities. _ Two-dimensional plots of discriminant sanalysis of the mineral content of vegetation 144 GREAT BASIN NATURALIST Vol. 46, No. 1 TABLE2. Differences in the mineral concentrations of soils associated with the life forms as determined by analysis of variance and Newman-Keul tests. Means with similar letters following indicate no significant differences for those means. Those with different letters indicate significant differences. Nutrient concentrations in life form soils Level of Mineral Grasses Forbs Shrubs significance ER Siy isc ee an aaee ras areas TT IG lege ceah coments eerie ae eae Nitrogen 2023a 2026a 1106a NS + 1105 + 1088 + 284 Manganese 10.3ab 15.3b Hole .05 + 8.87 se 8.51 at 2.3 Calcium 15287b 1123la 10368a .05 + 5195 + 2804 + 1497 Magnesium 2002a 1260ab 748b .05 + 1168 + 1309 + 42] Sodium 3428b 1233a 474a 01 + 2086 + 1065 + 209 Salt 7470a 325lab 639b .05 a) Oy + 2506 + 440 TABLE 3. Differences in the mineral concentrations of plant material (leaves) by life form. Differences determined by analysis of variance and Newman-Keul tests. Means with similar letters following indicate no significant differences for those means. Those with different letters indicate significant differences. Nutrient concentrations in life forms Level of Mineral Grasses Forbs Shrubs significance re erTiciiera teen on ahens tae este TING/KiGs 3 es rae tee ee Nitrogen 9908a 22033b 16283b .O1 + 3366 + 3614 + 4220 Phosphorus 943a 2471b 976a 01 + 507 + 608 + 244 Zinc 18.8a 33.1b 12.5a .O1 ate 8.4 ate 15.4 a2 3.06 Magnesium 2903a 7285b 3308a 01 + 2091 + 2414 + 1519 Copper 10.2ab 13.5b 8.3a 01 + Shil ste 3.0 =te ie? Manganese 50.9a 89.6b 72.8ab .O1 eo =f 48.1 ste 47.8 Potassium 4264a 20183b 24175b .O1 +1916 + 7911 + 5472 Sodium 3060a 29079b 51967b 01 + 1467 + 16199 + 9060 from our data that differences do exist in min- ences. These findings are conten) to the re- |). eral uptake for the different life forms. sults of Christie (1979) and Hallock et al. |) Nitrogen content in the plant tissue of C, (1965) and the hypothesis of Brown (1978). In | / and C, species showed no significant differ- fact, the C, shrub A. confertifolia showed less _ Frequency January 1986 1 11 1111122 1111122 1111122 1111122 Frequency Discriminant MORDEN ET AL.: PLANT ECOLOGY 145 Function 1 Fig. 5. Histogram of discriminant analysis for soil minerals. 1 = C, dominated habitats, 2 = C, dominated habitats. 2 2 1 1 2 2 112222122 1122 2214122 Discriminant C3 Function 1 Fig. 6. Histogram of discriminant analysis for vegetation (leaf) minerals. 1 = C, — species, 2 = C3 — species. -nitrogen than the C; shrub S. vermiculatus. The effect of species being adapted to similar habitats may influence this phenomenon and needs to be further investigated. The C; species of this study were growing in soils significantly higher in iron content than _the C, species (Table 4), yet no significant differences in iron and manganese concentra- tion of the vegetation was noted. This may be due to some attribute of C, plants that allows them to live in soils deficient in iron and still assimilate enough for existence. It could also _ be due to iron being present in luxury quanti- ties in both habitat soils and the plants assimi- ‘lating iron without any restrictions. Results of ‘this study also showed calcium to be signifi- scantly higher in C, vegetation than C, vegeta- ‘tion (Table 4). Concentrations of calcium in the soil of both community types is not signifi- cantly different, indicating differential uptake of calcium by the species possessing the differ- ent photosynthetic pathways. Also the C, spe- cies in this study assimilated significantly higher quantities of potassium than did C, species (P < .005). This is in contrast to the results obtained by Hallock et al. (1965). Sodium may be essential as a micronutrient for C, species based upon studies of halo- phytic C, species and nonhalophyte C; spe- cies (Brownell and Crossland 1972). These workers showed that the C, species they stud- ied developed lesions and some species would not flower in the absence of sodium, whereas the C, species grew normally. However, our field results indicate that C, plants do not necessarily require sodium, but that high 146 GREAT BASIN NATURALIST Vol. 46, No. 1 TABLE 4. Summary of mineral concentrations for vegetation and soil data. Means are for vegetation, soil, and plant:soil ratios for C; and C, plants across all 36 study sites. Significant differences are indicated by the presence of an asterisk (*) next to the means values. One asterisk indicates p < 0.05; two asterisks indicate p < 0.01. mg/Kg Vegetation Soil Plant:soil ratio C; C, C; C, C; (Cy N 16127.0 16022.0 1667.0 1771.0 15.3 10.6 + 6910.0 + 5611.0 + 1198.0 a2 ol) +15.2 + 4.9 P 1489.0 1438.0 47.0 29.0 79.7 79.0 + 825.0 + 915.0 + 104.9 ae 32.3 + 49.2 + 54.0 Zn 22.8 21.0 3.5 8 18.0 21.3 Sel oll se) UL! ete 6.6 =e 2.0 +11.5 +13.9 Fe 269.0 349.0 16.1* 6.5* 28.1 60.3 + 220.8 + 504.0 =e ASA! ste 5.1 + 27.0 +63.3 Mn 82.9 59.4 14.4 7.4 9.5 8.6 a= Sf) + 48.6 ao 9.3 ste 3.7 + 9.6 a= ©) 7/ Cu 10.2 11.1 2.5 1.5 5.4 8.5 ae 3.6 ste 3.2 ae 1s) ar 6 as @hl + 3.9 Ca 10089. 0* 13017.0* 12306.9 12644.0 0.9 2.0 + 4301.0 + 5875.0 + 3852.7 + 4609.7 + 0.4 = ES Mg 4390.0 4602.0 1457.0 1217.0 4.6 5.5 + 3420.0 + 2162.0 a 1A 5) + 848.2 as Zt Il + 5.0 K 13433.0** 18965.0** 1022.0 1370.0 19.6 16.4 + 8274.0 +11589.0 ae Ohya) + §889.1 +21.0 +19.5 Na 30778.0 25292.0 1766.0 1658.0 76.8 42.7 + 22431.0 + 23463.0 + 1750.7 + 1995.9 + 88.1 +53.3 Salt 3921.0 3653.0 + 5083.0 + 4025.0 pH 8.3 8.3 ate 0.5 Ste 0.3 Clay 33.4 30.8 + 13.0 as) + lle),c8) Silt 46.3 47.6 =ts 16.3 = 17.8 Sand 20.2 Pls a at 11.8 Ete 15.5 sodium uptake is characteristic of plants use efficiency than plants possessing the C; | adapted to the saline habitat (Table 4). pathway, which would aid in survival of the | Salinity would have the same effect on plant in semiarid regions (Ludlow 1976). C, plant-water relations as increasing plant plants may have a competitive advantage in — drought: the more salt present in a soil, the — saline environments due to their high water- wetter the soil must be to dilute the salt and use efficiency. Although no significant differ- | i, ih prevent salt hindrance to growth (Donahue et _ ences between salt or sodium levels in the two‘ al. 1977). Plants possessing the C, photosyn- _ habitat types and in the C, and C, plants were — thetic pathway typically have a higher water observed, it is important to note that the C3 1 i January 1986 MORDEN ET AL species of this study had slightly higher mean levels of sodium in their tissues and higher plant:soil ratios than the C, species. The C; spe- cies also flowered one to two months earlier in ‘the summer, when moisture conditions in the | hahitat were more conducive to their growth. though grasses grew in soils with high con- centrations of soluble salts (Table 2), their tissue ‘concentrations of sodium and potassium (Table ~3) were much lower than either forbs or shrubs. Although many grasses adapted to saline envi- ‘ronments possess salt glands (Liphschitz et al. 1974, Hansen et al. 1976), Puccinellia nuttal- ‘liana and Sporobolus airoides do not and as a result must restrict the amount of sodium and potassium entering their tissues. On the other ‘hand, the shrubs and forbs of this study do pos- ssess salt glands or become succulent (Luttge /1971) and thus are able to tolerate higher quanti- ties of sodium and potassium in their tissues. Both growth form and photosynthetic type showed habitat differences relative to mineral uptake. The detailed physiological basis for these differences must now be further investi- gated. | | | | LITERATURE CITED BLENKINSOP, P. G., AND J. E. DALE. 1974. The effects of nitrate supply and grain reserves on fraction I protein levels in the first leaf of barley. J. Expt. Bot. 25:913-926. Bovuyoucos, G. J. 1951. A recalibration of the hydrometer method for making mechanical analysis of soils. J. Agron. 43:434—438. | Brown, R. H. 1978. A difference in N use efficiency in C; and | C, plants and its implications in adaptation and evolu- tion. Crop Sci. 18:93-98. BROWNELL, P. F., ANDC. J. CROSSLAND. 1972. The requirement | for sodium as a micronutrient by species having the C, | dicarboxylic photosynthesis pathway. Plant Physiol. 49:794-797. | CuristIE, E. K. 1979. Ecosystem processes in semiarid grass- lands. I. Litter production, decomposition, and nu- trient dynamics. Aust. J. Agric. Res. 30:29—42. CRONQUIST, A., A. H. HOLMGREN, N. H. HOLMGREN, J. L. RE- VEAL, AND P. K. HOLMGREN. 1977. Intermountain flora, Vol. 6, Monocotyledons, Columbia University Press, New York. 584 pp. .. PLANT ECOLOGY 147 DAUBENMIRE, R. 1959. A canopy-coverage method of veg- etational analysis. Northwest Sci. 33:43—46. DONAHUE, R. L., R. W. MILLER, J. C. SCHICKLUMNA. 1977. Soils, an introduction to soils and plant growth. Prentice-Hall, New Jersey. HALLOCK, D. L., R. H. BROWN, AND R. E. BLASER. 1965. Relative yield and composition of Ky. 31 fescue and coastal bermudagrass at four nitrogen levels. Agron. J. 57:539-542. HEssE, P. R. 1971. Textbook of soil chemical analysis. Wm. Clowes and Sons, Ltd., London. 520 pp. HANSEN, D. J., P. DAYANANDAN, P. B. KAUFMAN, AND J. D. BROTHERSON. 1976. Ecological adaptation of salt marsh grass, Distichlis spicata (Gramineae), and environmental factors affecting its growth and dis- tribution. Amer. J. Bot. 63(5):635—650. KLECKA, W. R. 1975. Discriminant analysis. Pages 434—467 in SPSS, statistical package for the social sciences. N. H. Nie, C. H. Hull, J. G. Jankens, K. Steinbrenner, D. H. Bent, eds., McGraw-Hill, New York. Linpsay, W. L., AND W. A. NORVELL. 1969. Development of a DTPA micronutrient soil test, Agron Ab- stracts. p. 84. Equilibrium relationships of Zn?* Fe**, Ca’*, and H* with EDTA and DTPA in soil. Soil Sci. Soc. Amer. Proc. 33:62—68. LipHSCHITz, N., ADIVA-SHOMER-ILAN, A. ESHEL, AND Y. WAISEL. 1974. Salt glands on leaves of rhodes grass (Chloris gayana Kth.). Ann. Bot. 38: 459-462. LupLow, M. M. 1976. Ecophysiology of C, grasses. Pages 364-386 in O. Lange. L. Kapper, E. Schulze, eds., Water and plant life. Problems and modern approaches. LutrcE, U. 1971. Structure and function of plant glands. Ann. Rev. Plant Physiol. 22:23—44. Orr, L. 1977. An introduction to statistical methods and data analysis. Duxbury Press, Massachusetts. SWENSON, J. L., W. M. ARCHER, K. M. DONALDSON, J. J. SHIOZAKI, J. H. BRODERICK, AND L. WOODWARD. 1972. Soil survey of Utah County, Utah. USDA Soil Conservation Service. 161 pp. WELSH, S. L., AND G. Moore. 1973. Utah plants—Tra- cheophyta, 3d ed., Brigham Young University Press, Provo, Utah. 474 pp. WILSON, J. R. 1975. Comparative responses to nitrogen deficiency of a tropical and temperate grass in the interrelation between photosynthesis, growth and accumulation of nonstructural carbohydrate. Neth. J. Agric. Sci. 23:104—112. WILSON, J. R., AND K. P. HayDock. 1971. The comparative response of tropical and temperate grasses to vary- ing levels of nitrogen and phosphorus nutrition. Aus. J. Agric. Res. 22:573—587. RESPONSE OF WINTERFAT (CERATOIDES LANATA) COMMUNITIES TO RELEASE FROM GRAZING PRESSURE Lars L. Rasmussen’ and Jack D. Brotherson? ABSTRACT. —Sixteen study sites were established in grazed and ungrazed stands of winterfat in Kane County, Utah. The area is located within the winter range of cattle and along U.S. Highway 89 between Kanab, Utah, and Page, Arizona. Road construction in 1957 dissected several winterfat communities, and following fencing part of the communities were released from grazing. Differences in species composition, vegetation, and soil characteristics between grazed and ungrazed sites were assessed. Major differences in site characteristics appeared due to the influence of winter grazing by cattle. Winterfat and Indian ricegrass showed increased cover on the nongrazed sites following release from grazing pressure. Winterfat also showed significant negative interspecific association patterns with all major species. Winterfat (Ceratoides lanata [Pursh] J. T. Howell) is considered a valuable forage com- ponent of winter ranges throughout western North America. Blauer et al. (1976) described winterfat as “superior nutritious browse for livestock and big game.” Griffiths (1910) pointed out that winterfat is “very much in- UTAH ARIZONA Fig. 1. Map of study site location in Kane County, Utah. Glen Canyon jured by overgrazing.” However, more recent research has provided somewhat conflicting information relative to the tolerance of win- terfat to grazing pressure. Holmgren and Hutchings (1971) indicated that percent plant cover represented by winterfat sharply de- clines under heavy grazing during late winter. Salt Lake City UTAH Kane Co. @ STUDY AREA ‘Ellensburg District Office, Soil Conservation Service, Ellensburg, Washington 98926. "Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 148 ( ( ] } \ j January 1986 RASMUSSEN, BROTHERSON: WINTERFAT COMMUNITIES 149 TABLE 1. Means and standard deviations (SD) of soil factors and significance levels for the difference between the means for the grazed and nongrazed winterfat populations in Kane County, Utah. Significance levels were computed by using the student's t-statistic. { | *Significant difference between means at .05 level. t | ) | Trlica and Cook (1971) showed that winterfat did not make good growth recovery under any defoliation treatment. Yet, Norton (1978) ‘states that winterfat is “relatively indifferent to heavy grazing.” The purpose of the present study was to -examine changes in winterfat communities following 26 years of release from grazing pressure on ranges in Kane County, Utah. STUDY AREA U.S. Highway 89 from Kanab, Utah, to Page, Arizona, was constructed during 1957, vand the right-of-way was fenced. This con- struction dissected winterfat communities 5 km east of the Paria River, creating grazed and ungrazed units (Fig. 1). The area is located within the Bureau of Land Management East Clark Bench allot- ment. This allotment has been utilized pri- marily as winter range for cattle since 1956. Although entry and removal dates for live- stock have varied, 1 November to 31 May was the general season of use until 1964, when the ‘removal date for livestock was moved back to April 30. The climatic conditions of the study area are similar to conditions at Glen Canyon City, Utah, 15 km east along U.S. 89. Average an- jaual precipitation in the area varies from 15 to 20 cm. There are two main periods of precipi- sation, one beginning in December and end- SITE FACTOR GRAZED UNGRAZED Mean SD Mean SD Sand (%) 83.8 2.42 83.1 2.94 Silt (%)* 8.2 1.12 9.2 0.83 Clay (%) 8.6 2.47 7.6 2.98 Organic matter (%)* 0.3 0.05 0.2 0.05 pH 8.0 0.05 7.8 0.05 EC x 10° 0.5 0.05 0.6 0.36 CEC (meq/100 g) 8.6 1.31 8.8 1.32 Calcium (ppm)* 5194.0 768.0 4375.0 1040.0 Magnesium (ppm) 109.5 32.6 177.0 26.5 Sodium (ppm) 18433 6.26 25.6 47.3 Potassium (ppm) 226.9 45.1 264.8 90.8 Iron (ppm) 1.4 0.24 1.3 0.11 Manganese (ppm) Doll 0.57 2.6 0.35 Zinc (ppm) 0.6 0.27 0.6 0.21 Copper (ppm) 0.4 0.05 0.4 0.07 | Phosphorus (ppm) 31.0 5.31 37.0 8.44 ing in March in the form of snow and the second beginning in August and ending in October in the form of rain (Green et al. 1981). The hottest month of the year is July, with an average tem- perature of 28 C. The coldest month is January, with an average temperature of 0 C. The frost- free period for the area begins in late April and ends in late October, averaging 190 days (U.S. Environmental Data Service 1968). METHODS Sixteen stands were sampled during June and July 1984 (eight each within grazed and un- grazed sites), and data were collected to repre- sent conditions within grazed and ungrazed win- terfat communities. Each stand was sub- sampled, with a total of 11 1-m* quadrats placed one every 3 m along 33-m transect lines. Tran- sects within the protected sites were placed par- allel to and equidistant between the fence line and U.S. Highway 89. Transects within the grazed sites were placed parallel to transects in protected sites and at equal distances from the fence line. Total living cover of vascular plants was esti- mated in each quadrat. Cover by life forms, soil cryptogams, litter, exposed rock, bare ground, and individual plant species were estimated us- ing Daubenmire’s cover classes (1959). Three soil samples were taken at 10-m inter- vals along each transect line from the top 20 cm of soil. The three samples were later combined % SIMILARITY 10 20 30 40 50 GREAT BASIN NATURALIST Vol. 46, No. 1 60 70 80 90 100. SSS SSS sf) | ' 5 Non-grazed 7 Non-grazed 3 Non-grazed 1 Non-grazed 4 Non-grazed 6 Non-grazed 8 Non-grazed 2 Non-grazed 9 Grazed 13 Grazed 15 Grazed iW 16 Grazed 10 Grazed 11 Grazed 12 Grazed | 14 Grazed Fig. 2. Cluster dendrogram of grazed and nongrazed winterfat stands in Kane County, Utah. Cluster is based on | similarity of plant species cover in site vegetation. for laboratory analysis. Ludwig (1969) found that the surface decimeter of soil yields 80% of the information useful in correlating plant re- sponse with concentrations of essential min- eral nutrients in the soil. Holmgren and Brewster (1972) showed that greater than 50% of the fine roots of plants (which included winterfat) in Utah desert communities are found in the top 15 cm of soil profile. Soil samples were analyzed for texture (Bouyoucos 1951), pH, soluble salts, mineral composition and organic matter. Soil pH was determined with a glass electrode pH me- ter.Soluble salts were determined with a Beckman electrical conductivity bridge. Ex- changeable calcium, magnesium, potassium, and sodium were extracted from soils with DTPA (diethylene triamine-penta-acetic acid; Lindsay and Norvell 1969). A Perkin Elmer Model 403 atomic absorption spectrophoto- meter was used to determine individual ion concentrations (Isaac and Kerber 1971). Phos- | phorus was extracted with sodium bicarbon- _ ate (Olsen et al. 1954). Organic matter was, , estimated from total carbon using methods | described by Allison (1965). Similarity indices comparing each stand tc. all other stands were calculated (Ruzickelf 1958). These indicies were then employed tc } cluster winterfat stands following Sneath anc, , Sokal (1973). Individual plant species were , also clustered on the basis of niche overlay | (Colwell and Futuyma 1971). Interspecific as, sociation patterns between plant species were! computed using Cole’s (1949) Index. Mean: , and standard deviations were computed for al biotic and abiotic variables across the 1¢ |, stands. Prevalent species were selected o1 . the basis of cover values (Warner and Harpe. 1972). Diversity indices were computed fol lowing Shannon and Weaver (1949) anc ' McArthur (1972.) Statistical differences be ; tween grazed and ungrazed sites were caleu ,. lated using Student’s t-statistic. January 1986 % NICHE OVERLAP 10 20 30 40 50 60 70 80 90 100 Yucca navajoa Ambrosia acanthocarpa Sphaeralcea coccinea Hilaria jamesii Oryzopsis hymenoides Ceratoides lanata Bromus tectorum Vulpia octoflora Menizelia albicaulis Sporobolus cryptandrus Xanthocephalum sarothrae Aristida purpurea Ephedra torreyana Opuntia phaeacantha Salsola iberica Phacelia ivisiana Cleome lutea Helianthus deserticola Plantago insularis Sitanion hystrix Fig. 3. Cluster dendrogram of plant species occurring in the study area. Cluster based on niche overlap relative toa species geographical distribution. RESULTS AND DISCUSSION There were few differences in edaphic fac- tors between grazed and ungrazed winterfat communities (Table 1). Significant differences between means were observed for percent silt, percent organic matter, and calcium con- centrations. The higher levels of silt in the ungrazed area are probably due to the pres- ence of the fence. The fence (net wire) would act as a barrier to blowing weeds and plant / material and thus as a barrier to drifting soil. The more-abundant vegetation on the un- | grazed sites would also create a barrier against ‘which windblown silt would tend to accumu- late. The differences in percent organic mat- 'ter are also probably a function of increased | vegetation cover. Mean differences in these | three factors are relatively small, and floristic ‘differences between grazed and ungrazed ‘stands are, therefore, not considered to be _ caused by these soil characteristics. _ Cluster analysis (based on vegetative simi- larity) clearly separated the grazed and un- (grazed transects into two groups (Figs. 2 and '3). This separation reflects the effects of long \periods of winter livestock grazing on vegeta- tive composition. Grazing-induced change is RASMUSSEN, BROTHERSON: WINTERFAT COMMUNITIES 151 also indicated by greater diversity within the grazed stands (Table 2). The greater plant di- versities observed among grazed winterfat stands is expected. Cox (1976) noted that Charles Darwin observed greater diversity within grazed lands when compared with non- grazed lands (Cox et al. 1976). Harper (1977) also indicates that the great floristic diversity within the Chalk grasslands of Britain owes its existence to the selective grazing of livestock on potentially dominant plant species. Fur- ther research may better establish the occur- rence of this phenomenon on other western ranges. A major difference between grazed and un- grazed winterfat communities was in shrub cover (Table 2). Total live cover and litter cover were also greater on the ungrazed areas. Differences in community response to release from grazing pressure is best shown by differ- ences in species cover (Table 3). The cover of winterfat on ungrazed areas was significantly greater than on grazed areas. Greater shrub cover on the ungrazed stands was entirely accounted for by the increased cover of win- terfat. Although winterfat has been described as being relatively tolerant to grazing and as a “good natural increasor” (Blauer et al. 1976), it demonstrates little tolerance to winter use by cattle on our study sites, where it was poten- tially the dominant shrub. Wide ecotypic variation is known to exist between winterfat populations (Stevens et al. 1977). Populations examined in this study were characterized by relatively large plants (up to three feet in height) with a growth form similar to big sagebrush (Artemisia triden- tata). The genetic differences of this ecotype may account in part for its susceptibility to grazing. Cover of cool-season grasses on the un- grazed sites was greater than on grazed sites, with the cover values of Indian ricegrass (Ory- zopsis hymenoides) showing the greatest dif- ference. This difference was probably due to grazing pressure during the late winter season when Indian ricegrass actively grows. Grazing while the grass is actively growing would stress the plant and reduce its capacity to compete. The warm-season grass species, gal- leta (Hilaria jamesii), which does not actively grow during the late winter grazing season, maintained nearly equivalent cover values be- 152 GREAT BASIN NATURALIST Vol. 46, No. 1 TABLE 2. Means and standard deviations (SD) of site factors and significance levels for the difference between the means for the grazed and nongrazed winterfat populations in Kane County, Utah. Significance levels were computed using the student's t-statistic. SITE FACTOR GRAZED Mean SD Total life cover (%) 29.3 4.74 Exposed rock (%) 0.1 0.18 Bare soil (%) 24.9 3.67 Litter cover (%) 15.2 8.38 Cryptogram cover (%) 40.5 9.72 Shrub cover (%) CP 3.18 Perennial grass cover (%) 20.4 eK Annual grass cover (%) 4.2 3.26 Perennial forb cover (%) 0.3 0.54 Annual forb cover (%) 3.6 4.85 Diversity: Shannon-Weaver 3.3 0.4 MacArthur 4.8 1130) TABLE 3. Means and standard deviations (SD) of plant species cover occurring in grazed and nongrazed winterfat communities in Kane County, Utah. SPECIES GRAZED Mean Ambrosia acanthocarpa 0.1 Aristida purpurea 1.6 Bromus tectorum 0.4 Ceratoides lanata BS Cleome lutea 0.1 Ephedra torryana 1.0 Helianthus deserticola 0.1 Hilaria jamesii 15.9 Mentzelia albicaulis 1.0 Opuntia phaeacantha 0.2 Oryzopsis hymenoides 6.7 Phacilia ivesiana 0.1 Plantago insularis 0.3 Salsola iberica 2.4 Sitanion hystrix 0 Sporobolus cryptandrus Be) Sphaeralcea coccinea 0.1 Vulpia octoflora 4.1 Xanthocephalum sarothrae Dull Yucca navajoa ili tween the two sites. Conversely, sand drop- seed (Sporobolus cryptandrus), demonstra- ted increased representation on the grazed sites. This may be due to the pressure of live- stock foraging, reducing competition on the grazed sites by opening up the vegetation cover and thus allowing room for expansion of sand dropseed. Sand dropseed is well adapted to sandy soils and will increase or even invade SIGNIFICANCE UNGRAZED LEVEL Mean SD 32.6 4.14 0.10 0.1 0.18 NS 25.1 5.95 NS 26.1 4.81 0.005 34.7 8.66 NS 13.4 4.25 0.10 18.8 8.21 NS 5.1 3.79 NS 0.1 0.25 NS 1.3 1.23 NS 9,9) 0.5 0.10 ood 1.4 0.10 UNGRAZED SD Mean SD | 0.09 (es 1.47 0.3 0.95 0.48. 0.7 11 1.44 13.9 4.83 0.09 0 2.24 0 h 0.31 T 8.07 16.1 6.47 [ 1.20 1.1 1.19 0.49 0 | 2.88 10.5 4.21 i 0.03 0.1 0.03 | 0.81 0.1 0.14 \| 4.13 0.2 0.49 i 0.1 0.21 | 1.97 0.1 0.17 i 0.18 0.1 0.24 | 2.88 4.5 3.46 Ih 2.80 Li 2.16 I 0 I if the proper conditions are present. Other | : species showing increases on the grazed sites ||! were snakeweed (Gutierrezia sarothrae) and | | Russian thistle (Salsola iberica). Both species are considered as increasers and/or invaders on rangelands in the western United States. | Percent cover of forbs was also greater | among grazed stands. Russian thistle is pri- | " marily responsible for the increased represen- | January 1986 GROUP A Menizelia albicaulis Salsola iberica Opuntia Sages Phacelia ivisiana Bromus tectorum Vulpia octoflora RASMUSSEN, BROTHERSON: WINTERFAT COMMUNITIES 153 GROUP B Plantago insularis Sporobolus Lee Xanthocephalum | cryptandrus = sarothrae Ephedra torreyana Aristida purpurea Fig. 4. Cluster of plant species associated with grazed and nongrazed sites as determined by Cole’s (1949) Index. The more lines between the species, the greater the association. All associations are statistically significant. tation of forbs on the grazed areas. Brotherson and Brotherson (1981) also reported greater cover of forbs in grazed sagebrush communi- ties than in ungrazed communities. Increases in forb cover were due primarily to exotic annuals. To further understand species interaction, ‘niche overlap values were clustered to assess zeographical association patterns among the species (Fig. 4). Five groups clustered to- gether, three of which are of particular inter- 2st. These three groups centered around spe- cies that can be labeled decreasers, increa- sers, and opportunists. The remaining two roups seem of little significance. The de- sreasers group was characterized by winterfat ‘and Indian ricegrass. This group also included “species (winter annuals) that are not generally considered decreasers. However, all species on the group showed less cover on grazed than jon ungrazed areas. The increasers include snakeweed, sand dropseed, purple threeawn Aristida purpurea) and Torrey mormontea (Ephedra torreyana). Each of the species in this group had greater cover on the grazed sites and displayed some degree of domi- nance. The opportunists are generally consid- ered unpalatable and showed increased cover on the grazed sites. This group is character- ized by Russian thistle and prickly pear (Op- untia phaeacantha). To define these relationships more pre- cisely, we employed the use of Cole’s (1949) Index of interspecific association (Table 4). In this case, two groups of species were apparent from the analysis. Group A contains seven species, five of which are annuals, whereas group B contains species that are mostly perennial. The species of group B are gener- ally more important in the grazed sites with respect to cover values, and the species in group A show little preference for either side of the fence. Each group contains species that show positive affinities for species within that group and negative relationships for species found in the opposite group. The two groups 154 GREAT BASIN NATURALIST Vol. 46, No. 1 TABLE 4. Results of Cole’s Index analyses with respect to the interspecific association patterns of species found growing in conjunction with winterfat populations in Kane County, Utah. Significance levels of the chi-square values are as follows: 0.05 = p= 3.85, 0.01 = p = 6.64, and 0.01 = p= 11.21. Positive associations Negative associations Species Species X’ Coef. SD Species X’? Coef. SD Group “A” Bromus tectorum Salsalo iberica 19.5 0.286 0.064 Ceratoides lanata 6.0 0.299 0.122 Sporobolus cryptandrus 11.8 0.263 0.076 Hileria jamesii Mentzelia albicaulis 10.1 0.124 0.039 Oryzopsis hymenoides 9.5 0.578 0.187 Plantago insularis 13.3 0.076 0.021 Vulpia octoflora 6.1 0.411 0.167 Salsola iberica 4.2 0.054 0.026 Mentzelia albicaulus Salsola iberica 6.7 0.130 0.050 Aristida purpurea 5.6 0.702 0.296 Opuntia phaecantha Phacelia iviciana 43.2 1.000 0.152 Salsola iberica 4.5 1.000 0.470 Phacelia iviciana Salsola iberica 8.9 0.694 0.233 Ceratoides lanata 5.2 1.000 0.441 Salsola iberica Sporobolus cryptandrus 11.4 0.302 0.089 Ceratoides lanata 14.9 0.551 0.143 Vulpia octoflora 4.6 0.551 0.157 Oryzopsis hymenoides 4.4 0.185 0.088 Vulpia octoflora Ceratoides lanata 5.2 0.309 0.136 Hileria jamesii 6.1 0.412 0.167 Group “B” Aristida purpurea Ephedra torreyana 6.1 0.080 0.032 Ceratoides lanata 8.6 0.551 0.187 Sporobolus cryptandrus 8.5 0.343 0.118 Mentzelia albicaulis 5.6 0.702 0.296 Ephrdra torreyana Sporobolus cryptandrus 13.1 1.000 0.277 Oryzopsis hymenoides 13.4 1.000 0.272 Xanthocephalum sarothrae 4.9 1.000 0.451 Plantago insularis Hileria jamesii 13.3 0.076 0.021 Sporobolus cryptandrus 4.1 0.224 0.112 Sporobolus cryptandrus Xanthocephalum sarothrae 28.0 0.651 0.123 Ceratoides lanata 19.4 0.530 0.120 Xanthocephalum sarothrae are bridged by two species: sand dropseed and, toa lesser extent, desert plantain (Plantago insu- laris). The existence of the two groups indicates the species belonging to each group are doing quite different things with respect to their present environment. The underlying reasons for the groupings are unknown. Also of interest from the analysis is the fact that neither winterfat nor Indian ricegrass showed any positive correlations. In both cases, all indicated relationships with other species were negative. Winterfat, for example, had a total of 16 negative correlations out of a possible total of 20. Of these, 9 were significant (p < 0.05). With release from grazing, the individual plants of winterfat grow to be much larger in stature and increase in density (8,409 individu- als/ha in the nongrazed area vs. 2,414 individu- als/ha in the grazed areas). Such changes place winterfat plants in a highly competitive position with respect to other understory species. These changes would increase winterfat’s crowding and shading ability. Most of the species showing negative association patterns with winterfat are shade intolerant. Ceratoides lanata 6.8 0.201 0.077 Smith (1959) indicates that patterns of in- terspecific association between species can change with varying degrees of grazing pres- sure. Further, Cook and Hurst (1962), in a \ study done in the Escalante deserts of south- — ern Utah, showed that negative association | patterns intensified between winterfat and — the two species Indian ricegrass and yellow- brush (Chrysothamnus stenophyllus ). The in- tensified negative relationships that devel- oped with yellowbrush happened because it and winterfat responded differently to varying grazing pressures. Winterfat was shown to decrease in the face of heavy grazing pressure _ whereas yellowbrush increased under similar grazing conditions. The intensification of the © negative associations between winterfat and © Indian ricegrass developed for opposite rea- sons. In this case both species showed in- creased prominence to release from heavy | grazing, but under heavy grazing conditions | their association patterns were essentially neutral. This suggests the development of strong competition between the two species | when they are released from grazing and January 1986 growing sympatrically. Both cases appear to be happening with respect to winterfat and its in- terspecific association patterns as measured in our study. In the grazed areas of our study, win- terfat is being eliminated as a result of winter use, and other species are expanding into the vacated space, thus creating the opportunity for increased competition and negative associa- tions. Conversely, in the ungrazed sites winter- fat is expanding in prominence, thereby creating conditions for the intensification of competition between itself and other species. Reasons are not always apparent or easily understood. To gain a total explanation, further studies of the autecology of the species involved seems neces- sary. It is evident that release from winter grazing on the East Clark Bench allotment has had major impacts on the winterfat communities exam- ined. Following 26 years without grazing pres- sure, floristic diversity decreased within the winterfat communities. Winterfat and Indian ricegrass showed dramatic increases in cover when released from grazing pressure. These species are likely the primary decreasers under the present management system, demonstrating lowered tolerance to grazing. It is reasonable to assume that damage to these species is due to late winter season utilization. Holmgren and Hutchings (1972) also report marked decreases in winterfat cover when the species was grazed during late winter after its growth had begun. LITERATURE CITED _ ALLISON, L. E. 1965. Organic Carbon. Pages 1320-1354 - inC. A. Black, D. D. Evans, J. L. White, L. E. Ensmigner, F. E. Clark, and R. C. Dinauer, (eds.), Methods of soil analysis—chemical and mi- crobiological properties. Agronomy Series No. 9, Part 2. American Society of Agronomy, Inc. Madison, Wisconsin. BLAUER, C. A., A. PLUMMER, E. MCARTHUR, R. STEVENS, AND B. C. Giunta. 1976. Characteristics and hy- bridization of important intermountain shrubs: I, chenopod family. USDA, Forest Service Pap. INT-177: Intermountain Forest and Range Exper- iment Station, Ogden, Utah. Bouyoucos, G. J. 1951. A recalibration of the hydrometer method for making mechanical analysis of soils. J. Agron. 43:434—438. SBROTHERSON, J. D., AND W. T. BROTHERSON. 1981. Graz- ing impacts on the sagebrush communities of cen- tral Utah. Great Basin Nat. 41:335—340. RASMUSSEN, BROTHERSON: WINTERFAT COMMUNITIES 155 Cote, L. C. 1949. The measurement of interspecific asso- ciation. Ecology 30:411—424. Cook, C. W., AND R. Hurst. 1962. A quantitative measure of plant association on ranges in good and poor condition. J. Range Manage. 15:266—273. CowELL, R. K., AND E. J. FuruyMa. 1971. On the mea- surement of niche breadth and niche overlap. Ecology 52:567—576. Cox, C., I. N. HEALEY, AND P. D. Moore. 1976. Biogeogra- phy, 2d ed. John Wiley and Sons, New York. 194 DAUBENMIRE, R. 1959. A canopy coverage method of veg- etational analysis. Northwest Sci. 33:43-66. GREER, D. C., K. D. GurGLE, W. L. Wa.ouist, H. A. CHRISTY, AND G. B. PETERSON. 1981. Atlas of Utah. Brigham Young University Press, Provo. 300 pp. GriFFITHS, D. A. 1910. A protected stock range in Ari- zona. USDA, Bur. Plant Bull. 177. 28 pp. Harper, J. L. 1977. Population biology of plants. Aca- demic Press, New York. 892 pp. HOLMGREN, R. C., AND S. F. BREWSTER. 1972. Distribu- tion of organic matter reserve in a desert shrub community. USDA, Forest Service Res. Pap. INT-130. Intermountain Forest and Range Exper- iment Station, Ogden, Utah. HOLMGREN, R.C., ANDS. S. HUTCHINGS. 1972. Salt Desert shrub response to grazing use. Pages 153-164 in C. M. McKell, J. P. Blaisdell, and J. R. Goodin, eds., Wildlife shrubs—their biology and utiliza- tion. USDA. Forest Service General Technical Report INT-1, Intermountain Forest and Range Experiment Station, Ogden, Utah. Isaac, R. A., AND J. D. KERBER. 1971. Atomic absorption and flame photometry techniques and uses in soil, plant, and water analysis. Pages 17—38 in L. M. Walsh, ed., Instrumental methods for analysis of soils and plant tissue. Soil Sci. Soc. Amer. Proc. Madison, Wisconsin. Linpsay, W. L., AND W. A. NORVELL. 1969. Development of DTPA micronutrient soil test, page 84 in Agron. Abstracts. Equilibrium relationships of Zn", Fe®*, Ca’* and H* with EDTA and DTPA in soil. Soil Sci. Soc. Amer. Proc. 33:62-68. Lupwic, J. A. 1969. Environmental interpretation of foothill grassland communities of northern Utah. Unpublished dissertation, University of Utah, Salt Lake City, Utah. 100 pp. MACARTHUR, R. H. 1972. Geographical ecology: patterns in the distribution of species. Harper and Row Publishers, New York. 269 pp. Norton, B. 1978. The impact of sheep grazing on long- term successional trends in salt desert shrub vege- tation of southwestern Utah. Proc. Intern. Range- land Cong. 1:610-613. OLSEN, S. R., C. V. COLE, F.S. WATANABE, AND L. A. DEAN. 1954. Estimation of available phosphorus in soils by extraction with sodium bicarbonate. U.S. Dept. Agric. Cir. No. 939. Ruzicka, M. 1958. Anwendund mathematisch-statistis- cher methoden in der geobutanik (Synthetische bearbeitung von aufnahmen). Biolgia, Bratsil. 13:647-661. ; SHANNON, C. E., AND W. WEAVER. 1949. The mathematical theory of communication. University of Illinois Press, Urbana. 177 pp. 156 SmiTH, D. R. 1959. Changes in interspecific associations re- lated to grazing pressures. J. Range Manage. 12:309-311. SNEATH, R. H. A., AND R. R. SOKAL. 1973. Numerical taxonomy: principles and practice of numerical classification. W. M. Freeman Co., San Francisco. 573 pp. STEVENS, R., B. C. GiunTA, K. R. JORGENSEN, AND A. P. PLUM- MER. 1977. Winterfat. Publication 77-2, Utah Divi- sion of Wildlife Resources, Salt Lake City, Utah. GREAT BASIN NATURALIST Vol. 46, No. 1 TrRLICA, M. J., AND C. W. Cook. 1971. Defoliation effects on carbohydrate reserves of desert species. J. Range Manage. 24:418—425. U.S. ENVIRONMENTAL Data SERVICE. 1968. Climatic atlas of the United States. U.S. Govt. Print. Off., Washington, D.C. 80 pp. WARNER, J. H., AND K. T. HaRPER. 1972. Understory char- acteristics related to site quality for aspen in Utah. Brigham Young Univ. Sci. Bull., Biol. Ser. 16(2):1—20. ISOZYMES OF AN AUTOPOLYPLOID SHRUB, ATRIPLEX CANESCENS (CHENOPODIACEAE) E. Durant McArthur’, Stewart C. Sanderson!, and D. Carl Freeman? ABSTRACT.— Diploid, tetraploid, and hexaploid populations of Atriplex canescens (x =9) were examined for 18 isozyme systems. Of 24 interpretable loci, only one locus (Per,) was polymorphic. Another locus (Per) showed a dosage effect. Genetic distance values, D, ranged from near 0 to 0.05, which are in the normal range for local species races. Results from clonal ramets gave identical results. The data and analyses support an essentially autopolyploid origin for the polyploid populations examined. Atriplex canescens (Pursh) Nutt. is a widely distributed shrub in western North America. It occurs in chromosome races of 2x—12x. Te- traploids (4x =2n =36) are most common over a majority of the range of the species, but diploids and hexaploids are not infrequent in some widely distributed areas (Stutz and Sanderson 1979, McArthur and Freeman 1982). Higher polyploids are restricted in dis- tribution. Stutz et al. (1975) presented mei- otic evidence (2.18 IVs/cell, range of 0-6) in- dicating that the 4x populations are autopo- ‘lyploid. Diploid (2x) A. canescens is essen- tially dioecious; 4x and 6x forms are trioe- cious—?, 36 and [¢%, do] (McArthur 1977, McArthur and Freeman 1982). As might be expected in an outcrossing species, much morphological variation is evident within and between populations (McArthur et al. 1983). We undertook a study to examine isozyme patterns among polyploid levels, genders, and ecologically separated subpopulations of A. canescens. MATERIALS AND METHODS We selected four populations (Table 1) for study. One, Kingston Canyon, was divided into two subpopulations because of strikingly different ecological conditions (plant commu- nities, slope, moisture relationships, soils— McArthur et al. unpublished data) and be- cause Freeman et al. (1976) showed that plants of the related species A. confertifolia tend to segregate 2 versus ¢ on environmen- tally different sites. Seven plants from each sexual state were randomly sampled from each population. Thus, we sampled 14 plants from the Little Sahara Sand Dunes, 21 from Spanish Fork Canyon, 35 from Kingston Canyon, and 21 from near Grantsville (Table 1). Plant material for isozyme analysis con- sisted of actively growing leaves from rooted cuttings (McArthur et al. 1984) growing in a greenhouse. Isozyme procedures followed Leonard et al. (1981) using a vertical poly- acrylamide preformed gradient gel (Pharma- cia PAA 4/30)°. We also followed Leonard et al. (1981) for isozyme staining of all systems except for shikimate DH (Linhart et al. 1981) and NADP-MDH (Henderson 1966). Eigh- teen isoenzyme systems were examined (Table 2). For heterozygous loci (only peroxi- dase Per, in this study) the “dose” of each allele for polyploid plants was detected visu- ally by observation of staining intensity of two to five replicated gels. Standard genetic distance, D, was calcu- lated using the formula D = —log.I, where i= Vig ele and J with its subscripts is the probability that alleles under consideration are identical (Hartl 1980). Allele frequency differences fo’ 1USDA Forest Service, Intermountain Research Station, Shrub Sciences Laboratory, Provo, Utah 84601. Department of Biological Sciences, Wayne State University, Detroit, Michigan 48202. *The use of trade, firm, or corporation names in this paper is for the information and convenience of the reader. Such use does not constitut “nendorsement or approval by the U.S. Department of Agriculture or of Wayne State University of any product or service to the exclusion of oth usuitable. 157 158 GREAT BASIN NATURALIST Vol. 46, No. 1 TABLE 1. Characteristics of sampled populations of Atriplex canescens. Chromosome Sexual Collection sites Site descriptions numbers" states Little Sahara Sand Dunes, Large rolling sand dunes 2x 2,6 Juab Co., Utah Spanish Fork Canyon, Steep (> 50°) canyon slope. 4x 2.6, Utah Co., Utah Rocky, unconsolidated soil. [2,3] Kingston Canyon, Two subpopulations: 4x 2.6, Piute Co., Utah (1) alluvial fan on canyon [2.6] floor; (2) steep (> 50°) talus slope. Near Grantsville, Valley floor, sedimentary 6x 2 6s Tooele Co., Utah clay soils. [2,3] “Little Sahara Sand Dune population and Grantsville population determined by Stutz et al. (1975 and 1979, respectively). We did counts for several plants each at Spanish Fork Canyon and Kingston Canyon following the methods of Stutz et al. (1975) and confirmed the counts for the other locations. TABLE 2. Isozyme systems tested. Isozyme* Success Peroxidase Yes Glutamate DH Yes Malate DH Yes Shikimate DH Yes Indole Phenol Oxidase Yes NADP - MDH Yes LAP Yes PGI Yes PGM Yes G-6-PDH Yes Amylase Yes Esterase Yes RBPC* Yes ADH No Catalase No Alkaline Phosphatase No Acid Phosphatase No GOT No “See Henderson 1966, Leonard et al. 1981, and Linhart et al. 1981 for full isozyme name descriptions except for RuDPC, which is Ribulose diphosphate j carbohylase. > additional loci may be present but were difficult to resolve and appeared to be monomorphic. “From our total protein analysis—apparently RBPC. the Per, locus were analyzed using the Student- Newman-Keuls multiple range test following analysis of variance procedures (Woolf 1968). RESULTS AND DISCUSSION Of the 13 systems (24 interpretable loci) we were able to analyze, only one locus was poly- morphic (Table 2). The Per, locus had slow, s, and fast, f, alleles. Under our experimental con- ditions (see Leonard et al. 1981) the Per,—s al- lele migrates about 35 mm and the Per, —f allele about 41 mm from the origin. The isozyme data support the Stutz et al. (1975) and Stutz and Sanderson (1979) sugges- tion that A. canescens forms an autopolyploid Genetics 1 locus, polymorphic 1 locus, monomorphic 1 locus, monomorphic 1 locus, monomorphic” 2 loci, monomorphic 2 loci, monomorphic 1 locus, monomorphic 2 loci, monomorphic 1 locus, monomorphic 2 loci, monomorphic 6 loci, monomorphic* 2 loci, monomorphic* 1 locus, monomorphic many loci, not interpretable 1 locus, monomorphic? loci, identical in each locus in each population, suggests genetic homogeneity inherent in auto- polyploid complexes. In a similar study, Oliver and Ruiz Rejon (1980) also found identical isozymes at various polyploid levels in the appar- * ent autoploid Muscari atlanticum (Liliaceae). In | their study, they also found that esterase | isozymes stained more intensely with increasing | polyploid levels. We found an analogous situa- | tion with our Per, locus (54 mm from origin): all © A. canescens plants had Per , monomorphically _ and showed a distinct dosage effect. Diploids stained lightly, tetraploids darker, and hexaploids darkest of all. Visual observation of — Per, from a few plants is enough to ascertain the © population ploidy level. | complex. The preponderance of monomorphic | January 1986 MCARTHUR ET AL.: AUTOPOLYPLOID SHRUB 159 TABLE 3. Genetic distance, D, among the Atriplex canescens populations. Distance values Collection sites 1 2 3 4 5 Little Sahara Sand Dunes (1) — Kingston Canyon slope (2) .003 Kingston Canyon flat (3) .004 Spanish Fork Canyon (4) .008 Grantsville (5) .024 Ordinarily, polyploids have high levels of isozyme heterozygosity (Hamrick et al. 1979, Hunziker and Schaal 1983). That Atriplex canescens does not supports its probable auto- ploid condition. Autoploidy aside, it is an- other example, following Ledig and Conkle (1983), that some woody long-lived perennials have more isozymic homozygosity than previ- ously thought, e.g., Hamrick et al. 1979. ‘Sanderson and Stutz (1984) have recently dis- .covered that diploids have a consistently dif- ferent flavonoid pattern than do tetraploids for both Atriplex canescens and A. confertifo- lia. Their data can be interpreted as meaning the two ploidy levels have different flavonoid physiology. _ Table 3 shows that genetic distance values vamong A. canescens populations are minimal but are in the range (D = nearly 0 to 0.05) that ‘Nei (1976) suggested for local races of a spe- cies. All tetraploid populations had D values of nearly 0. The hexaploid population was set further from the tetraploid populations than was the diploid population. The hexaploid population may, interestingly, have some in- trogression from A. tridentata (Stutz et al. 1979), whereas the tetraploid populations ap- pear to be strict autopolyploids. We point out that our genetic distance values reflect varia- ‘tion at only one of the 13 loci examined. _ We examined two or more clonal ramets for _ isozymes from each plant and obtained identi- cal results in each case. These results are simi- lar to those of Sternberg (1976), who showed that separated clones of Larrea tridentata _ maintained identical isozyme patterns. | There was no statistically significant fre- _ quency difference of the Per, locus among the _ sexual phenotypes of A. canescens at the study | sites (Table 4) even though these phenotypes ! differ in morphological and physiological _ characteristics (McArthur and Freeman 1982; McArthur et al. 1984; McArthur et al. unpub- “) .000 — .000 .000 — O11 O11 .007 — lished data). However, in four of the five popula- tions d's have a higher frequency of the Per,—s allele than do 2’s. The change in frequency of the Per, —s allele from diploid to tetraploid to hexaploid is interesting and warrants further at- tention. Interesting, too, is that A. canescens is dioe- cious or trioecious. These sexual systems have long been considered to have evolved due to inbreeding depression (Grant 1975, Lloyd 1982). Given the monomorphic isozyme data for loci that are normally highly polymorphic (Table 2), it is difficult to see how inbreeding depres- sion, in this case, could be as potent as is re- quired to create the dioecious state. Krohne et al. (1980) discounted inbreeding depression as the driving force in the gynodioecious breeding system of Plantago lanceolata. ACKNOWLEDGMENTS The study was facilitated by National Sci- ence Foundation Grant DEB 81-11010 and Pittman Robertson Wildlife Restoration Pro- ject W-82-R. We thank Drs. F. T. Ledig, Y. B. Linhart, J. B. Mitton, and H. C. Stutz for constructive criticism of earlier versions of this manuscript. LITERATURE CITED FREEMAN, D. C., L. G. KLICKHOFF, AND K. T. HARPER. 1976. Differential resource utilization by the sexes of dioecious plants. Science 193:597—599. Grant, V. 1975. Genetics of flowering plants. Columbia University Press, New York. 514 pp. Hamrick, J. L., Y. B. LInHart, AND J. B. Mitron. 1979. Relationships between life history characteristics and electrophoretically detectable genetic varia- tion in plants. Ann. Rev. Ecol. Sys. 10: 173-200. Hart_, D. L. 1980. Principles of population genetics. Sinauer Associates, Inc., Sunderland, Massachu- setts. 488 pp. HENDERSON, N. S. 1966. Isozymes and genetic control of NADP-Malate Dehydrogenase in mice. Arch. Biochem. Biophys. 117:28—33. 160 GREAT BASIN NATURALIST Vol. 46, No. 1 TABLE 4. Frequency of the Per, —s allele. Population Mean + se Little Sahara Sand Dunes (2x) .921 + .057A* Kingston Canyon slope (4x) .647+ .071B Kingston Canyon flat (4x) .643 + .081B Spanish Fork Canyon (4x) .583 + .052B Grantsville (6x) .258 + .040C Frequency 2 3 [256] .850 1.00 — .643 .667 .625 625 .667 — .036 .643 oll .286 .238 .250 “Different letters after mean frequency values indicate significantly different (p < .01) means. Hl HUNZIKER, J. H., AND B. A. SCHAAL. 1983. Isozyme varia- tion in diploid tropical and octoploid subtropical- temperate species of Bulnesia. J. Hered. 74: 358-360. KROHNE, D. T., I. BAKER, AND H. G. BAKER. 1980. The maintenance of the gynodioecious breeding sys- tem in Plantago lanceolata L. Amer. Midland Nat. 103:269-279. LEDIG, F.T., AND M. T. CONKLE. 1983. Gene diversity and genetic structure in a narrow endemic, Torrey Pine (Pinus torreyana Parry ex Carr.) Evolution 37: 79-85. LEONARD, R. L., E. D. MCARTHUR, D. J. WEBER, AND B. W. Woop. 1981. Electrophoresis of isoenzymes of 16 western shrubs: technique development. Great Basin Nat. 41:377-388. LINHART, Y. B., M. L. DAvIs, AND J. B. MITTON. 1981. Ge- netic control of allozymes of shikimate dehydroge- nase in ponderosa pine. Biochem. Genetics 19: 641-646. LLoyp, D.C. 1982. Selection of combined versus separate sexes in seed plants. Amer. Nat. 120:571—585. McArTHUR, E. D. 1977. Environmentally induced changes of sex expression in Atriplex canescens. Heredity 38:97-103. MCARTHUR, E. D., A.C. BLAUER, AND G. L. NOLLER. 1984. Propagation of fourwing saltbush (Atriplex canes- cens [Pursh] Nutt.) by stem cuttings. Pages 261-264 in A. R. Tiedemann, E. D. McArthur, H. C. Stutz, K. L. Johnson, and R. Stevens (compil- ers), The biology of Atriplex and _ related chenopods, Proceedings, Wildland Shrub Sympo- sium. USDA Forest Service Gen. Tech. Rep. INT-172. 309 pp. MCARTHUR, E. D., AND D.C. FREEMAN. 1982. Sex expres- sion in Atriplex canescens: genetics and environ- ment. Bot. Gaz. 143:476—482. 2,3] ramets from Kingston Canyon flat were not available for study at the same time as the other populations; consequently they were not included. McArTHUR, E. D., R. STEVENS, AND A. C. BLAUER. 1983. Growth performance comparisons among 18 acces- sions of fourwing saltbush (Atriplex canescens ) at two sites in Central Utah. J. Range Manage. 36:78-81. NEI, M. 1976. Mathematical models of speciation and genetic distance. Pages 723-765 in S. Karlin and E. Nevo, eds., Population genetics and ecology. Academic Press, New York. 832 pp. OLIVER, J. L., AND M. Ruiz REJON. 1980. The relation between isozymes and ploidy level: its application to biogeo- graphical studies of Muscari atlanticum (Liliaceae). | Taxon 29:27-32. | SANDERSON, S. C., AND H. C. Stutz. 1984. Flavonoids of diploid and polyploid Atriplex confertifolia. Pages 34-38 in A. R. Tiedemann, E. D. McArthur, H. C. Stutz, K. L. Johnson, and R. Stevens (compilers), The biolegy of Atriplex and related chenopods, Proceedings of Wild- } land Shrub Symposium: USDA Forest Service Gen. | Tech. Rep. INT-172. 309 pp. STERNBERG, L. 1976. Growth forms of Larrea tridentata. Madrono 23:408—417. | Stutz, H.C.,J.M. MELBY, ANDG. K. LIvINGsTON. 1975. Evolu- | tionary studies of Atriplex: a relic gigas diploid popu- | lation of Atriplex canescens. Amer. J. Bot. | 62:236-245. | StuTz, H. C., C. L. Pore, AND S. C. SANDERSON. 1979. Evolu- |, tionary studies of Atriplex: adaptative products of the } natural hybrid 6n A. tridentata x 4n A. canescens. |. Amer. J. Bot. 66:1181—1193. | Stutz, H. C., AND S. C. SANDERSON. 1979. The role of poly- ploidy in the evolution of Atriplex canescens. Pages } \ 615-621 in J. R. Goodin and D. K. Northington, eds., }), Arid land plant resources, Proceedings of Interna- | tional Arid Lands Conference, Texas Tech Univer- ) sity, Lubbock. 724 pp. Woo LF, C. M. 1968. Principles of biometry. D. Van Nostrand i: Company, Princeton, New Jersey. 359 pp. WINTER NUTRITIVE CONTENT OF BLACK SAGEBRUSH (ARTEMISIA NOVA) GROWN IN A UNIFORM GARDEN! Barbara Behan? and Bruce L. Welch® ABSTRACT.— Winter crude protein content, in vitro digestibility, and productivity were determined for seven accessions of black sagebrush (Artemisia nova) grown in a uniform garden. No significant differences were detected among the accessions for any of these attributes. Mean crude protein was 6.8% of dry matter. Accessional range was from 5.8% to 7.3%. Mean in vitro digestion was 54.8% of dry matter; accessional range, 51.9% to 57.2%. Mean current year s growth (a measurement of productivity) was 4.3 cm; accessional range, from 3.7 to 5.1 em. In comparison to other winter forages, black sagebrush ranks high for winter levels of crude protein and very high in winter digestible dry matter but low in productivity. Protein and energy-producing compounds are two of three nutrients commonly listed as being deficient in the winter diet of ruminants on native ranges (Dietz 1965, Halls 1970, Nagy and Wallmo 1971, Welch and McArthur 1979a). Plants that retain significant amounts of green leaves during the winter usually con- tain higher levels of protein and are more digestible than those that shed their leaves (Ensminger and Olentine 1978, Welch 1983). We have reported significant differential pref- erence of wintering mule deer (Odocoileus hemionus hemionus ) among seven accessions of black sagebrush (Artemisia nova) (Behan sand Welch 1985). Significant variation in win- ter nutrient levels among accessions of a re- ‘lated species, big sagebrush (A. tridentata), grown in a uniform garden has been reported (McArthur and Welch 1982, Welch and McArthur 1979b, Welch and Pederson 1981), but there has been little information until now concerning variation in winter nutrient levels -among accessions of black sagebrush. We un- dertook this study to determine the winter nutritive content of seven accessions of black sagebrush grown in a uniform garden. MATERIALS AND METHODS ~ On a uniform shrub garden located at the ‘Gordon Creek Wildlife Management Area’ aear Helper, Utah, seven accessions were se- lected to determine in vitro digestibility, pro- ductivity, and levels of crude protein. The accessions had been transplanted as seedlings from various native source locations (Table 1). Within each accession, seven individual plants were randomly selected to furnish the vegetative tissue needed for testing. Because of heavy grazing on twig tips by wintering mule deer, composite sampling had to be used for the Spring Valley and Wingate Mesa accessions. Only twigs with terminal buds and leaves were collected from the plants. Sam- pling occurred on 3 December 1982. Vegetative samples (current years growth of stems and leaves) were collected from each plant, placed in separate paper bags and frozen on site with dry ice. Individual samples were placed in separate plastic bags tied and sealed in a second bag. The double-bagged samples were stored at —35 C until ground. The samples were ground while submerged in liquid nitrogen in a motorized mortar and pestle. Th was done to prevent loss of volatile su stances such as monoterpenoids that may suppress cellulolytic microorgan- isms and to aid in grinding the samples (1/2 mm, Hobbs et al. 1985). Next the ground samples were stored in airtight containers at —35 C until needed for protein determination or digestion trials. Crude protein levels were determined by the Kjeldahl method (Association of Official this article was written by a United States government employee and a federal cooperator on official time and is therefore in the public domain. Department of Range Science, Colorado State University, Fort Collins, Colorado 80521. 3USDA Forest Service Intermountain Research Station, Shrub Sciences Laboratory, 735 North 500 East, Provo, Utah 84601. f | *The shrub garden at the Gordon Creek Wildlife Management Area is cooperatively maintained by the Utah Division of Wildlife Resources (Wildlife Restoration funds W-82-R, job 1) and the Intermountain Research Station. | 161 162 GREAT BASIN NATURALIST Vol. 46, No. 1 TABLE 1. Locations by county, state, and landmark, or town where each of seven accessions of black sagebrush (Artemisia nova) were collected. Accession Pine Valley Ridge Manti Black Mountain Spring Valley Dove Creek Wingate Mesa Fremont Junction TABLE 2. Mean winter crude protein, in vitro digesti- bility, and productivity for seven accessions of black sage- brush (Artemisia nova) grown in a uniform garden. Protein and digestibility data are expressed as percent of dry matter. Productivity data are expressed in length of leader growth in centimeters. Means for all measure- ments were found not to be significantly different. Crude Leader protein Digestibility | growth Accession (%) (%) (cm) Spring Valley’ 5.8 54.6 BY Manti 6.5 57.2 4.1 Black Mountain 6.9 59.9 oer Fremont Junction 6.9 55.6 5.1 Dove Creek Doll 59.6 4.6 Wingate Mesa’ To 51.9 3.9 Pine Valley Ridge U8 53.5 4.5 ‘Samples of the Spring Valley and Wingate Mesa accessions were com- posited and not included in the analysis of variance. Analytical Chemists 1980). Crude protein data are expressed as a percentage of dry mat- ter. We used the in vitro digestion procedure as outlined by Pearson (1970), except 1.0 g of fresh tissue was placed in digestion tubes. Rumen inoculum was collected from a slaugh- terhouse steer fed a ration of alfalfa hay and corn. Welch et al. (1983) have reported that inoculum source has little effect on the rank- ing of the digestibility of range forages. Re- sults of the digestion trials are expressed as a percentage of dry matter digested. Percentage values for the crude protein were transformed (arcsin) before performing a completely random analysis of variance. Per- centages for in vitro digestion were in the range that did not require transformation prior to the analysis of variance. For signifi- cant F ratios, Student-Newman multiple range test (P < 0.05) was used to determine differences among treatment means. Because of the composite sampling, the Spring Valley and Wingate Mesa accessions were not in- cluded in the analysis of variance for crude protein and in vitro digestion. Location Millard, Utah (Desert Experimental Range) Sanpete, Utah (Manti) Sevier, Utah (Salina) White Pine, Nev. (Jct. US-93, 6, and 50) Dolores, Colo. (Dove Creek) San Juan County, Utah (Fry Canyon) Sevier, Utah (Fremont Jct.) Prior to deer use, the plants were used to determine current year’s growth, an indicator of production. Current year’s growth was de- termined by measuring the annual leader | length of 15 leaders per plant. Leader lengths — were measured to the nearest centimeter — from the terminal leaf bud scars to the tip of | the current terminal leaves. The leaders were i selected at random over the entire crown of | the plants. A plant mean was calculated from © the 15 measurements. Current year growth | data were expressed as centimeters and were | | statistically analyzed as described for crude | protein and in vitro digestion. ( RESULTS AND DISCUSSION are given in Table 2. Mean winter crude pro- | tein content for black sagebrush was 6.8%. — Accessional range was from 5.8% to 7.3%. The | Pine Valley Ridge accession contained the — highest amount of crude protein at 7.3%. No. significant differences among the accessiqusa | were detected. Our crude protein levels are ~oneiderali less than the 11.7% level reported by Sheehy i) (1975) and less than the 8.5% reported by the National Academy of Sciences (1964). We are’ not sure that the latter figure was for the win-. i ter period. Averaging the three studies, black! sagebrush winter crude protein content ): would be about 9.0% of dry matter. A winter || crude protein level of 9.0% ranks high amon { winter range forages (Table 3). Winter in vitro digestibility of the sever. i accessions of black sagebrush is given in Table — 3. Mean in vitro digestibility was 54.8% of dry i matter digested. Accessional range was fron }; 51.9% to 57.2%. The Manti accession was the )) most readily digested at 57.2%. No significan / differences were detected among the acces )) Results of the crude protein determinations | | y! i { January 1986 BEHAN, WELCH: BLACK SAGEBRUSH 163 TABLE 3. Mean winter crude protein content (percentage of dry matter) of some range plants. Plant Crude protein Range Reference Agropyron desertorum 15.0 19* (green-regrowth) Artemisia tridentata 11.4 (9.9-14. 2) 1, 2, 3, 4, 6, 8, 12, 16, 19 Cercocarpus ledifolius 10.1 (9.6—10.6) Seal Atriplex canescens 9.6 ll Artemisia nova 9.0 (6.9-11.7) 12, 20, 17 Prunus virginiana 8.7 (7.6—9.9) 3, 5, 10, 15 Cowania mexicana 8.6 (8.48.8) 5, 13 Purshia glandulosa 8.5 (8.0-9.0) 3, 13 Juniperus scopulorum 8.4 l Populus tremuloides 7.8 (6.5—9.5) Sy IO, Chrysothamnus nauseous 7.8 (5.9-7.8) 1, 10 Cercocarpus montanus 7.8 (7.2—8.4) Ly &) Purshia tridentata 7.8 (6.7—9.1) 1, 3, 4, 7, 8, 10, 13 Atriplex confertifolia eh 9 Juniperus osteosperma 6.6 (5.9-7.6) Son Chrysothamnus viscidiflorus 5.9 19 Amelanchier alnifolia 5.9 (5.5-6.2) 3, 10 Rosa woodsii 5.8 (5.4—6. 1) 15, 18 Quercus gambelii 5.3 (5.1—5.4) 5, 16 Fallugia paradoxa 4.8 13 Amelanchier utahensis 4.8 15 Agropyron desertorum 3.9 10 Native grass 3.6 3 Stipa comata 3.5 (2.9-4.0) 9, 10 Oryzopsis hymenoides 3.0 (2.5-3.5) 10, 17 *Reference: 1. Dietz et al. 1962 11. Welch and Monsen 1981 2. Welch and McArthur 1979b 12. Sheehy 1975 3. Tueller 1979 13. Welch et al. 1983a 4. Bissell et al. 1955 14. Welch and Monsen 1984 5. Smith 1957 15. Dietz 1972 6. Smith 1950 16. Kufeld et al. 1981 7. Smith 1952 17. National Academy of Sciences 1964 8. Medin and Anderson 1979 (Data converted to dry matter basis) 18. Welch and Andrus 1977 9. National Academy of Sciences 1975 19. Urness et al. 1983 10. National Academy of Sciences 1958 20. This study sions. Our mean in vitro digestibility com- pares favorably with reports by Sheehy (1975) at 53.1% and with Welch et al. (1983b), also at 53.1%. Mean in vitro dry matter digestibility for the three studies is 53.7%. Black sage- brush ranks very high in digestibility among winter range forages (Table 4). Ammann et al. | (1973) estimated that dry-matter digestibility of 50% would provide sufficient energy for maintenance. Mean current years growth was 4.3 cm, accessional range, 3.7 to5.1 cm. The Fremont Junction was the most productive at 5.1 cm (Table 2). No significant differences among 'the accessions were detected. Black sage- brush is not as productive as other winter range forages such as big sagebrush (Artemisia \tridentata), antelope bitterbrush (Purshia tri- dentata), fourwing saltbush (Atriplex canes- ‘cens), and true mountain mahogany (Cerco- carpus montanus) (McArthur and Welch 1982, McArthur et al. 1983). Black sagebrush is adaptable to sites where the more produc- tive species do not grow. Black sagebrush ranks high in winter levels of crude protein and very high in digestible dry matter in comparison to other forages. Phosphorus content is probably high also (Na- tional Academy of Sciences 1964). From a qualitative point of view winter nutrient con- tent of black sagebrush is exceeded only by big sagebrush (Tables 3 and 4; Welch 1983). Lack of significant differences among the seven accessions for the three characters tested suggests that breeding and selection schemes stressing improvement of these at- tributes would be fruitless. We have reported earlier that wintering mule deer significantly preferred the Pine Valley Ridge accessions over the other accessions tested (Behan and GREAT BASIN NATURALIST Vol. 46, No. 1 TABLE 4. Mean winter in vitro digestion of some range plants. Data are expressed as a percentage of dry matter 164 digested. Dry matter Plant digested Artemisia tridentata 57.4 Artemisia spinescens 57.0 Artemisia nova 53.7 Sporobolus cryptandrus 53.2 Agropyron smithii 50.2 Oryzopsis hymenoides 50.0 Cercocarpus ledifolius 49.1 Rosa eglanteria (hips) 49.1 Hilaria jamesii 48.2 Stipa comata 48.1 Agropyron spicatum 45.5 Ceratoides lanata 44.7 Chrysothamnus nauseous 44.4 Atriplex confertifolia 43.4 Amelanchier utahensis 41.0 Prunus virginiana 38.8 Atriplex canescens 38.3 Cowania mexicana 37.6 Purshia glandulosa SL} Amelanchier alnifolia 34.6 Kochia prostrata 32.2 Fallugia paradoxa 29.8 Quercus gambelii 28.1 Purshia tridentata 25.4 Cercocarpus montanus 24.3 *References: 1. Dietz 1972 2. Kufeld et al. 1981 3. Sheehy 1975 4. Urness et al. 1977 5. Wallmo et al. 1977 6. Welch and Pederson 1981 7. Pederson and Welch 1982 Welch 1985). Also, Clary and Beale (1983) noted that pronghorn and domestic sheep both pre- ferred black sagebrush that grows on the Desert Experimental Range in Pine Valley. This is the same kind of black sagebrush as our collection from the Pine Valley Ridge (just north of the Desert Experimental Range). We will be testing the adaptation range of the Pine Valley Ridge accession in preparation for releasing it through the Soil Conservation Service’s plant material program as a superior cultivar of black sagebrush for improving winter ranges for domestic sheep, pronghorn, and mule deer. LITERATURE CITED AMMANN, A. P., R. L. Cowan, C. L. MOTHERSHEAD, AND B. R. BAUMGARDT. 1973. Dry matter and energy intake rela- tion to digestibility in white-tailed deer. J. Wildl. Manage. 37:195-201. ASSOCIATION OF OFFICIAL ANALYTICAL CHEMISTS. 1980. Official methods of analysis. W. Horwitz, ed., 13th Ed. As- soc. Off. Anal. Chem., Washington, D.C. Range Reference (49.9-67.0) 2, 3, 4, 5, 6, 7, 10* 8 (53. 1-54.0) 3,8, 14 8 10 (45.7-54.2) 8, 10 (44.7-53.5) 4,6 6 8 10 10 8 10 8 1 (26.3-51.3) 1 Wi 9 12 12 10 13 12 2 (19.8-30.0) 4, 6, 10, 12 (20.0—28.5) 4,6 8. Welch et al. 1983b e 9. Welch and Monsen 1984 10. Ward 1971 11. Uresk et al. 1975 12. Welch et al. 1983a 13. Welch and Davis 1984 14. This study BEHAN, B., AND B. L. WELCH. 1985. Black sagebrush: | mule deer winter preference and monoterpenoid | content. J. Range Manage. 38:276—277. BISSELL, H. D., B. HARRIS, H. STRONG, AND F. JAMES. 1955. | Digestibility of certain natural and artificial foods — eaten by deer in California. California Fish and — Game 41:57-78. Crary, W. P., AND D. M. BEALE. 1983. Pronghorn reac- | tions to winter sheep grazing, plant communities, _ and topography in the Great Basin. J. Range Man- | age. 36:749-752. Dierrz, D. R. 1965. Deer nutrition research in range man- | agement. Trans. North Amer. Wildl. and Nat. — Resour. Conf. 30:274—285. Intermountain Forest and Range Exp. Sta., Og-. den, Utah. DIETZ, Chemical composition and digestibility by mule deer of selected forage species, Cache La Poudre Range, Colorado. Colorado Game and Fish Dep. Tech. Publ. 14. 89 pp. | . 1972. Nutritive value of shrubs. Pages 289-302 in» C. M. McKell, J. P. Blaisdell, and J. R. Goodin, eds., Wildland shrubs—their biology and utiliza- | tion. USDA For. Serv. Gen. Tech. Rep. INT-1. ; D. R, RH Upatt, anp L. E. YeaGer. 1962. | January 1986 ENSMINGER, M. E., AND C. G. OLENTINE, JR. 1978. Page 147 in Feeds and nutrition. Abridged. Ensminger Publ. Co., Clovis, California. Hats, L. K. 1970. Nutrient requirement of livestock and game. Pages 10-18 in H. A. Paulsen, Jr., E. H. Reid, and K. W. Parker, eds., Range and wildlife habitat evaluation—a research symposium. USDA _ For. Serv. Misc. Publ. 1147. | Hosss, N. T., B. L. WELCH, AND T. E. REMINGTON. 1985. Ef fects of big sagebrush on in vitro digestion of grass cell wall. Pages 186-189 in Proceedings—symposium on the biology of Artemisia and Chrysothamnus. USDA, For. Serv. Gen. Tech. Rep. INT-200. Intermountain Research Station, Ogden, Utah. 401 pp. | KUFELD, R. C., M. STEVENS, AND D. C. BOWDEN. 1981. Winter variation in nutrient and fiber content and in vitro digestibility of Gambel oak (Quercus gambelii) and big sagebrush (Artemisia tridentata) from diversified sites in Colorado. J. Range Manage. 34:149-151. . McArtuur, E. D., R. STEVENS, AND A. C. BLAUER. 1983. Growth performance comparisons among 18 acces- sions of fourwing saltbush (Atriplex canescens ).at two sites in central Utah. J. Range Manage. 36: 78-81. MCARTHUR, E. D., AND B. L. WELCH. 1982. Growth rate differ- ences among big sagebrush (Artemisia tridentata ) ac- cessions and subspecies. J. Range Manage. 35: 396-401. MepIN, D. E., AND A. E. ANDERSON. 1979. Modeling the dy- namics of a Colorado mule deer population. Wildl. Monogr. 68. 77 pp. Nacy, J. G., AND O. C. WALLMO. 1971. Deer nutrition prob- lems in the USA. Proc. World Exhib. Hunting, Int. Sci. Conf. Game Manage., Sect. 1:59-68. University Press, Sopron, Hungary. NATIONAL ACADEMY OF SCIENCES. 1958. Composition of cereal grains and forages. Natl. Res. Counc. Publ. 585. Washington, D.C. —_. 1964. Nutrient requirements of domestic animals. | No. 5. Nutrient requirements of sheep. 3d Ed. Natl. Res. Counc. Publ. 1193. Washington, D.C. ———. 1975. Nutrient requirements of domestic animals. No. 5. Nutrient requirements of sheep. 5th Ed. Natl. Res. Counc. Publ. 74-899. Washington, D.C. | PEARSON, H. A. 1970. Digestibility trials: in vitro techniques. Pages 85-90 in H. A. Paulsen, E. H. Reid, and K. W. Parker, eds., Range and wildlife habitat evaluation— a range snmpOstE. Publ. 1147. USDA For. Serv., Washington, D.C. | PEDERSON, J.C., AND B. L. WELCH. 1982. Effects of monoter- penoid exposure on ability ofrumen inocula to digest a set of forages. J. Range Manage. 35: 500-502. | SHEEHY, D. P. 1975. Relative palatability of seven Artemisia taxa to mule deer and sheep. Unpublished thesis, Oregon State University, Corvallis. Situ, A. D. 1950. Sagebrush as winter food for mule deer. J. Wildl. Manage. 14:285-289. ——. 1952. Digestibility of some native forages for mule deer. J. Wildl. Manage. 16:309-312. -—— . 1957. Nutritive value of some browse plants in winter. J. Range Manage. 10:162-164. TUELLER, P. T. 1979. Food habits and nutrition of mule deer on Nevada ranges. University of Nevada, Reno. | BEHAN, WELCH: BLACK SAGEBRUSH 165 Uresk, D. W., AND H. E. MESSNER. 1975. Constituents in in vitro solution contribute differently to dry matter di- gestibility of deer food species. J. Range Manage. 28:419—421. Urness, P. J., D. D. AusTIN, AND L. C. Fiero. 1983. Nutritional value of crested wheatgrass for wintering mule deer. J. Range Manage. 36:225—226. Urness, P. J., A. D. Smiri, AND R. K. WATKINS. 1977. Compari- son of in vivo and in vitro dry matter digestibility of mule deer forages. J. Range Manage. 30: 119-121. WALLMO, O. C., L. H. CARPENTER, W. L. REGELIN, R. B. GILL, AND D. L. BAKER. 1977. Evaluation of deer habitat on a nutritional basis. J. Range Manage. 30:122-127. Warp, A. L. 1971. In vitro digestibility of elk winter forage in southern Wyoming. J. Wildl. Manage. 35:681-688. WELCH, B. L. 1983. Improving the nutritive value of winter range forage. Pages 158-164 in S. B. Monsen, and N. Shaw, compilers, Managing intermountain range- lands—improvement of range and wildlife habitats. Gen. Tech. Rep. INT-157. USDA For. Serv., In- termt. For. and Range Exp. Sta., Ogden, Utah. WELCH, B. L., AND D. ANDRUS. 1977. Rose hips—a possible high-energy food for wintering mule deer? Res. Note INT-221. USDA For. Serv., Intermt. For. and Range Exp. Sta., Ogden, Utah. WELCH, B. L., AND J. N. Davis. 1984. In vitro digestibility of Kochia prostrata (L.) Schrad. Great Basin Nat. 44:296-298. WELCH, B. L., AND E. D. McArTHUR. 1979a. Feasibility of improving big sagebrush (Artemisia tridentata) for use on mule deer winter ranges. Pages 451—473 in Arid land plant resources. Texas Tech University, Lubbock. —___.. 1979b. Variation in winter levels of crude protein among Artemisia tridentata subspecies grown in a uniform garden. J. Range Manage. 32:467—469. WELCH, B. L., AND S. B. MONSEN. 1981. Winter crude protein among accessions of fourwing saltbush grown in a uniform garden. Great Basin Nat. 41: 343-346. ____. 1984. Winter nutritive value of accessions of fourwing saltbush (Atriplex canescens ) grown in a uniform gar- den. Pages 138-149 in A. R. Tiedemann, E. D. McArthur, H. C. Stutz, R. Stevens, and K. L. John- son, compilers, Proceedings—symposium on the bi- ology of Atriplex and related chenopods. Gen. Tech. Rep. INT-172. USDA For. Serv., Intermt. For. and Range Exp. Sta., Ogden, Utah. WELCH, B. L., S. B. MONSEN, AND N. L. SHAW. 1983a. Nutritive value of antelope and desert bitterbrush, stansbury cliffrose, and apache-plume. Pages 173-185 in A. R. Tiedemann, and K. L. Johnson, compilers, Proceed- ings—research and management of bitterbrush and cliffrose in western North America. Gen. Tech. Rep. INT-152. USDA For. Serv., Intermt. For. and Range Exp. Sta., Ogden, Utah. WELCH, B. L., AND J. C. PEDERSON. 1981. In vitro digestibility among accessions of big sagebrush by wild mule deer and its relationship to monoterpenoid content. J. Range Manage. 34:497—500. WELCH, B. L., J.C. PEDERSON, AND W. P. Ciary. 1983b. Ability of different rumen inocula to digest range forages. J. Wildl. Manage. 47:873-877. WINTER FOOD HABITS OF THE PINE MARTEN IN COLORADO Christine C. Gordon! ABSTRACT. —Eighteen pine marten (Martes americana) stomachs and 29 intestinal tracts, collected between October 1983 and March 1984 from northern Colorado, were examined for food items. Voles (Microtus spp.) occurred most frequently, followed by shrews (Sorex spp.), insects, and vegetative matter. The pine marten (Martes americana) in- habits mature stands of coniferous forests of North America (Lensink et al. 1955, Marshall 1951). Although the pine marten is an impor- tant furbearer in many states, few studies have specifically examined winter food habits (Hargis and McCullough 1984, Zielinski et al. 1983, Lensink et al. 1955, Cowan and Mackay 1950). Winter is a critical period when many food items are unavailable and prey popula- tions reach yearly lows. Lensink et al. (1955) reported that changes in the abundance or availability of food may be reflected in move- ments, productivity, and choice of habitat. The objectives of this study were to identify winter foods and their frequencies in the diet with respect to availability. STUDY AREA AND METHODS The study area is in the Roosevelt National Forest 29 km west of Rustic, Larimer County, Colorado. Elevation ranged from 3168 to 3780 m. The trap sites were dominated by Engel- mann spruce (Picea engelmannii) and sub- alpine fir (Abies lasiocarpa). The main under- story vegetation consisted of vaccinium (Vaccinium spp.). The study was conducted feathers. Hair impressions were used when — no other fragments were identifiable (Moore | et al. 1974). Prey items were identified to | genus whenever possible. Percent occur- — rence (number of occurrences of a prey item/ _ total number of stomachs or intestines x 100) was determined for each prey item. RESULTS Twelve different food items were identified | in stomach and intestinal samples (Table 1). | Mammals composed the highest percentage | of items found. Microtus spp. were the most important food item, occurring in 83% of the samples. Although insects and vegetative ma- | terials were represented in 17% of the sam- | ples, actual amounts per scat were negligible. | Birds, squirrels (Tamiasciurus hudsonicus), cervids, and fish were present in 11% of the diets. Mustelids constituted 10%; however | this is believed to have been ingested inciden- _ tally. Both snowshoe hare (Lepus americana) and beaver (Castor canadensis) items oc- | curred in 7%. DISCUSSION { (| Microtus spp. were also the most important’ food item in previous studies (Douglass et al. | 1983, Zielinski et al. 1983, Soutiere 1979, — from October 1983 to March 1984. The winter was severe with greater snowfall and colder temperatures than average. Thirty-two martens were collected using conibear and leghold traps. Food samples were taken from both the stomach and intes- tine. Samples were analyzed using the proce- dure described by Johnson and Hanson (1977). Reference collections were used to identify diagnostic bones, teeth, hair, or ‘Department of Fishery and Wildlife Biology, Colorado State University, Fort Collins, Colorado 80523. 166 Weckwerth and Hawley 1962, Murie 1961, Quick 1955, Lensink et al. 1955, Cowan and: | Mackay 1950). The results of small mammal, trapping have shown microtines to be the’ most abundant small mammals in the aree | during winter (Palmer, progress report). Al- though shrews were quite abundant on the | January 1986 TABLE 1. Stomach and intestinal contents from 32 Col- orado pine marten, October 1983—March 1984. Percent Percent occurrence occurrence in stomach _ in intestine (N=18) (N=29) Mammals Microtus spp." 83 79 Sorex spp. 39 45 Tamiasciurus hudsonicus ll 7 Lepus americana 5 7 Castor canadensis 5 5 Cervidae 11 3 Mustelidae 5 10 | Birds® 11 7 Fish 11 3 Vegetation 11 17 | Insects 17 14 Unknown 0 3 "Includes M. pennsylvanicus, M. longicaudus, M. montanus, Clethriono- mys gapperi, Phenacomys intermedius. ‘Includes S. hoyi, S. vagrans, S. cinereus, S. palustris, Cryptotis parva. ‘Includes Turdis migratorius, Junco hyemalis caniceps, and two samples keyed only to Passeriformes. » study site, the high prevalence in marten diets was unusual (Palmer, progress report). Gener- GORDON: PINE MARTIN Foop Hasits 167 Marten food habits are affected by availabil- ity, preference, and availability of alternate prey (Weckwerth and Hawley 1962). Zielinski et al. (1983) reported that seasonal activity of the marten appeared to be synchronized with the activity of its prey. Marten activity varies seasonally. They are crepuscular during the summer, but the approach of winter forces them to become nocturnal as alternate food sources become less available (Zielinski et al. 1983). The availability of invertebrates, fruits, birds, and diurnal mammals increases during warm spring and summer months. During winter, insects decrease in abundance, mi- grant bird populations have gone, and fruit no longer remains on bushes (Weckwerth and Hawley 1962). Colder temperatures and deep snows force many mammals underground; therefore, few prey items are still available to the martens. Nocturnal activity increases the martens interactions with prey species such as voles, shrews, or snowshoe hares. It is therefore not surprising that voles and shrews were taken most frequently. It appears that food items on the study area were taken in ally shrews are not taken, although they may be _ quite abundant, because they have musk glands \ that emit a strong odor (Lensink et al. 1955, proportion to their abundance and availabil- ity. This study, like others, shows that martens at this particular study area are op- Cowan and Mackay 1950). The high percentage of shrews may be directly related to the severity of the winter and prey abundance. Colder win- ters and deeper snows reduce aboveground ac- tivity of mammals (Ewer 1973), and the marten \ minimizes activity by taking the most abundant food source. Sorex spp. were the second most abundant food source available during winter. Birds occurred infrequently, suggesting that ‘they were taken only when chance permitted. /Similar results were reported by Weckwerth »and Hawley (1962), Lensink et al. (1955), and | Cowan and Mackay (1950). A relatively low per- centage of the samples contained pine squirrel. _ Although no index of abundance was available ‘for comparison, pine squirrels are believed to _ exhibit periods of torpor during winter and are virtually unavailable to the marten (Cowan and Mackay 1950). The cervid noted was most likely _ carrion. The fish and beaver were probably ob- tained from other trappers’ bait or discarded - portions of carcasses left in the area. Snowshoe vhares, though occurring on the study area, were ‘seldom taken by martens. Other studies have ‘reported similar results (Douglass et al. 1983, Lensink et al. 1955, Cowan and Mackay 1950). portunistic in nature during critical periods in order to optimize net energy gained (Quick 1955, Lensink et al. 1955, Weckwerth and Hawley 1962). ACKNOWLEDGMENTS I express my gratitude to M. W. Schehrer and G. E. Graves for their help and encour- agement. I also thank Terry Foppe and the Colorado State University Composition Labo- ratory for use of facilities and Charles Chase III, curator of ornithology, Denver Museum of Natural History. LITERATURE CITED Cowan, L. M., AND R. H. Mackay. 1950. Food habits of the marten (Martes americana) in the Rocky Moun- tain region of Canada. Can. Field-Nat. 64: 100-104. Douc ass, R. J., L. G. FISHER, AND M. Mair. 1983. Habi- tat selection and food habits of marten, Martes americana, in the Northwest territories. Can. Field-Nat. 97:71—74. Ewer, R. F. 1973. The carnivores. Cornell University Press, Ithaca, New York. 494 pp. 168 Harcis, C. D., AND D. R. MCCuLLOUGH. 1984. Winter diet and habitat selection of marten in Yosemite National Park. J. Wildl. Manage. 48:140-146. JOHNSON, M. K., AND R. M. HANson. 1977. Comparison of point frame and hand separation of coyote scats. J. Wildl. Manage. 41:319. KOEHLER, G. M., AND M. G. HORNOCKER. 1977. Fire ef- fects on marten habitat in the Selway-Bitteroot wilderness. J. Wildl. Manage. 41:500—505. LENSINK, C. J., R. D. ScooG, AND J. L. BUCKLEY. 1955. Food habits of marten in interior Alaska and their significance. J. Wildl. Manage. 19:364—367. MARSHALL, N. H. 1951. Pine marten as a forest product. J. Forestry 49:899—905. Mookrg, T. D., L. E. SPENCE, C. E. DUGNOLLE, AND W. G. HEpwortH. 1974. Identification of the dorsal guard hairs of some mammals of Wyoming. Wyo- ming Game and Fish Dept. Bull. 14. 117 pp. GREAT BASIN NATURALIST Vol. 46, No. 1 Murig, A. 1961. Some food habits of the marten. J. Mam- | mal. 42:516—521. PALMER, D. A. 1984. Investigation of the home range, territory size, and daily and seasonal movements | of two congeneric small owls. Colorado Div. Wildl. Progress Report. 7 pp. Quick, H. F. 1955. Food habits of marten (Martes ameri- cana) in northern British Columbia. Can. Field- Nat. 69:144-148. SOUTIERE, E. C. 1979. Effects of timber harvesting on marten in Maine. J. Wildl. Manage. 43:850—860. WECKWERTH, R. P., AND V. D. HAWLEY. 1962. Marten food | habits and population fluctuations in Montana. J. Wildl. Manage. 26:55—74. ZIELINSKI, W. J., W. D. SPENCER, AND R. H. BARRETT. 1983. Relationship between food habits and activity pat- | terns of pine martens. J. Mammal. 64:387—396. FOLIAGE AGE AS A FACTOR IN FOOD UTILIZATION BY THE WESTERN SPRUCE BUDWORM, CHORISTONEURA OCCIDENTALIS Elizabeth A. Blake' and Michael R. Wagner! ApsTRACT.—The influence of current year foliage age on food consumption and utilization by the western spruce | budworm, Choristoneura occidentalis (Lepidoptera:Tortricidae). was examined. Larvae were fed immature foliage of | Douglas-fir (Pseudotsuga menziesii var. glauca), Engelmann spruce (Picea engelmannii), and corkbark fir (Abies | lasiocarpa var. arizonica) in June and August of 1981 and Douglas fir in June and July of 1982. All larvae feeding on e early season (June) foliage reached maturity. Larvae feeding on middle (July) and late (August) season foliage died | before reaching pupation. Relative growth rate and efficiency of conversion of ingested food decreased with foliage age ) in both the 1981 and 1982 experiments. Relative consumption rate increased with foliage age in the 1981 and decreased ) in the 1982 experiment. The normal free-feeding period for western spruce budworm, Choristoneura occidentalis | Freeman, larvae begins in early spring at host » bud burst. The larvae feed primarily on imma- ture host foliage, ingesting mature foliage only when the preferred diet is unavailable. Nutrient content and ease of ingestion and ‘digestibility are often greater in immature fo- liage than in mature foliage (Heron 1964, Scriber and Slansky 1981). Host defenses such as lignins, silica, tannins, oils, waxes, and -resins often increase with foliage age. Such changes in food quality may be responsible for the western spruce budworm’s preference for early season foliage. _ The purpose of this study was to determine ‘the influence of the age of current year foliage on consumption and utilization by the west- ‘ern spruce budworm. | METHODS Feeding experiments were conducted 13 June-4 July and 10 August-25 August in 1981 and again 16 June-10 July and 10 July-30 July in 1982. Larvae were collected from the field and reared individually in 150 25 mm petri dishes at 16L:8D and 24-26 C during the eeding experiments. Sixteen replicates were set up for each year and season. Host tree and arvae selection and calculation of nutritional ndices followed Wagner and Blake (1983). Sollection of larvae and foliage, preparation of oNorthern Arizona University, School of Forestry, Flagstaff, Arizona 86011. foliage for feeding, and calculation of ingested foliage differed in 1981 and 1982. To standardize the host phenological stage, early season feeding experiments were begun when the foliage was at the “brush” stage (bud cap gone, needles flaring but no shoot growth so needles appear to arise from one location) (Shepherd 1983). The middle season feeding experiment conducted in 1982 was begun af- ter all the larvae used in the early season experiment had pupated. The 1981 late sea- son feeding experiment was conducted in early August, after the natural insect popula- tion had ceased feeding in the field. 1981 Feeding Experiment When C. occidentalis larvae were collected in the field, no attempt was made to segregate them based on the host foliage from which they were collected. Third instar larvae were selected for the first seasonal experiment us- ing head capsule measurements (Bean and Batzer 1957, Wagg 1958). The remaining lar- vae were reared to pupation, mated, and al- lowed to oviposit. When the eggs hatched, the larvae were placed on artificial diet to feed. Those larvae that did not diapause were selected from the population and reared to the fourth instar on artificial diet. Larvae were moved to natural foliage 48 hours before the second seasonal experiment was begun. A minimum of 24 hours was necessary for suc- cessful acceptance and assimilation of a food 169 170 source by the budworm larvae (Jacqueline Lee Robinson, personal communication). Foliage was collected at random from the lower crown of Douglas-fir, Pseudotsuga men- ziesii var. glauca (Beissn.) Franco, corkbark fir, Abies lasiocarpa var. arizonica (Merriam) Lemm., and Engelmann spruce, Picea engel- mannii Parry ex. Engelm. trees located ap- proximately 16 km north of Flagstaff, Arizona. Previous years needles were clipped from the stem, and the twigs, with only current year foliage attached, were weighed. The foliage twigs were replaced every 72 hours, and the twigs that had been fed upon were oven dried (60 C until no further weight loss occurred) and weighed. Foliage ingested was calculated on a dry weight basis as follows: W,=(W; - X)p—-DW, where: W,=dry weight of foliage ingested (mg) W,-=fresh weight of the foliage twig be- fore feeding (mg) X=% dry weight of an aliquot foliage twig DW, =dry weight of the foliage twig af- ter feeding (mg) The aliquot twig used to calculate the initial dry weight of each feeding twig was selected for uniformity in size and phenological stage from the same host tree that provided the feeding twig. Each replicate was terminated at pupation or when the larvae stopped feeding. Pupae and total feces for each replicate were oven dried and then weighed. Total foliage in- gested was calculated and duration was noted for each replicate. Data were analyzed using a one-way analysis of variance (AOV) with alpha equal to 0.10. 1982 Feeding Experiment Third instar larvae were collected. from Douglas-fir trees in the Kaibab National Forest, North Kaibab Ranger District, Ari- zona. Larvae used for the early season experi- ment were allowed to feed on Douglas-fir fo- liage and advance to the fourth instar. Larvae used for the middle season feeding experi- ments were placed in cold storage at the time of collection to retard their development. These larvae were brought to room tempera- GREAT BASIN NATURALIST | Vol. 46, No. 1 | ture five days before the experiment was to | begin and allowed to advance to the fourth | instar. Foliage used for the 1982 feeding experi- | ment was collected from a single medium | vigor (Waring et al. 1980) Douglas-fir tree. Paired foliage samples, selected for unifor- | mity in size and phenology, were collected | from midcrown at the four cardinal directions. One twig was used to determine the average — dry weight per needle; the other foliage twig was used to feed one budworm larva. The | average weight per needle was later used to | estimate the dry weight of the foliage con- | sumed by the insect. Foliage was replaced _ every 72 hours to assure freshness and accept- — ability. i Each replicate was terminated at pupation | or when feeding ceased. Total foliage ingested — was calculated; pupae and feces were oven — dried and then weighed After calculating the’ nutritional indices, data were analyzed using a. one-way AOV, itl alpha equal to 0.10. | RESULTS AND DISCUSSION | Total foliage ingested was eliminated from — the data analysis because the length of feeding | time was highly variable between the feeding © seasons. Nutritional indices for early, or nor- mal, season feeding were calculated for larvae\_ that had pupated. All the larvae feeding or: . middle (1982) and late (1981) season foliage | died before reaching pupation. However, the © nutritional indices were still calculated for lar vae that fed for at least 10 days. The percent o. - larvae that survived until pupation was calcu’ lated for each experiment. | Results of the 1981 experiment using Doug) ; , las-fir foliage showed significant difference: between early and late season foliage for al — the nutritional indices calculated (Table 1). Budworms feeding on late season foliage ap) pear to have increased their relative con’ ): sumption rate (RCR) to compensate for a de’ ~ crease in efficiency of conversion of ingestec | food to body weight (ECI). Despite this effort | C. occidentalis larvae were unable to maintail | a relative growth rate (RGR) statistically equa ~ to budworms feeding on early season foliage. | Larvae feeding on Engelmann spruce fo, q liage followed the same pattern of food utiliza jj tion as larvae feeding on Douglas-fir foliag) F-Prob. January 1986 TABLE 1. Effect of Douglas-fir foliage age on food uti- lization by western spruce budworm (1981). Food utilization indices Relative Relative Survival consumption growth Season % rate rate ECI Early (16,4)” 25 3.00 A” 0.09A 5.08A Late (16,4) 0 9.28 B 0.01B 0.16B 0.07 0.02 0.02 One-way AOV, a = 0.10, values followed by different letters are signifi- cantly different. ‘Numbers in parentheses are, respectively, initial number of larvae and number of larvae used to calculate nutritional indices. | ECI = Efficiency of ingested food to body weight. Designations apply for three following tables. TABLE 2. Effect of Engelmann spruce foliage age on food utilization by western spruce budworm (1981). Food utilization indices Relative Relative Survival consumption growth Season % rate rate ECI | Early (16,3) 19 LSS O.15A 11658 Late (16,2) 0 4.19B —0.05B — 1.34B 0.01 0.01 0.01 | F-Prob. | i f TABLE 3. Effect of corkbark fir foliage age on food utilization by western spruce budworm (1981). Food utilization indices Relative Relative Survival consumption growth Season % rate rate ECI , Early | (16,2) 13 2.67A 0.13A 4.78A | Late (16,5). 0 4.70A 0.06A 3.24A 0.50 0.24 0.56 ¢ | F-Prob. d | Nh iP TABLE 4. Influence of Douglas-fir foliage age on food utilization by western spruce budworm larvae (1982). Food utilization indices Relative Relative Survival consumption growth Season % rate rate ECI Early \(16,6) 38 1.58 A 0.08A 5.08A (Middle i (16,2) 0 1.05 B —0.08B —8.02B _F-Prob 0.02 0.0002 0.0014 BLAKE, WAGNER: WESTERN SPRUCE BUDWORM all (Table 2). Although their average RCR in- creased significantly, the mean larval weight was significantly lower in the late season ex- periment when compared to the mean weight of larvae reared on early season foliage. In fact, the larvae that were fed late season fo- liage lost weight, probably as a result of a negative average ECI. No significant differences were found be- tween seasons for the three indices reported when C, occidentalis larvae were fed corkbark fir foliage (Table 3). However, the pattern of change in food utilization was similar to the results of the experiments using Douglas-fir and Engelmann spruce. Significant differences were found in uti- lization between early and middle season feeding in 1982 (Table 4). RGR and ECI de- creased significantly, which follows the pat- tern of the 1981 experiment. However, RCR also decreased significantly during the middle season feeding rather than increasing as pre- dicted by Waldbauer (1968) and the 1981 feeding experiments. The larvae may have encountered chemical or physical feeding de- terrents in the middle season foliage that re- sulted in low RCR. This result could also have been due to the period of cold storage used to slow larval development, which may have been retarded to the point where the larvae could not recover. The statistical results of the foliage age ex- periments conducted in 1981 and 1982 can only be compared as to their relative patterns. Actual numbers should be disregarded due to the inconsistency in initial larval stage. CONCLUSIONS Host foliage phenology appears to greatly influence the food utilization and develop- mental success of C. occidentalis. Although no chemical tests were conducted on the fo- liage used for these experiments, the signifi- cant changes in RCR, RGR, and ECI suggest that chemical and physical changes may be occurring during the growing season. As pre- dicted by Waldbauer (1968) and Scriber and Slansky (1981), RCR was significantly greater and ECI was significantly lower for budworms feeding on late season foliage than for those feeding on early season foliage in 1981. De- spite this apparent effort to compensate for a W72 suboptimal food source, the budworm larvae feeding on late season foliage were unable to complete their development. The overall pattern of decreased RGR and ECland the survival rate of the larvae feeding on middle season foliage in 1982 are comparable to those of the 1981 experiments. The decrease in RCR from the early to the middle season experi- ment does not follow the 1981 pattern. This may be due to chemical or physical feeding deter- rents that could not be overcome by the bud- worm larvae. Budworm larvae feeding on middle and late season foliage experience retarded development and mortality, though many feed as long as suc- cessful larvae that feed on early season foliage. These findings are consistent with those of Heron (1964), who studied C. occidentalis feed- ing on mature white spruce needles. The feeding behavior of C. occidentalis ap- pears to be adapted for making the most of the nutrients provided by the foliage of its host early in the growing season, while avoiding the physi- cal and chemical defenses that may increase as the foliage matures. Studies of the physical and chemical changes in the foliage of these host trees as the growing season progresses are needed to better understand the feeding behav- ior of the western spruce budworm. ACKNOWLEDGMENT Work leading to this publication was funded in whole or in part by a program sponsored by the GREAT BASIN NATURALIST Vol. 46, No. 1 USDA Forest Service entitled Canada/United States Spruce Budworm Program—Science and Education Administration Agreement 59-2042- 1-3-013-0. LITERATURE CITED BEAN, J. L., AND H. O. BATZER. 1957. Mean head width for spruce budworm larval instars in Minnesota and_ | associated data. J. Econ. Ent. 50:499. HERON, R. 1964. The role of chemotactic stimuli in the | feeding behavior of spruce budworm larvae on white spruce. Canadian J. Zool. 43:247—269. SCRIBER, J.M., AND F. SLANSKy, JR. 1981. Nutritional ecol- _ ogy of immature insects. Ann. Review Ent. | 25:183-221. SHEPHERD, R. F. 1983. A technique to study phenological | interactions between Douglas-fir buds and emerg- | ing second instar western spruce budworm. Pages | 17—20 in Proceedings of the CANUSA workshop, | Forest Defoliator-Host Interaction: a comparison — between spruce budworm and gypsy moth. New | Haven, Connecticut. | Wace, J. W. B. 1958. Environmental factors affecting | spruce budworm growth. Forest Lands Research | Center. Corvallis, Oregon. Research Bulletin 11. 27 pp. | WAGNER, M. R., AND E. A. BLAKE. 1983. Western spruce | budworm consumption-effects of host species and | . foliage chemistry. Pages 49-54 in Proceedings of the CANUSA Workshop, Forest Defoliator-Host | Interaction: a comparison between spruce bud- | worm and gypsy moth. New Haven, Connecticut. | WALDBAUER, G. P. 1968. The consumption and utilization | of food by insects. Adv. Insect Physiol. 5:229-288. | WarING, R. H., W. G. THIES, AND D. MuscatTo. 1980. | Stem growth per unit of leaf area: a measure of. tree vigor. For. Sci. 26(1):112—117. | APPLE MAGGOT (RHAGOLETIS POMONELLA) ADAPTATION FOR CHERRIES IN UTAH Clive D. Jorgensen’, Darin B. Allred', and Richard L. Westcott” ABSTRACT.—The apple maggot, Rhagoletis pomonella (Walsh) is reported from Utah County, Utah, where it has adapted to sour cherries. It has been taken repeatedly from pheromone traps i the vicinity of hawthorn (Crataegus douglassii), but there are no Utah data that suggest it has adapted to apples. The apple maggot, Rhagoletis pomonella (Walsh), was first collected in Utah in 1976 froma Malaise trap in the Willard Basin of Box Elder County, Utah. This site was close to hawthorn, but 5 mi from the nearest apple trees. The apple maggot was not reported from Utah again until 1983 when the Utah Department of Agriculture (Edward J. Bi- anco, personal communication) took numer- ous specimens in the Mapleton (Utah County) area while trapping cherries for the western cherry fruitfly (Rhagoletis indifferens Cur- ran). Trapping for apple maggots was intensi- fied on cherries and apples in 1984. Continuous trapping (Pherecon® AM traps) in an unsprayed cherry orchard (1984) re- sulted in an emergence curve that seemed optimally synchronized with sour cherries (Fig. 1). Emergence was somewhat late for _ sweet cherries, unless they were left on the trees or were late varieties (e.g., Lambert). Perhaps the most interesting observation was the general paucity of adults collected when most commercial apples in Utah were ripen- ing. There was a slight increase in the number of trapped specimens in late September that » may have resulted from a partial second gen- eration, but this has not been confirmed. Sour Cherry (early emergence) Hawthorn Department of Zoology, Brigham Young University, Provo, Utah 84602. 4g Plant Division, Oregon Department of Agriculture, Salem, Oregon 97310. Early apples would be expected to ripen dur-ing the latter part of this emergence curve (Fig. 1). The apple maggot has apparently become adapted to cherries in the Mapleton, Utah, area. Although it is tempting to suggest genetic adap- tation has occurred in the population, it is more likely the adaptation to cherries is a phenological phenomenon within the population. Phenologi- cal adaptation of this type by apple maggots is certainly not new. It has been reported by Illing- worth (1912), Pickett and Neary (1940), Bush (1969), Reissig and Smith (1978), Diehl (1983), and others. Bush (1974), Reissig and Smith (1978), Prokopy et al. (1982), and Diehl (1983) suggested this allochronic isolation was impor- tant in evolution of the two sympatric host races in eastern North America, one for apples and the other for hawthorn (Crataegus spp.). Although we have found synchrony in apple maggot emergence with sour cherry develop- ment in Utah, it is not yet clear if an alternative host is present that provided the original source of apple maggots. Our work in 1985 has demon- strated a possible alternate host (hawthorn, Crataegus douglassii) that could provide this niche (Allred, unpublished data). It seems rea- sonable to suppose that allochronic isolation could develop in two directions from hawthorn in Utah: Hawthorn Apple (late emergence) 173 174 R o) 65 49 2 eee ee ew ee ee ee es 25 AM/TRAP/DAY No. 2 Lom 16 WzG 3 wT 19 JULY AUGUST GREAT BASIN NATURALIST Vol. 46, No. 1 Apple Harvest 4 12 2O° 26 2 RT 26 SEPTEMBER OCTOBER Fig. 1. Emergence curve for apple maggot (Rhagoletis pomonella) and its synchrony with sweet (cross-hatch) and sour (stipple) cherry harvest in central Utah, 1984. A 50% harvest for each is indicated. This possible phenomenon seems even more plausible when one considers the fact that almost all Utah apples are the late, hard varieties—Golden Delicious, Red Delicious, and Roman Beauty. The economic implica- tions of a possible synchronization with sour cherries, hawthorn, and apples are substan- tial. Such an extended synchronization to in- clude three fruits by the apple maggot in Utah seems rather unlikely, if data reported in east- ern North America are transferrable to our population. Bush (1969) and Reissig and Smith (1978) reported that apple host races emerged several weeks before hawthorn host races, with emergence peaks from 4 to 5 weeks apart. Since females are likely to de- velop and retain host fidelity after oviposition has started (Prokopy et al. 1982a,b), it is rea- sonable to suppose host adaptation for domes- tic fruits that mature before hawthorn is more likely than for fruits that mature after haw- thorn. Accordingly, adaptation to early-ma- turing apples grown in much of eastern North America and to cherries in Utah is more likely than adptation to the hard apple varieties that mature in late summer. LITERATURE CITED Busu, G. L. 1969. Sympatric host race formation on the speciation in rugivorous flies of the genus Rhago- letis (Diptera, Tephritidae). Evolution 23:237— 251. mation in the true fruitflies. Pages 3-23 in M. J. D. White, ed., Genetic mechanisms of speciation in insects, Australian and New Zealand Book Co., Sydney. DIEHL, S. R. 1983. Host race formation and sympatric species speciation in Rhagoletis (Diptera: Teph- retidae). Unpublished dissertation, University o! Texas, Austin. ILLINGWoRTH, J. F. 1912. A study of the biology of eel apple maggot (Rhagoletis pomonella), together; . 1974. The mechanism of sympatric host race for- | ( | | | with an investigation of methods of control. Cor- |. nell Univ. Agric. Expt. Sta. Bull. 324:129-187. PickeETT, A. D., AND M. E. Neary. 1940. Further studies | on Rhagoletis pomonella (Walsh). Sci. Agric. 20° | 551-556. Proxkopy, R. J., A. L. AVERILL, S. S. COOLEY, AND C. AN ROITBERG. 1982a. Associative learning in egglay- ing site selection by apple maggot flies. Science: 218:76-77. Proxopy, R. J., A. L. AVERILL, S. S. COOLEY, C. A. Ror! BERG, AND C. KALLET. 1982b. Variation in hos’) acceptance pattern in apple maggot flies. Proc. 5t/ Internat. Sympos. Insect-Plant Relationships, Wa Sean 1982. Pudoc, Wageninger: p. 123-129) REIssic, W. H., and D. C. SmirH. 1978. Bionomics 0) aes pomonella in Crataegus. Ann. Ento, mol. Soc. Amer. 71:155-159. i \ { NEW RECORDS FOR MONOTROPA HYPOPITHYS (ERICACEAE) FROM COLORADO William Jennings’, Loraine Yeatts!, and Velma Richards! ABSTRACT. —Monotropa hypopithys L., rarely collected in Colorado, was taken five times during 1984. Coupled ' with previous collections, it is now known from 10 Colorado counties. The cool, moist August of 1984 may have contributed to its apparent ubiquity. Ae a hypopithys L. in habitat. Fig. 1. Photo: Monotrop The saprophytic plant Monotropa hypo- pithys L. (Ericaceae), popularly known as pinesap (Fig. 1), has been described as rare in Colorado (Harrington 1954, Weber 1976). The Colorado Natural Heritage Inventory (Peterson 1984) maintains it on the list of Plant Species of Special Concern as a plant that may be in danger of extirpation from Colorado be- cause of the scarcity of occurrence. A survey of three regional herbaria shows a paucity of specimens (four at University of Wyoming, two at Colorado State University, and three at University of Colorado), generally supporting its characterization as a rare plant. During the summer of 1984, M. hypopithys was collected by us in five Colorado counties: Routt, Jackson, Douglas, Mineral, and Las Animas. These new observations, coupled with our previous sightings and collections and the existing herbarium specimens, show that M. hypopithys is probably more wide- spread than previously thought. As shown on the map (Fig. 2), it is now known from 10 counties in Colorado. In addition, B. E. Nel- ‘Kathryn Kalmbach Herbarium, Denver Botanic Gardens, 909 York Street, Denver, Colorado 80206. 175 176 wor rat ) Nos arnt x = | GRAND m0 BLANCO | ee ee es ___|avams gai 7 EAGLE a A CART ARF HELD A wESA Pe Sey OLAKE ae al WA ‘ore is Z |GUNNISON — i | i ie aa CHAFFEE WONTROSE SAGUACHE ° AY ee Ns CUSTER SAN MIQUEL MINSOALE MINERAL A ' GREAT BASIN NATURALIST Clee eee Ses: ne FREMONT RIO GRANDE [ALAMOSA Vol. 46, No. 1 FFER | le CHE VENNE | ARAPAHOE DOUGLAS] ELBERT LINCOLN A! Miso : | HUERFANO Pin LAS ANIMAS OSTILLA Fig. 2. Colorado distribution of Monotropa hypopithys L.: @® Herbarium Specimens (COLO, CS, RM) (Table 1); A author's collections and observations (Table 2). son of the Rocky Mountain Herbarium at Uni- versity of Wyoming concurs that it was much more widespread during 1984; he observed it in the Medicine Bow Mountains of southern Wyoming. The details on existing herbarium speci- mens are presented in Table 1; the details of our recent observations are tabulated as Table 2. All specimens were deposited at the Kathryn Kalmbach Herbarium of Denver Botanic Gardens and/or University of Colo- rado—Boulder. Photographs are in our col- lections. Habitat for M. hypopithys is in damp conif- erous woods where there is little understory vegetation, but rather forest duff composed of rotting logs, twigs, and conifer needles. Usu- ally such a habitat is found on a north-facing slope at high elevation. Most of our collection or observation sites follow this characteriza- tion. The 1983 and 1984 Jackson County sites, the 1974 and 1979 Boulder County sites, the 1984 Las Animas County site, and the 1980. Hinsdale County site were all on moist, heav- | ily wooded, northeast- , north- , or northwest- | | facing slopes above 8,000 ft (2,400 m). The 1984 Mineral County site was on a steep densely forested, but southwest-facing slope. The 1984 Routt County site was on flat ground | in open woods, with a number of rotting logs. This site was adjacent to a national forest campground and presumably much of the | deadwood had been gathered as firewood. | contributing to the openness. However, the! : 1984 Douglas County site was a dry but heav- © ily wooded hillside at 6,800 ft (2,070 m) with \ but the most likely habitat compared with the surrounding area. | The plant was associated with pinedrop: — (Pterospora andromedea Nutt.), another eri’ cad, and with coralroot orchids (Corallorhize sp.), which are usually in fruit when M. hypo: pithys is at its prime. At the Jackson County | sites the plants were associated with Rhodo | January 1986 TaBLE 1. Herbarium specimens of Monotropa hypopithys L. (CS, RM, COLO). JENNINGS ET AL.: SAPROPHYTIC PLANT 77 Collector and County Location Date accession number Douglas or E] Paso* Palmer Lake — August 1914 Bethel (CS33885) Larimer East Twin Lake 3 August 1974 Kopp (CS26043) Mineral or Archuleta* —_N of Pagosa Springs 14 August 1917 Payson (RM101307) Fremont Sec 31 T47N R12E 20 August 1938 Lemmon (RM-USFS) Fremont Sec 13 T47N R1OE 25 August 1960 Gierisch (RM-USFS) La Plata Sec 30 T37N R6W 25 July 1934 Loughridge (RM-USFS) Fremont Phantom Canyon 21 August 1971 Howard (COLO256444) Boulder Sec 23 T2N R73W 28 July 1966 Clark & Arp (COLO214312) Routt Big Red Park 7 September 1969 Stevenson (COLO244531) _ *County of collection not listed on herbarium sheet. TABLE 2. Collections and observations of Monotropa hypopithys L. Date Collector Castlewood Canyon County Location Douglas Jackson TON R82W Sec 14/15 Las Animas NW of Lake Dorothy Routt TION R86W Sec 11 Boulder West of Peaceful Valley Jackson Helena-Grizzly Trail near Bear Creek Mineral T37N RIE Sec 8 Hinsdale Piedra River T37N R2W Sec 5 dendron albiflorum Hooker; at the Mineral County site with Goodyera oblongifolia Raf. and Pyrola picta Smith ex Rees; and at the Las Animas County site with Goodyera repens (L.) R. Brown, G. oblongifolia, and Coral- lorhiza sp. Described by Harrington as “pink or red- _ dish, sometimes yellowish, ” all degrees of col- oration from red through pink to a creamy yellow have been noted by us. In fact, such _ variations can be seen ata single site, in plants 1 | W] \y _ only a few feet apart. The reason for the seemingly ubiquitous nature of M. hypopithys during 1984 is not known. Perhaps our extensive travel in Colo- rado during 1984 had something to do with it. It may be that the atypically moist, cool Au- gust of 1984 permitted the plants to stay f fresher longer than normal, leading to an ap- parent but not necessarily real increase in the population. As a saprophyte, the plant is quite 26 July 1984 4 August 1984 25 August 1984 2 September 1984 3 August 1974 5 August 1979 16 August 1983 WF] (specimen) WF] (specimen) WF] (specimen) WF (specimen) WF (photos) LY (specimen) 25 August 1984 18 August 1980 LY & VR (specimen) LY (photo) fleshy. In a warmer, dryer August, M. hypo- pithys changes from red or yellow to crispy- brown rather quickly and effectively disap- pears into the featureless brown duff of the forest floor. Whatever the reason for its widespread ap- pearance during 1984, it seems clear that the plant isn't really rare (only during some years), but occurs in Scattered localities in many forested mountainous counties. LITERATURE CITED HARRINGTON, H. D. 1954. Manual of the plants of Colo- rado. Sage Books, Swallow Press, Chicago. NELSON, B. E. 1984. Personal communication. 6 Septem- ber 1984. PETERSON, J. S. 1984. Colorado Natural Heritage Inven- tory: plant species of special concern as of January 31, 1984. Rocky Mountain Natural Heritage Pro- gram, 1370 Pennsylvania St., Denver, Colorado. WEBER, W. A. 1976. Rocky Mountain flora. Colorado Assoc. Univ. Press, Boulder, Colorado. TREE DENSITIES ON PINYON-JUNIPER WOODLAND SITES IN NEVADA AND CALIFORNIA Susan Koniak! ABSTRACT.— The densities of singleleaf pinyon and Utah juniper trees in four diameter classes (1-9, 10-19, 20-29, and = 30 cm) were measured on 522 plots of 1/10 ha each throughout the Great Basin. Density distribution patterns of pinyon and juniper varied with aspect, elevation, and eastern (EGB) versus western Great Basin (WGB) locations. On most locations north and, toa lesser extent, west slopes supported higher densities of pinyon than south and east slopes, with high relative densities of small diameter trees on north slopes and large diameter trees on west slopes. Pinyon densities were higher on EGB than on WGB sites and on higher elevation than on lower elevation sites. Juniper densities were higher on EGB than on WGB sites and on lower elevation than on higher elevation sites. Juniper densities on low-elevation WGB sites were higher on south and west aspects than on north and east, with higher | relative densities in the 20-29 cm diameter class than in other diameter classes. On low elevation EGB sites, east and south slopes supported higher juniper densities than did north and west slopes, with comparatively higher relative densities in the 10-19 cm diameter class. significant among aspects on high elevation sites. Singleleaf pinyon-Utah juniper (Pinus monophylla and Juniperus osteosperma) woodlands occupy about 7.1 million ha within the Great Basin (Tueller et al. 1979). These woodlands are most prevalent on mountain slopes above sagebrush-dominated (primarily Artemisia tridentata tridentata ) communities of the valley bottoms. The woodlands may extend to the mountain tops or to the lower edge of high-elevation sagebrush (primarily Artemisia tridentata vaseyana) communities. Few studies have recorded the effect of aspect and elevation on tree distribution in mature pinyon-juniper woodlands (West et al. 1978, Tueller et al. 1979, Cooper et al. 1980, Tausch etal. 1981). This paper reports on variations in tree densities by diameter class over four as- pects, two elevation classes, and two sections of the Great Basin. FIELDS METHODS AND DaTA ANALYSIS In 1981 and 1982, the densities of pinyon and juniper trees in four diameter classes (1-9, 10-19, 20-29, and = 30 cm) were deter- mined on 522 plots of 1/10 ha each, on 20 areas in Nevada and California (Fig. 1). Tree diame- ters were estimated at stump height. Densi- ties were obtained on plots selected for an- Differences in relative densities between diameter classes were not other study in which vegetation on wildfires of various ages and adjacent unburned sites were recorded. Data from 112 unburned woodland sites were combined with data from 410 wildfire sites for analysis. On wildfire sites the number of remnant tree skeletons was re- corded to reflect tree densities. The smallest diameter class may be underestimated on burned sites. All wildfire sites were burned — within the last 30 years, with 76% of those — sites burned within the last 15 years. We sam- pled only those sites in which pinyon was or had been the dominant species and the under- | story had been or was assumed to have been _ (by extrapolation from conditions on adjacent — sites) substantially reduced by tree competi- — tion. Each plot contained a minimum of 10/9 trees, 5 with diameter = 20 cm. In each of the ’' | | 20 areas, tree cores were taken from a a mum of 3 mature dominant pinyon trees on) the unburned sites for a total of 91 cores. Plot) elevations ranged from 1,585 to 2,280 m, and | average slope for each area ranged from 16% ) to 64%. Estimated annual precipitation for each area ranged from 20 to 33 cm. Following Cronquist et al. (1972), the sam- pled areas were divided into three groups— the Reno (or western) section containing 10 sampled areas and 313 plots, the central Great | ‘At the time of this research, the author was with the Intermountain Research Station, Forest Service, U.S. Department of Agriculture, Ogden, Utah 84401.) I located at Reno, Nevada. 178 jaa i January 1986 CVA | FE. CLAN ALPINE VIRGINIA RANGE RANGE PINE NUT RANGE ¢? a | SWEETWATER PARADISE | RANGE / RANGE PINE GROVE RANGE KONIAK: PINYON-JUNIPER WOODLANDS NEVADA 179 RUBY MTNs. SCHELL CR. RANGE TOIYABE RANGE EGAN RANGE WILSON CR. RANGE Fig. 1. Location of 20 study sites in Nevada and California. | Basin and Tonapah sections (or central sec- tion) containing 5 sampled areas and 117 ‘plots, and the Calcareous Mountain (or east- ern Great Basin) section containing 5 sampled areas and 92 plots. Similarity of tree distribu- | tion between aspects in the western and cen- tral sections on both high- and low-elevation sites justified combining the data from these two sections for analyses (collectively desig- ' nated as the western Great Basin section). One-way and two-way analyses of variance were used to compare the number of trees per " plot (density) and the percent of the total tree density in each diameter class (relative den- \sity) on eastern and western Great Basin sites; on north, south, east, and west aspects; and at nigh ane| low dlevatiors, In the eastern Great Basin (EGB), the point selected to differenti- ate between high and low elevations was 2,160 m compared to 2,040 m in the western Great Basin (WGB). The difference in eleva- tional division reflects sectional differences in woodland belt width and elevational range. The lowest site measured in the eastern sec- tion was 2,030 m compared to 1,585 m in the western section. RESULTS AND DISCUSSION The average age of the pinyon trees in each area sampled ranged from 72 to 159 years. The median age of all pinyon trees sampled was 96 years, with 51% of the trees sampled falling within the 70—110-year age group. Age differ- 180 GREAT BASIN NATURALIST Vol. 46, No. 1 TABLE 1. Comparison of tree densities (number of trees per 1/10 ha) between eastern and western Great Basin sites, high and low elevations, and four aspects. Location North Total tree density: Eastern (high elevation) 28.0%" Eastern (low elevation) 30.2% Western (high elevation) 23.6 Western (low elevation) 92.94 Pinyon tree density: Eastern (high elevation) 94, 4% Eastern (low elevation) 25.0" Western (high elevation) oR Te? Western (low elevation) 21.4 Juniper tree density: Eastern (high elevation) 3.6% Eastern (low elevation) 5,gred Western (high elevation) 0.5° Western (low elevation) 1.0° Means followed by the same letters a, b, c, d, or e do not differ significantly at P < 0.05. Each block is a separate analysis. ences were not apparent between EGB and WGB. Most stands appeared to have established in the mid- to late 1800s or early 1900s (Tausch et al. 1981). During this period pinyon-juniper woodlands were heavily harvested to supply fuel and charcoal for the mining industry. Subse- quent regrowth, concurrent with limited har- vesting and intensive fire control, has tended to create large woodland areas dominated by trees of roughly equal age (Lanner 1980, Young and Budy 1979). Intensive livestock grazing, which reduced competition from herbaceous and shrubby plant species, may also have con- tributed to the increase in tree density and domi- nance during this period (Blackburn and Tueller 1970). Unburned stands exhibited few signs of major disturbance from this period until the pre- sent, suggesting that the burned areas may also have been relatively undisturbed until wildfire occurred. Tree densities on all aspects for both high- and low-elevation classes were higher on EGB sites than on WGB sites, with differences being smallest among north aspects (Table 1). Tree densities were not different between aspects or between elevation classes on EGB sites nor be- tween high- and low-elevation WGB sites of the same aspect. In the WGB, tree densities were higher on north slopes than on other aspects for both high and low elevations. Pinyon Densities Pinyon densities were generally highest on high-elevation sites and north aspects (Table Aspect East South West 33.2? 29.0% 99.8 28.0%¢ 30,77 27.57 17.6° 7 GEE 197 Fi 17.0° 18.5% 19:8 al 27.49 22. 6%e¢ 24.7% 16.5% gee 20.5°4 Anes 16.9°¢ norsk 16.4% 15.24 aegct 4,gbe4 6.4P¢ Beles 11.5% 10.5° 6.9? 0.4° 0.4° 0.4° 0.8° Byes 2.59 | ie | | 1). Cooler, moister environments and longer | periods of snow cover were characteristic of | these sites. Pinyon densities were also higher in the EGB than WGB, corresponding to_ higher average annual precipitation and lower | average annual temperatures in the EGB> (NOAA 1983). Pinyon dominance, contrary to | total tree density, was greatest in the WGB, ~ where summer precipitation is ata minimum. — In both EGB and WGB the second highest’ pinyon densities were consistently found on) west aspects. | High-elevation EGB sites, unlike other lo-. cations, supported higher pinyon densities on east aspects than on north aspects. This may — be related to the higher number of storms — from the east in the EGB than in the WGB. © However, the same relationship is not evident _ at lower elevations. q Juniper Densities The density of Utah juniper was greatest in © the EGB, on low-elevation sites, and on south © slopes (Table 1). The frequency distribution patterns indicate that juniper dominance is positively correlated to higher summer pre- © cipitation and greater diurnal fluctuation oj | soil moisture and temperature. There was — some evidence that the higher incidence o} © paleozoic sedimentary soil parent material in | the EGB may be related to higher densities 0! © juniper, but more comprehensive study is re- | quired to clarify the relationship. Higher an- | \ i ; | | January 1986 KONIAK: PINYON-JUNIPER WOODLANDS 181 PINYON JUNIPER NORTH EAST SOUTH WEST NORTH EAST SOUTH WEST WGB LOW ELEVATION SITES 2 WGB > HIGH rE ELEVATION nny) SITES az iw sa ww > = < led eal ay ve EGB LOW ELEVATION SITES 40 20 EGB HIGH ELEVATION SITES Oo 1 Within each block of 16 bars, bars with the same letters a,b,c,d,e,f, or g do notdiffer significantly at P< 0.05. * Diameter classes : S = 1-9 cm, MS = 10-19 cm, ML = 20-29 cm, L= 230 cm. This sequence is repeated for every group of four bars. j Fig. 2. Variation in pinyon and juniper relative densities among four diameter classes, four aspects, two elevations, and two locations (WGB = western Great Basin; EGB = eastern Great Basin). I nual precipitation on EGB sites compared to _low-elevation east-facing slopes in the EGB WGB sites probably was not directly related may be related to differences in prevailing _ ‘o higher juniper densities, since greater ju- summer weather patterns (Houghton 1969, , niper densities were not also evident on gen- Presley 1978). In the WGB summer storms _ erally moister sites at high elevations. are infrequent and primarily originate in the _ Relatively high juniper densities on low-el- west and southwest. In the EGB summer , vation west-facing slopes in the WGB and on _ storms occur frequently, often originating in 182 the southeast. Influence of aspect on juniper densities was less at high elevations, resulting in approximately equal tree densities on all aspects. Relative Densities According to Diameter Classes—Pinyon On north aspects, differences between rela- tive densities (percentage of total tree density in each diameter class) of pinyon in the 1-9, 10-19, and 20—29 cm diameter classes were small except on low-elevation WGB sites (Fig. 2). On these sites a higher proportion of pinyon trees were in the 1-9 cm class. Rela- tive densities in the = 30 cm class were signifi- cantly lower than those in the other diameter classes. North slopes, compared to other as- pects, tended to have higher proportion of pinyon trees in the 1-9 and 10-19 cm diame- ter classes. On east aspects the highest pinyon relative densities were generally in the 20-29 cm di- ameter class and the lowest in the = 30 cm class. Low-elevation WGB sites deviated from this pattern with high relative densities in the 1-9 cm class. Compared to other as- pects, east-facing slopes tended to have higher proportions of pinyon trees in the 20-29 cm classes. South aspects, like east slopes, had higher proportions of pinyon trees in the 20-29 cm diameter class than in the other classes for all locations. Unlike the distribution of pinyon on east aspects, south aspects also supported rel- atively high pinyon densities in the = 30 cm diameter class at most locations. South slopes rivaled east slopes for highest relative densi- ties in the 20-29 cm class and west slopes for highest relative density in the = 30 cm class. On west aspects, pinyon distribution dif- fered substantially between WGB and EGB sites. On WGB sites relative densities in the 1-9, 10-19, and 20—29 cm diameter classes were not significantly different, and those in the = 30 cm class were significantly lower. On EGB sites pinyon relative densities tended to be highest in the 20-29 and = 30 cm classes and lowest on the 1—9 and 10-19 cm classes. Compared to other aspects, west-facing slopes consistently supported high propor- tions of trees in the = 30 cm diameter class. GREAT BASIN NATURALIST Relative Densities According to Diameter Class—Juniper Because juniper densities were very low on | Vol. 46, No. 1 | high-elevation WGB sites, these data were | deleted from the analysis and discussion of density distribution patterns among diameter | classes. Juniper relative densities in the = 30. cm diameter class were lower than in the | other classes for all locations except high-ele- — vation EGB east and west slopes. On north | slopes juniper relative densities were consis- | tently higher in the 1-9 and 10-19 cm diame- — ter classes than in the 20-29 cm class. On west | aspects juniper relative densities were consis- | tently higher in the 20-29 cm class than in the. other classes. Distribution of juniper among_ the three smaller diameter classes on east and | south aspects exhibited no consistent pat-) terns. Relative densities in the two smaller) classes tended to be higher on north and east’ slopes than on other aspects. South and west aspects frequently supported higher juniper relative densities in the 20-29 and = 30 cm: class than did north and east aspects. ? { Elevational and Sectional Effects on Distribution Patterns The basic patterns of pinyon and juniper distribution among diameter classes were) similar between high and low elevation (Fig | 2). Several trends were apparent, but they were generally not significant. On WGB site: variation in relative densities between eleva — tion classes was not consistent between as. pects. On EGB sites all aspects generally ex! | hibited an increase in pinyon in the 1-9 and= i 30 cm diameter classes with increasing eleva | tion, whereas middiameter class relative den sities decreased. Juniper relative densities i the = 30 cm diameter class also tended ti ° increase at higher elevations on EGB sites) - accompanied by a decrease in the 10-19 cn | class. Consistently larger trees at high eleva’ - tion EGB sites may indicate older stands 0 better growing conditions at these elevations’ Distribution patterns of pinyon and junipe) among diameter classes were similar betwee’ EGB and WGB sites (Table 2). For bot} pinyon and juniper, distribution patterns 0! north aspects were almost identical betwee EGB and WGB sites. On other aspects ther’ i [ il January 1986 KONIAK: PINYON-JUNIPER WOODLANDS 183 TABLE 2. Comparison of pinyon and juniper relative densities on eastern (EGB) and western Great Basin (WGB) sites among four aspects and four diameter classes. Diameter class and location Aspect 1-9 cm 10-19 cm 20-29 cm => 30cm WGB EGB WGB EGB WGB EGB WGB EGB Pinyon North 32.8 30.5 28.9 28.8 28.9 STi 10.7 10.1 East 30.3 24.8 26.0 31.6 32.1 38.1 13.6 6.1 South 23.4 23.0 28.1 22.2 34.4 35.0 14.0 19.6 ’ West 29.7 21.3 26.6 23.3 29.2 30.9 Wee). PL al _ Juniper North 32.7 33.4 38.5 39.3 24.4 20.9 5.0 6.4 East 21.2 39.9 42.3 33.9 33.9 14.4 OL0 SSeS South 15.1 24.4 28.1 36.3 54.8 * 28.6 2 Oss West 28.4 Po 29.3 21.2 36.0 36.9 6.8 14.7 *Pairs denoted by an asterisk (*) are significantly different at P < 0.05. was wider variation; however, few of these differences were significant. WGB_ south slopes supported significantly greater juniper relative densities in the 20-29 cm diameter class than did EGB south slopes. Pinyon rela- tive densities on west-facing sites and juniper relative densities on south-facing and east- | facing sites were significantly higher in the = 30 cm class on EGB sites than on WGB sites. | The consistently higher proportion of trees in the = 30 cm class on EGB sites than on WGB sites may indicate better growing conditions » or an earlier onset of stand establishment. | Major Distribution Patterns | Among Diameter Classes _ Sites with the highest total juniper densi- ‘ties (i.e., low elevation south and east EGB sites and low elevation south and west WGB _. sites) displayed different distribution patterns ' among diameter classes (Fig. 2). In the WGB, - regeneration appeared to have been greatest _ in the early tree stage of the successional cy- _ cle, after the first generation of trees had _ reached seed-bearing age. Subsequent regen- _ eration may have been limited by competitive . interaction for water, nutrients, and space, or . by infrequent environmental conditions con- _ ducive to seed germination and seedling sur- _ vival. In the EGB the highest tree establish- , Ment rates apparently occurred later in the “successional cycle than they did in the WGB. , This may have been because environmental | conditions were favorable during this period, ; producing an unusually high regeneration , wate. Or perhaps the natural accretion of trees , ‘into seed-bearing age, the continued capabil- ity of the sites to support new trees, and cli- mate favorable to tree establishment may have yielded progressively higher propor- tions of small diameter trees, diminishing only in recent years when either adverse envi- ronmental conditions or competition limited regeneration. Interspecific competition may also have affected density distribution among diameter classes. For example, high propor- tions of pinyon in the larger diameter classes on low-elevation EGB sites may indicate early domination by pinyon and subsequent de- layed establishment of juniper. Sites with the highest pinyon densities, north and high-elevation EGB east aspects, displayed similar distribution patterns among diameter classes. Pinyon regeneration on these sites remained high after tree establish- ment and had not decreased substantially in recent years. The relatively low proportions of trees in the = 30 cm diameter class on these sites may indicate either that competition be- tween large numbers of small-diameter trees restricted or delayed the number of trees at- taining substantial girth or that stand estab- lishment occurred later on these sites than on other sites. Historically, north and east slopes may have been exposed to a greater frequency and severity of wildfires than have other as- pects because of the generally higher biomass to carry fire. Fire suppression policies in the early 1900s allowed the invasion of many sites previously dominated by shrub disclimax communities. Stand establishment on in- vaded shrub areas would occur slower than stand establishment after tree harvesting be- cause cut-over areas generally retain numer- 184 ous small diameter trees. Greater competi- tion from perennial species (Koniak 1985) on these slopes may also delay tree establish- ment. Neither the tree competition nor the late stand establishment hypotheses are clearly supported by tree diameter and age data in this study or in the literature (Meeuwig 1979, Cooper et al. 1980). More comprehensive study is needed to clarify the underlying processes. Of the four aspects, west slopes consistently supported the second highest pinyon densi- ties at all locations. Less competition or ear- lier stand development with less disturbance or both may explain the disproportionately high number of trees in the = 30 cm diameter class on west slopes. EGB west slopes sup- ported higher proportions of pinyon trees in the = 30 cm diameter class than WGB west slopes at both high and low elevations. In the WGB tree establishment has remained high on west slopes. On EGB west slopes competi- tion from large-diameter trees may have re- stricted regeneration, limiting the proportion of pinyon trees in the smaller diameter classes. Distribution of pinyon and juniper over as- pect, elevation, and eastern versus western Great Basin locations tended to be negatively correlated. However, variations in relative density distribution between diameter classes were often similar. For both pinyon and ju- niper, north and, to a lesser extent, east as- pects tended to support high relative densi- ties of small-diameter trees, whereas south and west aspects tended to support high rela- tive densities of large-diameter trees. Of the diameter classes, relative pinyon and juniper densities were generally lowest in the = 30 cm class. For both pinyon and juniper, rela- tive densities in the = 30 cm class tended to increase with elevation, especially on south and west aspects. One striking difference in the diameter class distribution is the consis- tently lower proportion of juniper trees in the = 30 cm classes compared to pinyon trees. Juniper appear to be more sensitive to compe- tition than pinyon (Meeuwig 1979) and may be slower to establish in communities where both species are represented. Most stands sampled had a substantial pro- portion (20% —35%) of both pinyon and ju- niper trees in the 1-9 cm diameter class, indi- GREAT BASIN NATURALIST Vol. 46, No. 1 cating these woodlands are still in the forma- tive stages of stand renewal (Meeuwig and Cooper 1981). At what point equilibrium with the natural environment will be reached, and the nature of the stand at that time, are largely matters of conjecture. Recording the process, however, will have value in our understand- ing the dynamics of this system. LITERATURE CITED BLACKBURN, W. H., AND P. T. TUELLER. 1970. Pinyon and juniper invasion in black sagebrush communities in east-central Nevada. Ecology 51:841—848. Cooper, S. V., R. O. MEEUWIG, AND P. T. TUELLER. 1980. Classification of pinyon-juniper woodlands in the Great Basin. Nevada Agric. Expt. Sta. Proj. 650, Final Rept. 60 pp. CRONQUIST, A., A. H. HOLMGREN, N. H. HOLMGREN, AND J. L. REVEAL. 1972. Intermountain flora: vascular plants of the Intermountain West, USA. Vol. 1. Hafner Publishing, New York. 270 pp. HOUGHTON, J. G. 1969. Characteristics of rainfall in the | Great Basin. Desert Research Institute, Univer- | sity of Nevada, Reno. 205 pp. Konlak, S. 1985. Succession in pinyon-juniper woodlands | following wildfire in the Great Basin. Great Basin — Nat. 45:556—566. LANNER, R. M. 1980. The pinon pine: a natural and cul- |— tural history. University of Nevada Press, Reno. | 208 pp. | MEEuwic, R. O. 1979. Growth characteristics of pinyon- juniper stands in the western Great Basin. USDA | For. Serv. Res. Pap. INT-238. 22 pp. MEEuwic, R. O. AND S. V. Cooper. 1981. Site quality and | _ growth of pinyon-juniper in Nevada. For. Sci. 27: | 593-601. | NATIONAL OCEANIC AND ATMOSPHERIC SCIENCES ADMINIS- | | TRATION. 1983. Climatological data annual sum- | mary, Nevada, Vol. 98, No. 13. Dept. of Com- | merce. 17 pp. | PRESLEY, A. R. W. 1978. Synoptic patterns and precipita- tion in the pinyon-juniper zone of the Great Basin. | Unpublished thesis, University of Nevada, Reno. | 149 pp. Tauscu, R. J., N. E. WEST, AND A. A. Nasi. 1981. Tree age and dominance patterns in Great Basin pinyon- | juniper woodlands. J. Range Manage. 34:259- 264. TUELLER, P. T., C. D. BEESON, R. J. Tauscu, N. E. WEST, AND K. H. REA. 1979. Pinyon-juniper woodlands of the Great Basin: distribution, flora, vegetal cover. USDA For. Serv. Res. Pap. INT-229. 22 pp. WEST, N. E., R. J. Tauscu, K. H. REA, AND P. T. TUELLER. 1978. Phytogeographical variation within juniper- | pinyon woodlands of the Great Basin. Pages| 695-705 in K. T. Harper and J. L. Reveal, eds., | Intermountain biogeography: a symposium. Great) | Basin Nat. Mem. 2. Brigham Young University | Provo, Utah. YOUNG, J. A., AND J. D. Bupy. 1979. Historical use of Ne- | vada’s pinyon-juniper woodlands. J. For. Hist. 23:| 112-121. NOTICE TO CONTRIBUTORS Manuscripts intended for publication in the Great Basin Naturalist or Great Basin Natural- ist Memoirs must meet the criteria outlined in paragraph one on the inside front cover. They should be directed to Brigham Young University, Stephen L. Wood, Editor, Great Basin Naturalist, 290 Life Science Museum, Provo, Utah 84602. Three copies of the manuscript 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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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. See Notice to Contributors on the inside back cover. 8-86 650 25552 ISSN 017-3614 ft The Great Basin Naturalist PUBLISHED AT PROVO, UTAH, By BRIGHAM YOUNG UNIVERSITY ISSN 0017-3614 VOLUME 46 | 30 April 1986 No. 2 BIOLOGY OF RED-NECKED PHALAROPES (PHALAROPUS LOBATUS ) AT THE WESTERN EDGE OF THE GREAT BASIN IN FALL MIGRATION Joseph R. Jehl, Jr." ABSTRACT.—Large numbers of Red-necked Phalaropes migrate overland across the Great Basin in fall, occurring commonly at highly saline lakes. Migrants occur at Mono Lake, California, from mid-July to mid-October. The earliest migrants are adult females, followed several weeks later by adult males, and finally by juveniles. Adults make up ca 75% of the population, with males outnumbering females by 5:4. From 1980 through 1984 an estimated 52,000—65,000 birds passed through the area each year, except in 1983, when only 36,000 were recorded. The low number might be attributable to high mortality on oceanic wintering grounds in the Southern Hemisphere in 1982 associated with the severe E] Nino. At Mono Lake the phalaropes concentrate near the shore and feed almost exclusively on brine flies. The migrants neither gain much weight nor accomplish much molt during their sojourn, which suggests that the average stay is only a few days. Some aspects of the molt pattern differ from those reported elsewhere. After the breeding season, many Red- necked Phalaropes (Phalaropus lobatus) mi- grate from breeding grounds in the arctic re- gions of the New World to pelagic wintering areas in the equatorial Pacific Ocean. Sub- ‘stantial numbers, presumably from the west and central Canadian Arctic, move overland, and each fall hundreds of thousands occur in the Great Basin of the western United States. Concentrations exceeding one million indi- viduals have been estimated at Great Salt Lake, Utah (Kingery 1982). Thousands also occur at Lake Abert, Oregon (K. Boula, per- sonal communication), Mono Lake, California Cogswell 1977, Winkler 1977), and Stillwater National Wildlife Refuge, Fallon, Nevada (S. Thompson, personal communication), and arge but unestimated numbers visit the Salton Sea (McCaskie 1970, Garrett and Dunn 1981). The existence of an overland -»nigration route in this species has long been i “mown (e.g., Fisher 1902, Bent 1927, Grin- nell and Miller 1944). What does not seem to have been appreciated, however, is that sa- line lakes are preferred stopping places. This report documents the fall migration at Mono Lake, California. It is based on studies that spanned the entire fall migration period, July through mid-October, for six consecutive years (1980-1985). The major goals were to document abundance, period of occurrence, and ecological requirements of the Red- necked Phalarope and to determine the com- position of the population throughout the mi- gration period. METHODS Mono Lake (surface elevation 6,380 ft in 1984) is a large (ca 160 km) basin lake at the western edge of the Great Basin in east central California. It is highly saline (surface salinity 75-90 0/00 during this study) and alkaline (pH = 10) and contains no fishes (see Hubbs and 1Hubbs Marine Research Institute, 1700 South Shores Road, San Diego, California, 92109. 185 186 Fig. 1 Juvenile (left) and adult (right) Red-necked Phalaropes. Miller 1948). Accordingly, the aquatic inver- tebrates (brine shrimp, Artemia sp., and brine flies, Ephydra hians) that provide food for the phalaropes and a few other species of migratory birds can attain great abundances. I made behavioral observations through the entire migration period. Specimens were weighed, measured, and examined for molt and external parasites. Stomach contents were identified using a binocular microscope. Fat-free weights were determined using stan- dard ether-extraction techniques. I estimated the percentage of molting feathers in each of the body tracts and determined molt scores for primaries and rectrices. A value of 0 is given to an unmolted feather, 5 for one that has been replaced, and 1 to 4 for intermediate stages (e.g., Morrison 1976). Thus, a bird that has replaced all its primaries and rectrices would receive scores of 100 (i.e., 5 < 10 pri- maries < 2 wings) and 60 (5 < 12 rectrices). CENSUSING.—Because of their localized distribution and preference for nearshore habitat, phalaropes are relatively easily cen- sused. Counts from a boat cruising ca 400 m offshore will reveal nearly all birds, except those hidden behind tufa formations. As the migration period progresses, it is not uncom- mon to find flocks of 30-400 birds 2 km or more from shore; these can be detected by routine transects. Censusing by boat pro- duces consistent and replicable results. I esti- GREAT BASIN NATURALIST Vol. 46, No. 2 | mate that errors did not exceed 15%—20%, | even when populations were large. For exam-_ ple, replicate censuses of birds on the eastern | half of Mono Lake on 31 August and 1 Septem- ber 1981 resulted in estimates of 6,800 and 7,053 birds, a difference of 3.5% AGE AND SEX RATIOS.—Male Red-necked | Phalaropes bear the responsibility for incuba- _ tion and caring for the young. Females leave | the breeding grounds about the time young | hatch and are followed by the adult males | several weeks later and, finally, by the juve- | niles (Hildén and Vulanto 1972). At Mono. Lake the migration period of each of these groups is protracted, perhaps because: (1) 1) the. | population is composed of birds from several | nesting areas, where the start of breeding may vary by a week or more; (2) the species is _ polyandrous, and females that obtain more. than one mate remain on the breeding, grounds longer than those that obtain a single , mate; (3) and males that lose their eggs or! young migrate earlier than successful nesters. | — Sex ratios in adults usually cannot be deter- | mined in the field because plumage differ-| ences, although pronounced on the breeding, grounds, dull with the onset of molt. Esti- mates in this report are derived largely from specimens collected with no conscious bias and are supplemented by banding data (Win- kler 1977). Age ratios, however, can be deter- mined by plumage (Fig. 1; Prater et al. 1977)» —_ ee saa i April 1986 JEHL: PHALAROPE BIOLOGY 187 1981 e 1982 = 1983 oO 1984 10 20 30 20 JULY AUGUST 30 10 20 30 0 20 = 30 SEPTEMBER OCTOBER Fig. 2. Population size of Red-necked Phalaropes at Mono Lake, California, 1982-1984. until mid-September, or later. Because the distribution of adults and juveniles is not nec- essarily similar, I determined age ratios in several areas of the lake, attempting to achieve a minimum total sample size of 100. _ Data obtained in that manner were similar to these obtained by collecting. RESULTS CHRONOLOGY.—In my experience, only small numbers of the Red-necked Phalarope pass through the Mono Basin each spring. I have seen flocks totaling 200 birds on several occasions on dates ranging from 30 April to 24 May, with a maximum of 800-1,000 on 13 May 1984. That much larger numbers may occur occasionally can be inferred from Ry- sers (1985) report of thousands killed near Reno, Nevada, in mid-May 1964 (see also Gaines 1977). One or two may summer in ‘some years. The few birds that appear in late ‘June are often molting heavily, which sug- gests that they may not have reached the nest- ing areas; certainly they could not have bred. Usually there was no significant influx of mi- grants until the last third of July, although in 1981 more than 2,000 were present by mid- July (Fig. 2). Numbers increase through early August, peak near 10 August at ca 12,000 individuals, and remain high for the next month. A decrease is evident by 10-15 Sep- tember; by early October fewer than 200 re- main; and by the last third of October the species has departed. The latest dates on which phalaropes were present (based on ir- regular surveys) were 17 October 1980, 9 Oc- tober 1981, 21 October 1982, and 14 October 1983. Only once did I see migrants depart. On 6 August 1982, four flocks totaling 450 birds left the mouth of a freshwater creek where they had been bathing, climbed high toward the Sierra Nevada, and disappeared to the west. This suggests that some migrants may move directly toward the Pacific Ocean. COMPOSITION OF THE POPULATION. —Adult females predominated among the early mi- grants (Fig. 3). The rapid population increase in late July (Fig. 2) results from the arrival of postbreeding males. By late July adult fe- males composed fewer than half the popula- 188 No. 12 10 18 25 13 10 20 30 10 JULY GREAT BASIN NATURALIST Vol. 46, No. 2 11 20 = 10 Estate 20 30 10 20 30 10 SEPTEMBER Fig. 3. Age and sex ratios of Red-necked Phalaropes based on specimens collected from 1980 through 1982. The | percentage of adult females determined by banding studies (Winkler 1977) is shown by @. The dotted line shows the inferred average percentage of adult females from late July through mid-September. tion, and their abundance decreased through the remainder of the fall. Juveniles began ar- riving in mid-August (earliest 13 August 1980, 10 August 1981, 2 August 1982, after 6 August 1983), predominated by the first of Septem- ber, and composed virtually all the population by early October. SIZE OF THE POPULATION.—The size and composition of the population at major phases of the migration period are shown in Table 1. This species does not use Mono Lake either as a molting or staging area (see below) and, because the arrival period of each age and sex class is protracted, I infer that the average stays of most migrants ranged from 5 to 14 days (see also Evans 1984, Kersten and Smith 1984). Short stop-overs were also suggested by weight data (see below). The bulk of the migration for any age and sex category is ac- complished in 30 days. Assuming an average stay of 10 days, one can estimate the total population at roughly three times the sum of the peak counts in each category. If so, 52,000 Red-necked Phalaropes passed through the Mono Basin in 1981, 65,500 in 1982, 36,000 in 1983; limited data from 1980 and 1984 suggest a population similar to 1981. Although the validity of the multipliers is unknown, the census data are sufficiently accurate to show that annual differences in abundance are real. | DISTRIBUTION. —Red-necked Phalaropes | are pelagic for much of the year, but at Monc Lake they tend to avoid midlake habitats in favor of shallow areas within 200 m of shore. | There they congregate over shallowly sub- merged rock formations, which provide a sub-. strate for pupating brine flies; brine flies of al | ages are their major food. In July migrants appear along the centra | north shore, where there are large expanses 0 tufa-encrusted rocks. As numbers increase the population spreads laterally, and by mid August birds are found almost anywhere or the periphery where there is submerged rock >. Distributional patterns were similar eacl — year (Fig. 4), with two exceptions. In early ~ September 1981 approximately 2,500 of 9,000 birds were flocking offshore, perhaps in antic { April 1986 JEHL: PHALAROPE BIOLOGY 189 TABLE 1. Size and composition of Red-necked Phalarope populations at Mono Lake, California, 1981-1984. Population size Composition* Number observed Estimated Adult Female Adult Male Juvenile Western” Eastern total % N % N % N 1981 2-3 Jul 9 303 400 [95] 380 [ 5] 20 14-15 Jul 805 1,070 2,050 [80] 1,640 20] 410 21 Jul ua ee 3,000 24Jul _—1, 100-1, 200 ND (60) [40] 29-30 Jul 500+ 1,500+ 3,600 [55] 1,980 [45] 1,620 4 Aug 3,445 ND 6,000 [48] 2,880 [52] 3,120 02 120 1] Aug 3,708 6,370 12,000 [35] 4,200 63] 6,840 2 960 19 Aug 4,300 2,130 8,500 [33 2,805 [47] 3,995 20 1,700 31 Aug 7,030 = =e se Bk 64 1-2 Sep 6,800 3 9,000 [13 1,110 [17] 1,530 70 6,300 22 Sep 50 165 250 — [20 50 80 200 9 Oct — 0 20 — —- — 100 20 Total (%) 14,995(35.8) 17,585(42.0) 9,300(22.2) 1982 2-4 May 200 ae 200 23 Jun 2 ee XID 2 see 5 Jul 5 5 [92] 5 8] 11 Jul 2} — 10 [80] 8 [20] 2 17-19 Jul 100 725 1,000+ [80] 800 [20] 200 25-27 Jul 100 1,100 1,500 [60] 900 [40] 600 2-4 Aug 5,040 2,900 8,500 [58] 4,930 42] 3,570 0.2 16 6-8 Aug 5,300 2,600 10,000 [58] 5,800 [42] 4,200 0.2 16 18-19 Aug 7,020 2,630 12,000 [30] 3,600 [66] 7,920 4 480 2-3 Sep 7,360 3,550 14,000 [ 7] 980 [35] 4,900 58 ~=—-8, 120 23 Sep 4,300 192 4,800 — — [20] 960 80 = 3, 840 21 Oct — 5 5 — — 100 5 Total % 17,025(32.8) 22, 352(43.1) 12,477(24.1) » 1983 | 14 May 200 al 200 | 13Jun 1 a 100 i — 0 21 Jun 6 Ge 100 6 an 0 4ful 1 10 ~—«[92] 9 [ 8] 1 12Jul 30 <50 —*(80] 40 [20] 10 i) 23 Jul 200 800 1,000 [60] 600 [40] 400 6Aug 2,400 500 3,200 [48] 1,536 [52] 1,664 10 Aug = as 8,000° [42] 3,360 [58] 4,640 21 Aug 5,748 100 6,300 [19] 1,200 [30] 13950 50P-woyllo0 15-16 Sep 1,800 3,000 5,000 [ 5] 250 [15] 750 80 4,000 12-14 Oct 2 = <10 — _ = 100 10 ) Total % 7,002(30.2) 9,415(39.0) 7,160(30.8) 1984 3 Apr 200 13 May 1,000 16 Jul 400 [80] 320 [20] 80 13 Aug 12,000 [3] 4,440 [63] 7,560 + | 7Sep 8,000 [10] 800 [32] ~ 2,560 58 6,400 28 Sep 2,000 [20] 400 80 —:1,600 Total (%) 5,560(23.0) 10,600(43.9) 8,000 (33.1%) a . . © = 4 ate Percentages of age or sex classes in brackets are derived from the average for any date as shown in Figure 1. Data not in _ observations. _ | For convenience in censusing, the lake is divided into western and eastern sectors. ‘Data provided by J. Bright, personal communication. } , |) ul brackets are based on field 190 2-3 JULY 1981 200 S GREAT BASIN NATURALIST Vol. 46, No. 2 29-30 JULY 198) 17-19 AUGUST 1981 See ae 22 SEPTEMBER 1981 2-4 Aug 1982 18-19 Aug 1982 Fig. 4. Distribution of Red-necked Phalaropes at Mono Lake, California, in 1981 and 1982. ipation of departure. In mid-September 1983, phalaropes foraged mainly at the southwest- ern and southeastern corners of the lake. In those areas the rising lake had inundated freshwater marshes, creating new substrate for flies in the form of submerged vegetation, 3 Sept 1982 \ a 6-8 Aug 1982 im 23 Sept 1982 and the drowned plants were blackened by masses of pupae. | Foop.—To study feeding habits, I col- ' lected birds in many localities, making delib: erate efforts to sample in offshore localities a: well as at sites remote from submerged rocks. April 1986 JEHL: PHALAROPE BIOLOGY 191 100 2 ea aoa 5 1980 [__] Brine Flies, Ad Brine Flies, Imm ce} % 50 ee ) 100 1981 % 50 fe) 100 3 1 4 9 3 1982 | % 50 O 10 20 30 10 20 30 10 20 30 JULY AUGUST SEPTEMBER Fig. 5. Diet (% volume) of Red-necked Phalaropes at Mono Lake, California, in 1980, 1981, and 1982. In 1982 the stages of brine flies were not determined after 8 August. Numbers above columns refer to sample size. {Cursory observations of birds feeding over or until water currents wafted a larva or pupa ‘swarms of brine shrimp could easily lead one __ within reach. If this did not occur within five ‘to infer that shrimp compose the bulk of the | seconds or so, the birds ceased their typical )phalarope diet. However, careful scrutiny “swim and peck’ foraging method and began » showed that the birds were ignoring shrimp spinning, which quickly brought larvae to the and waiting until an adult brine fly passed by _ surface. | 192 Volumetric analysis of stomach contents (Fig. 5) showed that brine flies made up essen- tially all the diet in July-August, and well over 90% later in the year. The predominance of dipterans was also shown by Wetmore (1925) at Great Salt Lake and by K. Boula (personal communication) at Lake Abert. I estimate that brine flies composed approximately 99% of the phalaropes diet at Mono Lake. Gravel was present in 70% of the samples, and seeds or unidentified plant parts in 5%. I found no age or sex differences in food preference. Brine fly larvae were the only prey found in the single spring sample (30 April 1984). In 1980 only | adult (14 August) had fed on brine shrimp. In 1981 phalaropes fed exclu- sively on flies through 21 August. At the next sampling period, 1-2 September, all of 12 birds collected offshore had fed mainly on shrimp, and 7 also contained fly parts; 1 of 7 birds on 21 September contained a few shrimp eggs. In 1982, to investigate the possi- bility of a seasonal shift in diet, I biased my sampling methods and collected only in off- shore sites, and mostly late in the season. Although these samples included birds feed- ing directly on or over a commercial trap for brine shrimp, no shrimp were encountered. I infer that they are taken mainly when brine flies are unavailable. WATER USE.—Red-necked Phalaropes visit freshwater sources on the periphery of Mono Lake, often in association with Wilson’s Phalaropes (P. tricolor), to drink and bathe. Their visitations seem irregular, and daily ac- cess to fresh water does not seem to be re- quired. Birds often bathe in the lake itself, sometimes over sublacustrine springs from which fresh water boils to the surface. For- merly, in the 1970s, Red-necked Phalaropes regularly visited small ponds just north of the lakeshore, where many hundreds (M. Mor- ton, personal communication) to several thou- sand might be seen (Winkler 1977). As the lake has receded, pond use has declined. In 1980 fewer than 500 frequented the ponds each day, in 1981 birds were less abundant, and since 1982 the ponds have been avoided. Visitations occurred mainly in late afternoon, or even after dark, when groups of 5 to 20 might fly in. After circling nervously for sev- eral minutes, they would alight and bathe or drink frantically for 30-60 seconds before re- GREAT BASIN NATURALIST Vol. 46, No. 2 turning directly to the lake. This nervous be- , havior was unlike their calm demeanor at | freshwater sources on the lakeshore, where | they might bathe and preen for an hour or | more. Presumably it was related to a greater _ risk of predation. | WEIGHT. —Weight data are shown in Fig- ure 6. Since Red-necked Phalaropes, unlike many migratory shorebirds, do not lay on | great fat stores at this point in their migration, it follows that they do not use Mono Lake as a — staging area but only as a stop-over location. | Fat-free weights of 6 adult females averaged | 29.3 g (range 24.8—32.0) and of 10 males 27.8 g | (range 24.2-33.4), which is ca 0.3 g less than | the mean arrival weights. MorRPHOMETRICS.—Mensural data are pre-— sented in Table 2. Although there is no recog- | nized variation in this species, it may eventu-— ally be possible to. infer the source of the migrants if mensural data from other areas reveal significant differences. | BEHAVIOR.—Phalaropes make sporadic vis- its to fresh water and sometimes come ashore > or climb on tufa boulders to rest and preen. — Otherwise, they remain continuously near feeding areas and fly only short distances each . day. It is not uncommon to see a half-dozen feeding over a single pupa-covered rock, pecking larval flies from the water column or lunging at adults as they come to the surface. Occasionally one to two birds will act as satel-’ lites to a Wilson’s Phalarope, waiting for food that the larger bird spins to the surface (Fig. — 7). Whether this commensal activity is an en- ergy-saving device or indicates that the Red- | necked Phalarope’s effectiveness in spinning. is less than that of the larger, longer-legea@ Wilson's is unknown. Mo.ttr.—Cramp and Simmons (1983: 639). reported that male Red-necked Phalaropes — undergo limited molt on the head and neck on | the breeding grounds but that females delay until after the start of migration (see alse! Hildén and Vuolanto 1972). They also re-| ported that (1) molt began in “moulting areas. near the breeding grounds,” (2) renewal oi | flight feathers did not usually commence unti| _ late September, and (3) southward migratior began in August or September. 7. Data from Mono Lake modify those conclu: _, sions. Migration begins in July, and the firs! , migrants appearing shortly after their disap: April 1986 Males 50, Weight (g) 50 Juveniles JEHL: PHALAROPE BIOLOGY 193 28.0-43.8 (347436)g N=50 31.0—43.5(35.6+39) 9. N=23 1 5 1 15 20 25 30 5 10 JULY AUGUST SEPTEMBER Fig. 6. Weights of Red-necked Phalaropes based on combined samples from 1980 through 1982. TABLE 2. Measurements (mm) of Red-necked Phalaropes at Mono Lake, California, 1980-1982. Exposed culmen N Rangeandmean S.D. N Adult female 47 20.5—24.9 (22.7) + 1.0 48 Adult male 53 20.2-24.8 (22.5) + 1.0 53 Juvenile female 13 22.2-93.5 (22.8) + 0.4 13 Juvenile male 9g 20.4-23.0 (21.7) + 1.0 9 pearance from North American breeding ar- eas is noted, (e.g., Jehl and Smith 1970). This schedule allows no time for stops at supposed molting areas. Adults of either sex begin molting on the nesting grounds, probably at about the same time. From 8 through 12 July 1980, five of eight newly arrived postbreeding females ‘showed light to moderate molt in most of the »body tracts, two had just started to molt on the vhead and neck, and one had not begun. By late July most females showed moderate to heavy molt on the body, and by mid-August ‘many appeared to be in basic plumage, al- Wing, flat Tarsus Range and mean_ S.D. 19.2-22.2 (20.7) + 0.7 19.4-22.2 (20.9) + 0.7 19.4-22.0 (20.9) + 0.8 19.5-21.4 (20.3) + 0.7 Range andmean S.D. N 105-116 (111.2)+2.3 48 100-113 (106.3) +2.9 54 100-114 (108.6) + 3.7. 13 100-109 (103.7) + 3.3 9 though many feathers had not yet been re- placed. The degree of molt in the earliest males, which arrive two weeks later than fe- males, is less than in females examined on the same date. As late as 8 August, however, afew males showed no molt in any tract. Molt of the primaries in each sex (Fig. 8) starts in early August; therefore, many females leave Mono Lake before replacing any. All but one female examined after late August, however, had re- placed the inner four or =i ve primaries on each wing. Wing molt in males begins slightly later; one male had not molted any primaries as late as 23 September. I found no evidence 194 GREAT BASIN NATURALIST Fig. 7. A Red-necked Phalarope feeding commensally with a Wilson’s Phalarope (foreground). of molt of secondaries or wing coverts, except for the irregular replacement of some greater secondary coverts. Tail molt (Fig. 9) in many females begins by early July, perhaps while they are still on the nesting grounds. All females examined after late July had molted some rectrices, but only one had completed molt by late August. In males the start of tail molt coincides with their arrival in late July, and all but two examined after 7 August showed some molt. One had replaced all rectrices by 5 September, whereas another on 23 September had not started. The central pair of rectrices is lost first, usually followed by the outer pair or pairs, but this is not invariable and molt is not necessarily symmetrical. One male taken on 28 August had 14 rectrices, of which the cen- tral and outer pair were half grown and the fourth pair was just starting to grow. Juveniles, which arrive in mid-August, have not started to molt body feathers. Some are in heavy molt by late August and nearly all show body molt before departing; one on 21 September, however, had none. Molt of the rectrices may start in late August, but even by late September it remains slight. Juveniles do | not replace primaries at this time. MortaLity.—I used beached bird censues to index mortality patterns. Although cen- | suses along 8-16 km of shoreline were made at two-week intervals (often more frequently) in’ several localities and were spread over the. entire migration periods of i981—1984 (irreg- — ular surveys were made in 1980 and 1985), I| : | found only two dead phalaropes. Evidently | this phase of the migration imposes no great). stress on the species. : | RED PHALAROPE.—The Red Phalarope (P. | fulicaria) is rare in the interior of the western states and provinces (Goossen and Busby 1979). The species remains in the Arctic late into the fall, and inland records, as for the Red-necked Phalarope, probably pertain to! | birds from the western and central Arctic that) | are en route to wintering areas in the Pacific. At Mono Lake one or two Red Phalaropes , have been recorded in four of six years since 1980, all but one of the sightings (1 September , 1980) occurring between 14 and 23 October (1981-1, 1982-2, 1983-1). Goossen and Busby . (1979) suggested that most fall migrants in the) April 1986 50 Wing Score 40. Females 30, 20 10 JEHL: PHALAROPE BIOLOGY AUGUST 195 15 20 25 30 5 Lo TH BO “ae SEPTEMBER Fig. 8. Wing molt scores of Red-necked Phalaropes based on combined samples from 1980 through 1982. interior were juveniles. Two of three that I have seen were juveniles; the third was an adult found dead. The juveniles seemed healthy and were feeding well offshore, ap- parently on brine shrimp. DISCUSSION The Red-necked Phalarope uses lakes in the Great Basin as a stopover point on the ‘southward migration and not as a molting or ‘staging area. At Lake Abert, Oregon, K. Boula (personal communication) reported )peak numbers of 5,000—7,000 in the first three weeks of August. At Mono Lake I found that )peak populations of ca 12,000 were attained ‘by early August and maintained into. early September, whereas Winkler (1977) reported ‘the greatest numbers in late August. From ‘shore-based observations, he estimated 21,600 on 30 August 1976, and H. Cogswell personal communication) estimated more ithan 40,000 on 22 August 1958. Whether \these apparent differences in abundance and iming may pertain to censusing methods, ex- “ceptionally large flights in 1958 and 1976, the availability of alternative stopping places in | some years, or other factors is unknown. In mid-September 1985, when only a few hun- dred birds were seen in an incomplete survey of Mono Lake, S. Thompson (personal com- munication) reported 16,000 near Fallon, Ne- vada. Historical data are too scanty to determine if there has been any change in the size of the population using Mono Lake. Fisher (1902) reported “countless hundreds.’ Grinnell et al. (1918) considered the species very common but barely mentioned Wilson’s Phalarope, which currently predominates; perhaps infor- mation available to them was inadequate to differentiate between the two species or was based on seasonal data obtained after the main migration of Wilson's Phalarope, which is largely completed by mid-August. The composition of the Red-necked Phalarope population remained fairly con- stant during this study. Adult males outnum- bered adult females 5:4. This might indicate a faster stopover period for females or their re- liance on different migration routes or staging areas, as may be the case for Wilson's Phalarope (Jehl, unpublished manuscript). I suspect, however, that the differences are real 196 GREAT BASIN NATURALIST Vol. 46, No. 2 | Score Females Tail Juveniles 5 10 615 200 6.2506«6330)=—Cl 55 10 JULY and indicate an unbalanced tertiary sex ratio, as would be expected in a polyandrous spe- CIES: Total population size varied each year from ca 52,000 to 65,000, except in 1983, when it dropped to ca 36,000. The lower numbers in that year cannot be ascribed to censusing er- ror, as techniques and observers remained constant. Nor can they be due to unfavorable food conditions (brine flies were exceptionally abundant) or toa poor breeding season (young were present in expected proportions). Be- cause the migrant population at Lake Abert was judged to be ca 30% below normal (K Boula, personal communication), I infer that regional events were involved. Two possibili- ties seem of potential importance. Fresh wa- ter was abundant in the far west in 1983, a result of the atypically wet winter of 1982. Asa result many aquatic habitats were renewed, and those may have provided alternative stop- ping points. Perhaps of far greater signifi- AUGUST Fig. 9. Tail molt scores of Red-necked Phalaropes based on combined samples from 1980 through 1982. 10 SEPTEMBER 15 20 5 20 25 cance, however, was an apparent drop in breeding populations in the spring of 1983. Low numbers were recorded in the ae Canadian Arctic (Churchill, Manitoba, J. Reynolds, personal communication) as well as, in the Gulf of Alaska (Middleton Island, P| Gould, personal communication). Birds nest: | ing in those areas presumably winter in the equatorial Pacific off South America, an are: | that was profoundly affected by an El Nino o unprecedented strength (Rasmusson 1985). The abnormally warm waters are known tc — have had dramatic effects on breeding birds ir i the central and eastern Pacific. Reproductior — failed at Christmas Island (Schreiber anc Schreiber 1984), and in areas influenced by | the Humboldt Current major oceanographic changes affected the distribution of fish. seabirds, and marine mammals (Barber anc. Chavez 1983). I suspect that phalaropes suf fered very high mortality on the wintering grounds in 1982, which could not be made uy | April 1986 by the production of young in the following breeding season, thus accounting for the low migrant populations of 1983. Schreiber and Schreiber (1984) noted that “field biologists must recognize that atmo- spheric circulation patterns that undergo ir- regular anomalies may affect their study re- gions and species far from marine eco- systems.” Events at Mono Lake illustrate this and point out the difficulties in trying to un- derstand the population dynamics of migra- tory species in the absence of information about condition and events throughout the range. ACKNOWLEDGEMENTS This study was funded by the Los Angeles Department of Water and Power, with a coop- erative agreement with the U.S. Fish and Wildlife Service. Additional support was re- ceived from Hubbs Marine Research Insti- tute, San Diego State University, National Geographic Society, Kawasaki Motors of America, and Sevylor Boats. Permits were issued by the California Department of Parks and Recreation and Fish and Game, U.S. Fish and Wildlife Service, U.S. Bureau of Land Management, and U.S. Forest Service. Many persons contributed to this study; D. Babb, K. Boula, S. Bond, B. Cord, H. Cogswell, J. Granito, R. Jarvis, D. Jehl, R. Kastelein, P. Kelly, D. Kent, C. D. Lit- tlefield, S. Mahoney, M. Messersmith, D. Shuford, B. Tillemans, S. Thompson, D. Winkler. Personnel from the Mono Lake Tufa State Reserve helped facilitate the field work. | J. Bright and the staff of the Cain Ranch, Lee Vining, California, provided outstanding sup- -port at all times. A draft of the manuscript benefited from ‘the criticism of K. Garrett, S. Hurlbert, A. | Pollak, R. Schreiber, and N. K. Johnson. LITERATURE CITED BARBER, R. T., AND F. P. CHAVEZ. 1983. Biological conse- quences of E] Nino. Science 222: 1203-1210. JEHL: PHALAROPE BIOLOGY 197 BENT, A. C. 1927. Life histories of North American shore birds, Part I. U.S. National Museum Bulletin 142. Cramp, S., AND K. E. L. Simmons. 1983. Handbook of the birds of Europe the Middle East and North Africa. ae III. Oxford University Press, Oxford, Eng- and. COoGSwELL, H. L. 1977. Water birds of California. Califor- nia Natural History Guides 40. University of Cali- fornia Press, Berkely, California. EVANS, P. R. 1984. The British Isles. Pages 261-275 in P. R. Evans, J. D. Goss-Custard, and W. G. Hale, eds., Coastal waders and wildfowl in winter. Cam- bridge University Press, Cambridge, England. FISHER, W. K. 1902. A trip to Mono Lake, ornithological and otherwise. Condor 4: 1-12. GAINES, D. 1977. Birds of the Yosemite Sierra. GRT Book Printing, Oakland, California. GARRETT, K., AND J. DUNN. 1981. Birds of southern Cali- fornia. Artisan Press, Los Angeles, California. GOOSSEN, J. P., AND D.G. Busby. 1979. Occurrences of the Red Phalaropes in the Prairie Provinces and adja- cent states. Canadian Field-Naturalist 93: 446-449. GRINNELL, J.. AND A. H. MILLER. 1944. The distribution of the birds of California. Pacific Coast Avifauna 27. HILDEN, O., AND S. VUOLANTO. 1972. Breeding biology of the Red-necked Phalarope Phalaropus lobatus in Finland. Ornis Fennica 49: 57-85. Husss, C. L., AND R. R. MILLER. 1948. The Great Basin, with special emphasis on glacial and postglacial times. II. The zoological evidence; correlation be- tween fish distribution and hydrographic history in the desert basins of western United States. Bulletin University of Utah 38: 17-166. JEHL, J. R. JR, AND B. A. SmiTH. 1970. Birds of the Chruchill region, Manitoba. Manitoba Museum of Man and Nature, Special Publication No. 1. KERSTEN, M., AND C. J. SMITH. 1984. The Atlantic coast of Morocco. Pages 276-292 in P. R. Evans, J. D. Goss-Custard, and W. G. Hale, eds., Coastal waders and wildfowl in winter. Cambridge Uni- versity Press, Cambridge, England. KiInGERY, H. 1982. Mountain West. American Birds 36: 202. McCaskIE, G. 1970. Shorebird and waterbird use of the Salton Sea. California Fish and Game 56: 87-95. Morrison, R. I. G. 1976. Moult of the Purple Sandpiper Calidris maritima in Iceland. Ibis 118: 236-246. PRATER, A. J., J. H. MARCHANT, AND J. VUORINEN. 1977. Guide to the identification and ageing of Holarctic waders. British Trust for Ornithology, Field Guide 17. RASMUSSON, E. 1985. El Nino and variations in climate. American Scientist 73: 168-177. RYSER, F. A., JR. 1985. Birds of the Great Basin. University of Nevada Press, Reno, Nevada. WINKLER, D. W., ed. 1977. An ecological study of Mono Lake, California. Institute of Ecology Publication 12, University of California, Davis, California. SOME RELATIONSHIPS OF BLACK-TAILED PRAIRIE DOGS TO LIVESTOCK GRAZING Craig J. Knowles’ ABSTRACT. —Relationships of black-tailed prairie dogs (Cynomys ludovicianus) to livestock grazing were studied | from 1973 to 1983 on the Charles M. Russell National Wildlife Refuge and the Fort Belknap Indian Reservation in | northeast Montana. A total of 154 prairie dog colony sites was examined, and most were in association with livestock watering sites and/or areas where the topsoil was disturbed by human activity. Roads and cattle trails were found in 150 — of the prairie dog colonies. Prairie dog colonies were found to be located significantly (p < 0.001) closer to livestock | water developments and homestead sites than randomly located points. Observations showed cattle to occur signifi- cantly (p < 0.05) more on quarter sections with prairie dog colonies as opposed to quarter sections without prairie dog | colonies. Forage utilization at one prairie dog colony was estimated at 90% by midsummer. Prairie dogs consumed | about a third of the vegetation, with grasses the predominant forage class used. Habitat characteristics of black-tailed prai- rie dogs (Cynomys ludovicianus) colonies have been reported on over a wide geographic region (Reid 1954, Koford 1958, Smith 1967, Hassien 1976). Prairie dogs are frequently as- sociated with areas of low-growing vegetation and areas intensively grazed by ungulates (Mead 1898, Osborn and Allan 1949, King 1955, Koford 1958, Smith 1967). Although many authors have commented on this rela- tionship, little quantitative information exists on the subject. Furthermore, it is not clear in the literature if prairie dog colonies develop at intensively grazed sites or if the presence of prairie dogs attracts ungulates to an area. There are documented cases of declining prairie dog numbers following reduction or elimination of ungulates from an area (Mead 1898, Osborn and Allen 1949, Uresk and Bjugstad 1983). Knowledge of the spacial dis- tribution and habitat use of ungulates and prairie dogs over a broad area is important to understanding prairie dog-ungulate relation- ships. The purpose of this study was to investi- gate the distribution, habitat use, and forage utilization of black-tailed prairie dogs and do- mestic livestock in northeastern Montana. STUDY AREA AND METHODS Data were gathered from 1973 through 1975 and from 1978 through 1980 on the Boulder, Montana 59632. 198 Charles M. Russell National Wildlife Refuge (CMRNWR) and during 1983 on the Fort | Belknap Indian Reservation (FBIR) in north- | east Montana. The CMRNWR is typified by | rough, river breaks country merging with | rolling prairies on either side of the Missouri — River. Coniferous forest habitats dominated | by ponderosa pine (Pinus ponderosa) and | Rocky Mountain juniper (Juniperus scopulo- | rum) are commonly found on the steeper. slopes along the Missouri River and cover about 36% of the land area. Shrub-grassland | and grassland habitats occur on the broad’! ridge tops and coulee bottoms that extend’ from the prairies onto the CMRNWR. Glaci- . ated prairies with relatively little topographic — relief compose the majority of the FBIR. | Shrub-grassland and grassland habitats domi- nate these sites. Coniferous habitats occur only on the foothills of the Little Rocky Moun-' tains that border the FBIR on the south. Prairie dog colonies on both the CMRNWR | and the FBIR are restricted to the shrub- \ grassland and grassland habitats. Western © wheatgrass (Agropyron smithii), blue grama_ (Bouteloua gracilis), green needlegrass (Stipa | viridula), and needle-and-thread grass (S. co- ] mata) are the predominant grasses in these — { wort (Artemisia frigida), plains prickly pear _ habitats. Common forbs include fringed sage- (Opuntia polycantha), and yellow sweet- ' clover (Melilotus officinale). The shrub layer © i 'Montana Cooperative Wildlife Research Unit, University of Montana, Missoula, 59812. Present address: Fauna West Wildlife Consultants, Box 113, al April 1986 is composed largely of big and silver sage- brush (Artemisia tridentata, A. cana) and greasewood (Sarcobatus vermiculatus). The CMRNWR and the FBIR are grazed primar- ily by cattle, although sheep and horses are present in a few grazing allotments. Native ungulates are pronghorns (Antilocapra ameri- cana), elk (Cervus elaphus), and mule and white-tailed deer (Odocoileus hemionus, O. virginianus ). The CMRNWR was surveyed for prairie dog colony sites in 1979. Colonies were -mapped on frosted mylar plastic over 1:24,000 aerial photos while driving or walking the perimeter of the colony. The area of each colony was determined with an electronic dig- itizer. Prairie dog colonies on the FBIR (sur- vey area boundaries described in Knowles and Knowles 1984) were surveyed in 1983 and were mapped on 7.5’ USGS _ topographic maps. Area for each colony was determined with a dot grid. At each prairie dog colony site, | recorded presence or absence of prairie dogs, livestock developments (reservoirs, wells, salt licks, and calf feeders), home- steading activity, roads, and well-established cattle trails. Complete survey coverage was made of each study area. However, on FBIR inactive colony sites were not investigated nor was one small colony located around a private residence. Special effort was made to study prairie dogs at the west end of the CMRNWR north of the Missouri River because of the number (36) and density (6.3 colonies/100 km’) of prairie dog colonies. All stock water develop- ments and homestead sites were accurately located on a map over a 570 km’ area. I used a ‘Students t-test to test the hypothesis that ‘mean distance from the geographic center of ‘each prairie dog colony in this area to the ‘nearest stock water development or home- ‘stead site was not different than the mean distance to such features for 120 randomly ‘chosen points. In this same area, I made weekly surveys of cattle each summer and fall ‘from 1973 to 1975 in two pastures (20,244 ha) sof the four-pasture Nichols Coulee rest-rota- ‘tion grazing system as part of another study (Knowles and Campbell 1982). Quarter sec- tion location, habitat type, and slope were | recorded for each cattle group when first ob- served. The quarter section distribution of KNOWLES: PRAIRIE DoG ECOLOGY 9 cattle was compared to the quarter section distribution of prairie dog colonies occurring in these two pastures using a chi-square test of homogeneity. Habitat type designation fol- lowed Mackie (1970) except for analysis pur- poses, where observations of cattle in the Xanthium strumarium and Agropyron- Symphoricarpos habitat types were com- bined and observations in the Artemisia longi- folia and Pinus-Juniperus habitat types were combined. Spring/summer forage utilization of prairie dogs, prairie dogs and other wildlife (primar- ily mule deer and elk), and prairie dogs and cattle was investigated at a prairie dog colony located next to a reservoir site in the Nichols Coulee allotment. A 7.7 ha area of the 16.4 ha colony was fenced to exclude cattle in July 1978. Ten agronomy cages were placed on each side of the fence that passed through the center of the colony in pairs at 5 m intervals. The 10 cages within the exclosure had a mesh of 25 x 50 mm, and the other 10 cages had a mesh size of 51 x 76 mm. The larger mesh allowed prairie dogs to enter the cages. The cages were placed on the site in November 1979 (pasture rested in 1979), and in early August 1980 1.2 m° area was sampled in each cage. In addition, 10, 1.2 m* areas were sam- pled on either side of the exclosure fence mid- way between each agronomy cage. Forbs and grasses were bagged separately, oven dried, and weighed to the nearest gram. A Kruskal- Wallis one-way analysis of variance was used to statistically test for differences among graz- ing regimes. RESULTS A total of 112 prairie dog colony sites was found on the CMRNWR (Table 1). Ninety-six of these colonies were active, occupying a total of 2,122 ha and averaging 22 ha in size (se + 47 ha, range < 1 — 307 ha). Approximately 0.6% of the land area was inhabited by prairie dogs, with 2.8 active prairie dog colonies per 100 km’. On the FBIR, 42 active prairie dog colonies were surveyed totaling 2,786 ha (x = 66 ha, se + 91 ha, range 3 — 372 ha). Prairie dogs occupied about 2.1% of the survey area on the FBIR, with 3.0 colonies per 100 km’. The majority of prairie dog colonies both on the CMRNWKR and FBIR were located in ar- 200 GREAT BASIN NATURALIST Vol. 46, No. 2 TABLE 1. Distribution of prairie dog colonies found in association with livestock developments, natural bodies of water, homesteads, and roads and cattle trails. Percentage of colony sites located next to Study Livestock area n developments CMRNWR' 112 62 FBIR® 42 60 1Charles M. Russell National Wildlife Refuge. Fort Belknap Indian Reservation. eas of intensive livestock grazing and/or areas of topsoil disturbance by human activity. Livestock developments (reservoirs, wells, salt licks, and calf feeders) were found at 62% of the colony sites on the CMRNWR and 60% of the colony sites on the FBIR (Table 1). Nine colonies on the CMRNWR had salt licks or calf feeders in them; four of these colonies were located away from a stock watering area. Many colonies at the east end of the CM- RNWR were adjacent to the waters of Fort Peck Reservoir. On the FBIR, 29% of the colonies were found along perennial streams or around dry lakes. These sites were areas of intensive livestock grazing on both study ar- eas. Homestead activity was found at 26% and 17% of the colony sites on the CMRNWR and FBIR, respectively. Stock watering areas and areas formerly cultivated were frequently found at homestead sites. Only one small colony (< 1 ha) was found in a grain field currently under cultivation. Included in this group is a prairie dog colony on the CM- RNWR that started at a site where gravel was removed for road construction and another that started on a greasewood bottomland site that was mechanically cleared and leveled to make a pasture for horses at a refuge field station. Roads (usually two-track vehicle trails) and/or well-established cattle trails were found at 109 of the 112 prairie dog colony sites on the CMRNWRand at 41 of 42 colonies on the FBIR. Roads alone intersected 88% of the prairie dog colonies on the CMRNWR. Roads and cattle trails on both the CMRNWR and the FBIR interconnected livestock water- ing areas and homestead sites. In the northwest portion of the CMRNWR, the mean distance from the geographic center of 36 prairie dog colony sites to the nearest stock water development or homestead site (0.5 km) was less than (p < 0.001, t= 1084.36, Streams Trails & lakes Homesteads & roads 18 26 97 29 17 98 154 d.f.) the mean (1.3 km) for the 120 random points. Thirty of the 81 stock water develop- | ments in this area had colonies at them. The | quarter section locations of 1,772 observations _ of cattle groups recorded from 1973 to 1975 in | the Nichols Coulee allotment were not dis- | tributed homogeneously with respect to quar- — ter sections with prairie dog colonies (p < 0.05. X’=4.90, 1 d.f.). Cattle were observed with | greater than expected frequency on quarter — 1 sections with prairie dog colonies. However >) | the distribution of cattle observations on quar- | ter sections with reservoir sites lacking prairie _ | dog colonies was homogeneous to the distri- |, bution of cattle observations on quarter sec- | tions with prairie dog colonies (0.50 < p <_ 0.75, X°=0.16, 1 d.f.), suggesting that the © concentration of cattle on quarter sections | pyron habitat types jie both summer and | fall (Table 2). Use of shrub-grassland habitats | averaged 85% over both seasons for all years. |), These habitats, as determined from aerial. photos, composed only 54% of the two pas- | tures. More than three-fourths of the observa- | tions of cattle were on slopes with inclinations | of less than 11 degrees. Cattle, for the most | land habitats along the main ridge tops and ), major drainages where water developments |} (reservoirs and wells) had been established. |), All prairie dog colonies in these two pastures || were located in shrub-grassland habitat types Cee eae yron, 88%; ane were located primarily on Sone of | i; less than 7 degrees. Prairie dog colonies occu- } pied 2.8% of these two pastures, which was ), considerably above the average for the CM- |. RNWR. Number of colonies per 100 km? (8.9). S arcobatus- | | April 1986 TABLE 2. Use of habitat type and slope by cattle during summer and fall in the Nichols Coulee allotment. Percentage of observations Habitat type Summer Fall Artemisia-Agropyron 44 42 Sarcobatus-Agropyron 29 Sf Agropyron-Symphoricarpos 13 15 Pinus-Juniperus 14 16 Degrees of slope 0-10 81 79 11-25 14 13 26-35 5 7 36+ repeat 6 ——_______________» repeat 16 —————»> repeat ———> repeat = 91 14] MIke ST w~ Total examinations = 349 Owing to the mountainous terrain where the grazing sites were located and because of unpredictable time restraints, no schedule could be prearranged when the examining team would be at a specific camp site. Never- theless, in most cases owners were contacted at least one day in advance to encourage them to allow ample time for the clinic and to fast their dog for approximately 12 hr prior to the actual examination. Such a practice allowed for an evacuation of most of the dog’s intestinal contents and made the subsequent purgation less severe. ; At the time of the examination, the atten- dant veterinarian gave educational brochures to each owner and explained the process to be undertaken. Because approximately 10% of all dogs purged and examined at clinics in Utah over the past 12 years have shown some adverse reactions (spasms, labored breathing, incoordination, accelerated heartbeat, etc.), and since many of the highly trained sheep dogs were considered of relatively high mone- tary value, a release of responsibility form was signed by each owner before his dog could be examined. Also, a questionnaire was filled out identifying the owner's name and address, the number of sheep in each herd, and the name, breed, sex, and age of each dog to be exam- ined. Each owner and each dog were given identification numbers to be used in all subse- quent identifications and statistical tabula- tions. Each dog was individually tethered with a collar choke chain and given an oral dose of 1.5% arecoline hydrobromide (3 mg/kg of body weight) to induce purging. One or two additional doses were given if no response occurred within an approximate 20-30 210 GREAT BASIN NATURALIST TABLE 3. Prevalence of specific cestodes in sheep dogs examined, 1982-1984. Neg. E. g T.h Year No. (%) No. (%) No. (%) 1982 46 10 15 (N = 91) (50.5) (11.0) (16.5) 1983 86 3 8 (N = 141) (61.0) (2.1) (5.7) 1984 83 3 11 (N = 117) (70.9) (2.6) (9.4) Total 215 16 34 (N = 349) (61.6) (4.6) (9.7) Legend: Neg. = negative E.g. = Echinococcus granulosus T.h. = Taenia hydatigena T.o.k. = Taenia ovis krabbei T.p. = Taenia pisiformis T.s. = Taenia serialis 25 @ Echinococcus granulosus a ZL Taenia hydatigena Lu O Taenia ovis krabbei Fe 20 © Taenia pisiformis O HB 7aenia serialis Ts 15 z 5 10 LW mE a 0 1982 1983 1984 YEAR Fig. 1. Prevalence of cestodes in sheep dogs from central Utah, 1982-1984. minute period following the initial dose. After a purge occurred, any large tapeworms present were carefully collected from the ground and placed into bottles containing 10% formalin. Also, any clear mucoid portion of the purge that might contain the small E. granulosus tapeworms was carefully placed in the same bottle, which was then labeled with all pertinent information and stored for labo- ratory examination. After purgation, each dog was given a subcutaneous injection of prazi- quantel (Droncit®; 5 mg/kg of body weight), a drug known to be highly effective in the re- moval of E. granulosus tapeworms (Andersen et al. 1978, 1979). All samples were taken to the Parasitology Laboratory at Brigham Young University for examination of the collected material. The larger taeniids were transferred to new con- tainers and then shipped to Kansas City, Mis T.o.k T. p T.s No. (%) No. (%) No. (%) 13 22 5 (14.3 (24.2) (5.5) 21 17 13 (14.9 (12.1) (9.2) 12 8 6 (10.2 (6.8) (5.1) 46 47 24 (13.2 (13.5) (6.9) souri, where one of us (LAJ) identified the | worms to species. The remaining fluid in each | original sample bottle was divided into two equal portions and each part examined with a. variable 7X-30X stereozoom microscope by a O 6 6 s { different investigator to detect the small E. | granulosus tapeworms that might be present. After the examinations and identifications were completed for each year of study, each dog owner was sent a letter detailing the re-. sults for each of his dogs and, when tape- worms were found, was also given an indica-— tion as to the most likely food item his dog had eaten to acquire such a tapeworm (e.g., sheep. carcass, deer, rabbit, etc.). All data collected. for each owner and each dog were entered onto tabular sheets designed for the project. and then transferred to computer tapes. All } categorization and tabulation of data were — done on the Research VAX computer at BYU: . with the aid of Statistical Analysis Systems — (SAS) programs. Analyses or statistical associ: — ations of specific cestode infections were. tested for (1) effect of breed, sex, age of sheep’ dog, and number of sheep in individual herds | (2) occurrence in dogs owned by individual: who submitted animals for an examination ei- ther two or three years of the three-year study, period; and (3) coexistence within all sheer. dogs examined during the entire program. RESULTS | During the three-year study period, 266 separate sheep dogs were examined a total 0 { | ) | | +) { | {\ ‘| Ul i | @ iw | } April 1986 ANDERSEN ET AL.: CESTODE INFECTIONS IN Docs 211 TaBLE 4. Breed and sex of sheep dogs examined for cestodes, 1982-1984. Sex All dogs examined Infected dogs Breed Male Female Number Percent Number Percent Australian Shepherd 4 6 10 2.9 5 Oar Australian Shepherd cross 5 4 9 2.6 2 1B) Border Collie 39 ll 50 14.3 26 19.4 Border Collie cross ii 2 9 2.6 4 3.0 Blue Heeler 3 i 4 Tell 2 1.5 Blue Heeler cross 2 0 2 6 = all a Collie 61 35 96 Ts) 36 26.9 Collie cross 108 45 153 43.8 54 40.3 Dachshund cross 0 2 2 6 0 0 German Shepherd 1 0 1 8 0 0 German Shepherd cross 2 0 2 6 1 ofl Kelpie 2 5 il 2.0 3 3,0) ~- Poodle cross 3 0 3 9 0 0 Terrier cross 0 iL 1 3 0 0 Total 237 112 349 100.0 134 100.0 TABLE 5. Age of sheep dogs examined for cestodes, 1982-1984. pec ok Number of separate examinations Number Percent dog when Pp Z E Total of dogs of dogs examined 1982 1983 1984 examinations infected infected 0.5 10 DT 26 63 14 22RD, 1 19 21 17 ol 22 38.6 2 16 31 19 66 32 48.5 3 12 18 16 46 20 43.5 4 9 10 10 29 10 34.5 5) 4 10 8 OP) 10 45.4 6 3 6 6 NS 8 53.0 Hi 5 4 2 ll 5 45.4 8 4 3 3 10 4 40.0 9 3 2, 2 fl 2} 28.6 10 4 3 tS) 12 3 25.0 ll 1 2 2 ») 3 60.0 | | 12 0 3 0 3 0 0.0 | 13 0 0 1 1 1 100.0 15 1 1 0 2 0 0.0 | Total 91 141 117 349 134 38.4 349 times; 205 dogs were examined only once, 48 dogs were examined two of the three years, _and only 16 dogs were examined each of the three years (Table 1). Dogs were examined ' from 49 different owners, who had an average of 1,269 sheep each. Tables 2 and 3 give various statistics as to \the cestode infection levels identified within |the dog population. Overall, 134 of 349 exami- ‘nations (38.4%) showed at least one species of cestode present, whereas 215 examinations (61.6%) were negative. The level of infection in all examinations over the three-year period decreased from 49.5% in 1982 to 39.0% in 1983 to 29.1% in 1984 (Table 2). The infection levels for specific cestodes found over the three-year periods were: Echinococcus gran- ulosus, 16 of 349 positive, 4.6%; Taenia hy- datigena, 34 positive, 9.7%; T. ovis krabbei, 46 positive, 13.2%; T. pisiformis, 47 positive, 13.5%; and T. serialis, 24 positive, 6.9%. Specifically, the level of infection of E. granu- losus: dropped from 10 of 91 dogs positive (11.0%) in 1982 to 3 of 141 positive (2.1%) in 1983 and then rose slightly to 3 of 117 positive (2.6%) in 1984 (Table 3). The prevalence of 212 GREAT BASIN NATURALIST Vol. 46, No. 2 TABLE 6. Relative decline in cestode infections in sheep dogs examined, 1982-1984. Number of infections! Cestode 1982 1983 Echinococcus granulosus 10 3 Taenia hydatigena 15 8 Taenia ovis krabbei 1Te3} 21 Taenia pisiformis 22 17 Taenia serialis 5 13 Total: 65 62 ‘Some dogs infected with more than one species of cestode. *Statistically significant decrease (SAS program). 1984 Gamma Standard Total statistic error 3 16 —0.522* 0.006 ll 34 —0.208 0.156 12 46 —0.122 0.281 8 47 —0.440* 0.118 6 24 —0.047 0.176 40 67 Summary: —0.269 0.087 TABLE 7. Statistical associations (uncertainty coefficient; SPSSX) of specific cestode infections tested against breed, | sex, age of sheep dog, and number of sheep in individual herds, 1982-1984. Standard Uncertainty Variable Cestode coefficient* error Breed Echinococcus granulosus 0.074 0.046 Taenia hydatigena 0.055 0.031 Taenia ovis krabbei 0.057 0.022 | Taenia pisiformis 0.042 0.016 | Taenia serialis 0.042 0.017 Summary 0.029 0.013 | Sex Echinococcus granulosus 0.012 0.018 Taenia hydatigena 0.006 0.010 Taenia ovis krabbei 0.021 5 0.018 Taenia pisiformis 0.001 0.006 Taenia serialis 0.002 0.014 Summary 0.003 0.004 Age Echinococcus granulosus 0.164 0.046 Taenia hydatigena 0.084 0.038 Taenia ovis krabbei 0.077 0.032 Taenia pisiformis 0.061 0.024 Taenia serialis 0.061 0.028 Summary 0.047 0.016 Number of Echinococcus granulosus 0.141 0.049 Sheep Taenia hydatigena 0.111 0.036 Taenia ovis krabbei 0.143 0.028 Taenia pisiformis 0.047 0.022 Taenia serialis 0.092 0.019 Summary 0.066 0.022 *Uncertainty coefficient values represent the reduction in uncertainty (1 to 0) in predicting the specific cestode infection when the specific test variable (i.e., , breed, sex, age, or number of sheep) is known. All values are extremely low (i.e., indicate poor predictive capability). infection for each specific cestode detected over the three-year study period is depicted in Figure 1. The dog population surveyed could further be categorized into 14 different breeds (Table 4) and into 237 male and 112 female dogs. The most popular breeds examined were Border Collie (14.3%), Collie (27.5%), and Collie cross (43.8%). The age of the dogs was catego- rized into 15 age groups between 0.5 and 15 years of age (Table 5), with 66% of all examina- tions being done on dogs 3 years of age or less. The age group categorization of dogs in- fected with E. granulosus showed that of 16 total infections over the three-year period, 3! each were identified in dogs one, two or three’ years of age; 2 infections in four-year-old dogs, four infections in six-year-old dogs, and one — infection in an eight-year-old animal. i Statistical analyses of the trends in cestode , infections over the three-year study period — showed an over-all decrease (SAS; summary ! gamma statistic of —0.269), with individual — decreases for all parasite categories. The greatest single decline in any species was for E. granulosus (gamma statistic of —0.522, | Table 6). Additional statistical analysis , showed that there were no significant correla: | 1 a April 1986 ANDERSEN ET AL.: CESTODE INFECTIONS IN Dogs 213 TABLE 8. Statistical association (Wilcoxon Test) for cestode infections in sheep dogs of owners who submitted dogs for examination two of the three years of the study, 1982-1984. Number of Cestode owners! Echinococcus granulosus i Taenia hydatigena 10 - Taenia ovis krabbei sit - Taenia pisiformis 17 - Taenia serialis 5 Number of dogs examined lst exam 2nd exam P value 10 ] 0.036* 15 3 0.012* 14 a 0.036* 24 4 0.000* 6 a 0.109 SS eee ee eee !Owners identified in this table are those individuals who had dogs infected with specific tapeworms at an initial examination (either in 1982 or 1983) and then submitted dogs for examination also in a subsequent year (either in 1983 or 1984). *Significant decrease at < 0.05 level. TABLE 9. Statistical association (Quade Test) for cestode infections in sheep dogs of owners who submitted dogs for examination each year of study, 1982-1984. Number of dogs Numbecot examined Statistical significance* Cestode owners’ 1982 1983 1984 1982 vs. 1983 1982 vs. 1984 1983 vs. 1984 Echinococcus granulosus 5 8 0 1 yes yes no _ Taenia hydatigena U 10 2 3 yes yes no | Taenia ovis krabbei 10 13 9 6 no no no Taenia pisiformis ll 18 10 4 yes yes yes Taenia serialis 3 4 8 1 no no no 1Owners identified in this table are those individuals who had dogs infected with specific tapeworms at an initial examination in 1982 and then submitted dogs for examination also in 1983 and in 1984. *Significant decrease at < 0.05 level. TABLE 10. Statistical association (Fisher's Exact Test) on coexistence of specific cestodes in all sheep dogs examined, 1982-1984. Taenia | Cestode hydatigena | Echinococcus granulosus .002* \ Taenia hydatigena = | Taenia ovis krabbei = | Taenia pisiformis = | *Significant positive association at < 0.05 level. tions in any of the cestode infection categories ‘over the three-year study period for breed, sex, or age of sheep dogs involved, nor with number of sheep owned by any individual rancher (Table 7). For owners who submitted dogs for two annual examinations during the three-year study period there was a significant decrease in the infection levels noted for E. granulosus, T. hydatigena, T. 0. krabbei, and T. pisiformis (at 0.05 level; Table 8). For own- ers who submitted dogs for examination all three years of the study, a significant decrease tin infection levels was noted for two of the ithree years for E. granulosus and T. hydati- ygena and for all three years for T. pisiformis (at /0.05 level; Table 9). A significant positive as- sociation in the coexistence of specific ces- _todes detected in the study was seen for E. 1 bal Taenia Taenia ovis Taenia krabbei pisiformis serialis .244 .461 .302 .790 .793 .492 — 1.000 1.000 a — .029* granulosus and T. hydatigena and for T. pisi- formis and T. serialis (at 0.05 level; Table 10). DISCUSSION Epidemiological factors on hydatid disease for this region, such as anthelmintic treatment regimes used in the dogs, sheep management practices, owner awareness of the life cycle and transmission of hydatid tapeworms, and willing- ness of local individuals to cooperate with rec- ommended preventive and control measures for the region, had been previously evaluated (Schantz and Andersen 1980, Condie et al. 1981, Crellin et al. 1982, Andersen et al. 1983), and were not assessed in the present study. How- ever, from the data collected in this project, several conclusions or trends can be identified. 214 Although it was relatively easy to convince ranchers to have their dogs examined for one year of our study, it became increasingly diffi- cult to obtain their cooperation in subsequent years. A total of 19 of 49 separate owners submitted dogs for examination in only one of three years of the study, 10 owners had dogs examined two of three years, and only 20 own- ers submitted dogs all three years. Conse- quently, even though we examined 269 differ- ent dogs during the project, 205 were tested only one year, 48 for two years, and only 16 were examined each year of the three-year period. Part of the apparent reason for a de- crease in repeat examinations was the incon- venience for ranchers to submit their dogs for examination when they needed the dogs on a daily basis. Generally dogs subjected to the rather harsh purgation used in our study were unable to do hard work for several hours fol- lowing the examination, and sheep ranchers rapidly became aware of this complication. Furthermore, the field team of veterinarians could not always arrive at the camp site on a planned schedule, could not always find the herd in a specific locale, and sometimes could not traverse the mountainous roads which were on occasion interspersed with drifted snow even into mid-August. Another factor which complicated our be- ing able to examine more dogs in subsequent years was the rather high turnover of dogs in this particular industry. The age profile cate- gorization showed a marked cluster of dogs in the early age brackets, with 66% being three years of age or less. Reportedly, many dogs are lost or stray from the camp sites, some are purposefully destroyed when they do not prove to be easy to train, some are acciden- tally killed (e.g., being run over by owner's truck), and some cannot withstand the rigors of that type work for more than a few months at a time. Dogs in the latter category are rou- tinely taken to the home residence of the owner and are allowed to recuperate for sev- eral weeks before being returned to the mountain rangeland. This decrease in dogs examined on a year-to-year basis naturally complicated our attempts to accurately iden- tify the prevalence of cestodes in this popula- tion of animals and to properly evaluate our overall program. However, for those owners who did submit dogs for more than one year, GREAT BASIN NATURALIST Vol. 46, No. 2 | there was a significant reduction in specific _ cestodes identified (Tables 8 and 9), which | fact suggests that such owners were imple- | menting recommended preventive and con- | trol measures in their overall sheep and dog management. The study also showed that the same group | of cestode species was identified in these | sheep dogs for all three years, as shown in previous surveys (Andersen et al. 1973, Love- less et al. 1978, Jensen et al. 1982). Of particu- lar importance was the fact that the greatest individual reduction in prevalence of any one species of cestode was observed for E. granu- | losus (gamma statistic of —0.522) and suggests | that ranchers are indeed concerned about hy- | datid disease. The observed reduction in the. prevalence of T. hydatigena (gamma = —(0.208) should have paralleled more closely. that of E. granulosus, since sheep serve as the’ principal intermediate host in the life cycle of both parasites. Jensen et al. (1982) found that. 25 of 76 mule deer (32.9%) examined in cen-, tral Utah harbored cysticerci of T. hydati-| gena, which fact could account for the availability of an additional source of infectior (other than sheep) for dogs in that region. A! consistent drop in the infection level also oc. curred for T. pisiformis (gamma = —0.440).. where the numbers of infected dogs droppec, from 22 in 1982 to 17 in 1983 to 8 in 1984. Suck. a decline could have been due to a drop in the. rabbit (intermediate host) population, since. rabbits go through regular fluctuations ir numbers. We did not see any statistical correlation ir, this study of specific cestode infections with) breed, sex, and age of sheep dogs (Table 7) | However, we did note a significant positive — association on coexistence of E. granulosu. and T. hydatigena and also of T. pisiformi. and T. serialis (Table 10). The fact that the firs two species of cestodes utilize sheep as princi. pal intermediate hosts and that the seconc two use rabbits in that role corroborates the presumed dietary habits of the dogs harboriny these cestodes and adds additional validation to the specific identification of the cestode — reported over the three-year period. lf With the mountainous terrain in this region , and the large numbers of sheep present, iti unlikely that ranchers will ever be able t. identify all sheep that die or are killed on th: | ul April 1986 grazing sites, nor will they be able to burn or bury all carcasses. However, our observations _ over the past several years have indicated that more ranchers are now using an increased amount of commercial food for their dogs rather than expecting them to scavenge for food, and that fewer sheep are now butchered for routine consumption by the herders than in former years. These practices would defi- _nitely reduce the availability of sheep viscera to sheep dogs at the camp site. The probable advantage of using praziquan- tel in our program to treat all dogs in the region, irrespective of whether or not they proved to be infected with cestodes, is some- what difficult to evaluate with the design of our project, and in particular with the lack of | large numbers of repeat examinations. Inas- much as the time required for adult worms to develop and become patent after cystic mate- rial is eaten by the dog is only 40-45 days (Rausch 1975), and since our program moni- tored the dogs only once per year, we are unable to substantiate the known efficacy of praziquantel as demonstrated in experimen- tally infected dogs (Andersen et al. 1978, 1979, Andrews et al. 1983). Nevertheless, our data show that there was a definite decline in the overall cestode infection level during the three-year study period, and a more frequent program of drug administration may be war- ranted. The use of praziquantel in our study was a clear signal to the owners and herders that chemotherapy must be incorporated into -an overall program to prevent and control _ hydatid disease. Certainly the use of this par- ' | ticular drug would have had some contribu- "tory effect to the general downward trend in _ levels of cestode infection noted. Recent test- , ing in our laboratory on the efficacy of a new | paste formulation of praziquantel (Andersen » etal. 1985) will be an additional benefit to the ranchers in this particular region, because , they will soon be able to purchase and use that formulation rather than to rely on either tablet ; or injectable formulations. Tablets are fre- / quently difficult to administer to certain un- { ‘manageable dogs, and injectable formulations ‘require an assortment of needles and syringes i yor the services of an attendant veterinarian. «In spite of the difficulty in completing this » particular surveillance project, the general at- | titude and cooperation of the owners and | ANDERSEN ET AL.: CESTODE INFECTIONS IN Docs 215 herders remained positive throughout. This same attribute has been evident from the very beginning of the hydatid disease control pro- gram in Sanpete County in 1971 and is un- doubtedly the most important factor in what- ever level of success the program _ has achieved. Inasmuch as the control program we have attempted to implement has been an entirely voluntary campaign, it has been es- sential that all measures advocated or under- taken be promoted in a cooperative and friendly manner. Many local individuals such as community and church leaders, educa- tional administrators and teachers, public health workers, and local veterinarians and physicians have all contributed markedly to the success of the program. The incorporation of a filmstrip and coloring books on hydatid disease into the school curricula has definitely enhanced the educational aspect of the cam- paign (Crellin et al. 1982, Andersen et al. 1983). However, since the last surgical case for removal of hydatid cysts from an individual in Sanpete County occurred in 1979 (Crellin et al. 1982), it is important the complacency during an era of diminished public health em- phasis not be allowed to occur. In 1971, 20%-25% of the dogs in several communities were infected with E. granulosus (Andersen et al. 1973), whereas it is now virtually impos- sible to find infected dogs from these same communities. Also, in 1971 approximately 10% of all sheep slaughtered at local abattoirs in central Utah had hydatid cysts (Andersen and Wallentine 1976), whereas 14 years later the number of sheep reported to have hydatid cysts is well below 1% of those slaughtered, and only rarely is it possible to collect hydatid cysts from abattoirs in this region. In 1984 only three dogs infected with E. granulosus were identified along the Skyline Drive, yet one of the dogs had the heaviest natural infec- tion seen to date in this county. This dog could well have been a source of infection for at least several sheep grazing in that herd, which in five to seven years could be the potential source for continued transmission to other dogs. If such a focus of infection is perpetu- ated, the chance for eventual human cases originating from any one dog source would be greatly increased. Thus, the project now ter- minates with a possible “tip of the iceberg” still evident. 216 It now seems that most of the individuals living in Sanpete County have had ample op- portunity to become educated about hydatid disease and the distinct characteristics that allowed the disease to become endemic in that region. Whether or not a concerted effort will continue by both individuals and commu- nity leaders in attempting to implement rec- ommended preventive and control measures for hydatid disease in central Utah remains to be seen. Hydatid disease has been controlled or eradicated completely in parts of the world such as Iceland and on the Island of Cyprus, but extensive efforts to eradicate the disease elsewhere, such as in an endemic site in New Zealand, have not succeeded _ entirely (Gemmell 1979). Many of the unique charac- teristics and epidemiological determinants which were identified for Sanpete County (Crellin et al. 1982) still exist. The only major factor that has changed through time has been the increase in public awareness and basic knowledge on the life cycle of the causative parasite. It is hoped management practices for both dog and sheep populations in Sanpete County will be modified sufficiently as to in- deed decrease the potential for human cases in the future. LITERATURE CITED ANDERSEN, F. L., G. A. CONDER, AND W. P. MARSLAND. 1978. Efficacy of injectable and tablet formulations of praziquantel against mature Echinococcus granulosus. Amer. J. Vet. Res. 39: 1861-1862. . 1979. Efficacy of injectable and tablet formulations of praziquantel against immature Echinococcus granulosus. Amer. J. Vet. Res. 40: 700-701. GREAT BASIN NATURALIST ANDERSEN, F. L., J. R. CRELLIN, C. R. NICHOLS, AND P. M. Vol. 46, No. 2 | SCHANTZ. 1983. Evaluation of a program to control | hydatid disease in central Utah. Great Basin Nat. | 43: 65-72. ANDERSEN, F. L., J. S. SHORT, AND H. D. McCurpy. 1985. | Efficacy of a combined paste formulation of prazi- quantel/febantel against immature Echinococcus | granulosus and immature Echinococcus multiloc- | ularis. Amer. J. Vet. Res. 46: 253-255. ANDERSEN, F. L., AND M. W. WALLENTINE. 1976. Hydatid | disease. Nat. Woolgrower 66: 16-18. ANDERSEN, F. L., P. D. WRIGHT, AND C. MORTENSON. 1973. Prevalence of Echinococcus granulosus in- fection in dogs and sheep in central Utah. J. Amer. Vet. Med. Assoc. 163: 1168-1171. | ANDREWS, P., H. THOMAS, R. POLKE, AND J. SEUBERT. 1983. Praziquantel. Med. Res. Rev. 3: 147-200. CONDIE, S. J., J. R. CRELLIN, F. L. ANDERSEN, AND P. M. | SCHANTz. 1981. Participation in a community pro- | gram to prevent hydatid disease. Publ. Hlth. Tom don 95: 28-35. | CRELLIN, J. R., F. L. ANDERSEN, P. M. SCHANTZ, AND S. J. ConDIE. 1982. Possible factors influencing distri- | bution and prevalence of Echinococcus granulo- | sus in Utah. Amer. J. Epidemiol. 116: 463-474. GEMMELL, M. A. 1979. Hydatidosis control—a | view. Aust. Vet. J. 55: 118-125. JENSEN, L. A., F. L. ANDERSEN, AND P. M. SCHANTZ. 1982. The prevalence of Echinococcus granulosus and other taeniid cestodes in sheep dogs of centaalt Utah. Great Basin Nat. 42: 65-66. JENSEN, L. A., J. A. SHORT, AND F. L. ANDERSEN. 1982. Internal parasites of Odocoileus hemionus in cen- tral Utah. Proc. Helminthol. Soc. Washington 49 317-319. LOVELESS, R. M., F. L. ANDERSEN, M. J. RAMSAY, AND R. K | HEDELIUS. 1978. Echinococcus granulosus ir dogs and sheep in central Utah, 1971— 1976 Amer. J. Vet. Res. 39: 499-502. Rauscu, R. L. 1975. Taeniidae. Pages 687-707 in Hub: bert et al., eds., Diseases transmitted from ani-. mals to man. Thomas Books, Springfield, Illinois. | SCHANTZ, P. M., AND F. L. ANDERSEN. 1980. Dog owner: | and hydatid disease in Sanpete County. Grea’ | Basin Nat. 40: 216-220. DAM-RAISED FAWNS, AN ALTERNATIVE TO BOTTLE FEEDING Kathrin M. Olson-Rutz', Philip J. Urness!”, and Laura A. Urness! ABSTRACT. —Rearing young ungulates for ecological studies is costly and time consuming. Doe-rearing mule deer | (Odocoileus hemionus) fawns is a viable alternative to the common method of bottle-feeding. Fawns tamed while ) nurtured by their tractable dams showed no marked difference in tractability over bottle-reared orphans. The » advantages of doe-rearing are better health for the young and convenience for the handler. The use of tame animals to obtain informa- tion on the foraging behavior and habitat se- lection of wild ungulates is increasing. This technique has been used with many native ‘North American species (Reichert 1972) as swell as several from Africa (Hutchison 1970). ‘Close observation of foraging animals offers \more precise dietary assessment than fecal/ rumen sample analyses or distant observa- | tion, yet time and monetary investments re- quired to rear and train experimental animals vare high. Several methods for successfully / rearing tractable animals have been reported. The most common is bottle-raising captured ‘neonates (Schwartz et al. 1976, Hobbs and | Baker 1979, Addison et al. 1983) or the young born to either penned wild or tame dams (Rei- ‘chert 1972, Knorre 1974). Another option is taming wild-caught yearlings, as discussed by ‘Kreulen (1977). _ The Utah Division of Wildlife Resources ‘has maintained a tame mule deer (Odocoileus ‘hemionus) herd for more than 30 years by )sottle-feeding fawns. In the last three years 14 vawns have been tamed while nurtured by | heir dams. We report this as a viable alterna- ‘ive to bottle-rearing of young, with advan- ages in better health and convenience. REARING The herd was maintained in a 1-ha com- ound on the foothills of the Bear River Range 't Logan, Utah. Beginning in early June the loes were carefully observed for signs of im- vending parturition. They were then put into yeparate 5 x 25 m outdoor runs each with an | ——————— Hl ‘Department of Range Science, Utah State University, Logan, Utah 84322. “Utah Division of Wildlife Resources, Logan, Utah 84322. open-air shed and hay bedding. The fencing must be stout to withstand aggressive behav- iors between does and of a fine mesh wire on the bottom meter to prevent fawns from es- caping. The sheds were cleaned of soiled bed- ding daily to reduce development and spread of disease. Green alfalfa hay, barley, bal- anced-ration pellets, and clean water were available ad libitum. To supplement the sparse forage growing in the runs, the animals were given freshly clipped mixed forbs (mostly alfalfa) morning and evening for a month postpartum and whenever a source was available thereafter. As noted by Schwartz et al. (1976) for pronghorn (Antilo- capra americana) fawns, the deer fawns also consumed small quantities of soil beginning at about one week of age. When possible, births were attended to as- sure the health of both fawn(s) and doe and to ensure that the fawn(s) obtained colostrum. If the doe did not attend the young, they were removed and subsequently _bottle-raised. However, the inability to provide milk alone did not dictate bottle-rearing. The doe’s lick- ing of the perianal region stimulated the fawn to seek out a teat (or nipple) and nurse. This response could be used to advantage when supplemental bottle-feeding was required. One doe, with mastitis, cleaned her fawn while it nursed from a bottle. This doe-han- dler cooperation carried through until wean- ing. During the first week does were kept with their fawns 24 hours a day. The young were exposed to a minimum of 3 hours human con- tact and gentle handling per day. By the sec- 2 218 ond week does were turned out in the morn- ing, returned at noon to clean and nurse their young, turned out again in the afternoon, and put in for the night in the evening. Generally, from the third week until fawns were weaned at three months, the dams were eager to get out in the mornings and were left out until evening. Any fawn handling or training could proceed as described by others (Reichert 1972, Parker et al. 1984). HEALTH The most common diseases encountered were of intestinal microbes causing diarrhea leading to rapid dehydration, emaciation, and death. Although Kramer et al. (1971) dis- cussed the occurrence of Escherichia coli in mule deer, and Schwartz et al. (1976) found Clostridium perfringens to be a problem in pronghorn, our major concern was with Coc- cidia spp. Upon detection of this protozoan, 12.5% sodium sulfamethazine was used to prevent and treat the Coccidia infections. The drinking water was treated for two days with 8ml/] water (1 0z/gal) at time of birth, at one week postpartum (when the fawns begin drinking water), at two weeks, and anytime thereafter when loose or watery feces were noticed. A change in the character of the feces is the cue to an intestinal infection. For a more detailed discussion of the normal changes that the feces of young growing fawns undergo see Schwartz et al. (1976). Although sulfamethazine is commonly used on livestock, Schwartz et al. (1976) noted that the drug may crystalize in the urine and kid- neys of young animals. As an alternative they recommended the use of Sulfaquinoxaline. Sulfamethazine was effective in controlling di- arrhea in all the nine fawns treated and we have, as yet, experienced no adverse effects. We do, however, recommend caution in the use of this drug. In their evaluation of fawn-rearing proce- dures, Halford and Alldredge (1978) con- cluded that doe-reared fawns had no health advantage over those bottle-raised. They ex- perienced 67% (6 of 9 total) mortality of dam- raised fawns to necrobacillosis (Fusibacterium necrophorum), whereas the mortality of hand-raised fawns was only 33% (3 of 9 total), entirely due to E. coli and Streptococcus spp. GREAT BASIN NATURALIST umbilical infections. Unlike the hand- raised fawns, however, losses due to this disease, as well as many 6 of the 9 dam-raised fawnel| | (67%) were: (1) kept in pens with no forbs or | grasses available, (2) at higher animal densi- | ties, and (3) nursing does that had been on> deficient diets. As reviewed by Hibler (1981), | necrobacillosis is often associated with poor range and crowded conditions. Therefore, the | others, may well be averted under better con- | ditions. DISCUSSION Over three years 14 fawns have been raised | by does, and we bottle-reared 7 orphans. There was no notable difference in the’ tractability of the animals reared by these two’ methods, but there was a marked difference Vol. 46, No. 2 | in favor of dam-reared fawns in their stature as _ yearlings and two-year-olds. This was particu-|_ larly noticeable in those raised as singles’ rather than twins by their dam. If given < choice, raising singles is preferable. They ex:' hibited a faster growth rate and were gener | ally more robust than twins. In addition, the! lactation drain on the doe was greatly re! duced. Our visual assessment agreed with Halfore. and Alldredge (1978), who reported signifi’ Al cantly higher (P < .001) mean body weight’ ‘ and growth rates of fawns raised by their dam‘ as compared to those bottle-reared. Our year’ ling bucks were equal to or larger in statur’ than the bottle-raised two-year-olds and wer of substantially heavier build than their bot tle-raised cohorts. A more quantitative indice tion of physical condition is the minimur breeding age of females (Mackie et al. 1982) Of two doe fawns sired by the same buck an. raised concurrently, the dam-raised one gav. birth to a fawn at one year of age. This is a rar occurrence and was not matched by her boi tle-fed half sister. There are two major advantages of doe’ rearing fawns: (1) health—there is no subst) tute for the dam’s nurturing, species specifi. colostrum, and doe’s milk, which has twic the nutritional value of cow's milk (Sho: 1981), and (2) time—time and inconvenienc spent in cleaning and preparing bottles thre to five times daily is eliminated, thus allowin more time for direct contact with the young. | 4 April 1986 It is unknown whether a key period for imprinting on a handler exists. Our fawns were first exposed to humans between 0 and 24 hours after birth. The animals were pre- dominately handled by two people, yet were in frequent contact with others. Several au- thors stress the bond formation between han- dlers and bottle-raised young (Schwartz et al. 1976, Addison et al. 1983). Without the de- pendence on a handler for feed, the develop- -ment of confidence between handlers and dam-reared fawns is very important. Initially, _preferential behavior was exhibited toward the handlers; yet, amity or distrust did de- velop toward anyone with whom the animals had contact. The fawns’ response to individu- als gradually moderated through their first year. The work reported herein was done with ' fawns born to tractable does. The presence of tame conspecifics eases the handling of new animals (Kreulen 1977). Some _ species, though, may not be suited for this method of rearing. As part of a project involving white- tailed (Odocoileus virginianus), mule, and black-tailed (O. h. columbianus) deer in New | | | i 3 | | i Hampshire, an effort was made to raise two .sets of twin white-tailed deer fawns on their dams. The does were the most tame of the herd; however, their fawns were never ap- »proachable despite constant human contact. | One set eventually brought about their own deaths in panicked flight (P. Pekins, personal communication). In time the adaptable spe- cies will be known. Until then dam-raising young should be considered as an option when rearing animals for ecological studies. ACKNOWLEDGMENTS We thank M. Urness for assistance in rear- ing of fawns; K. Udy and M. Powell for veteri- “nary advice; and J. C. Malechek, F. D. 1") / OLSON-RUTZ ET AL.: MULE DEER FAwns 219 Provenza, and M. L. Wolfe for helpful sugges- tions and review of this manuscript. Facilities, animals, and funding were provided by the Utah Division of Wildlife Resources through Federal Aid Project W-105-R. LITERATURE CITED ADDISON, E. M., R. F. MCLAUGHLIN, AND D. J. FRASER. 1983. Rearing moose calves in Ontario. Alces 19: 246-270. HALForD, D. K., AND A. W. ALLDREDGE. 1978. A method for artificially raising mule deer fawns. Amer. Midl. Nat. 100: 493-498. HIBLER, C. P. 1981. Diseases. In O. C. Wallmo, ed., Mule and black-tailed deer of North America. Univer- sity of Nebraska Press, Lincoln. 605 pp. Hopss, N. T., AND D. L. BAKER. 1979. Rearing and training elk calves for use in food habits studies. J. Wildl. Manage. 43: 568-570. Hutcuison, M. 1970. Artificial rearing of some East African antelopes. J. Zool. 161: 437-442. KNorrE, E. P. 1974. Changes in the behavior of moose with age and during the process of domestication. Le Naturaliste Canadien 101: 371-377. KRAMER, T. T., J. G. NaGy, AND T. A. BARBER. 1971. Di- arrhea in captive mule deer fawns attributed to Escherichia coli. J. Wildl. Manage. 35: 205-209. KREULEN, D. K. 1977. Taming of wild-captured wilde- beeste for food habit studies. J. Wildl. Manage. 41: 793-795. MackIE, R. J., K. L. HAMLIN, AND D. F. Pac. 1982. Mule deer. In J. A. Chapman and G. A. Feldhammer, eds., Wild mammals of North America—biology, management, and economics. Johns Hopkins Uni- versity Press, Baltimore. 1,147 pp. PaRKER, K. L., C. T. ROBBINS, AND T. A. HANLEY. 1984. Energy expenditures for locomotion by mule deer and elk. J. Wildl. Manage. 48: 474-487. REICHERT, D. W. 1972. Rearing and training deer for food habits studies. U.S. For. Serv., Rocky Mtn. For. Range Exp. Stn. Res. Note RM-208. ScHWaRTz, C. C., J. G. NAGy, AND S. M. Kerr. 1976. Rear- ing and training pronghorns for ecological studies. J]. Wildl. Manage. 40: 464-468. SHort, H. L. 1981. Nutrition and metabolism. In O. C. Wallmo, ed., Mule and black-tailed deer of North America. University of Nebraska Press, Lincoln. 605 pp. SUBSPECIFIC IDENTITY OF THE AMARGOSA PUPFISH, CYPRINODON NEVADENSIS, FROM CRYSTAL SPRING, ASH MEADOWS, NEVADA Jack E. Williams’ and James E. Deacon” ABSTRACT.—Samples of pupfish from Crystal, Marsh, and Point of Rocks springs, Ash Meadows, Nevada, were examined to determine the subspecific identity of Cyprinodon nevadensis presently inhabiting Crystal Spring. | Meristic and morphometric analyses indicate that Crystal Spring is inhabited by C. n. mionectes. The presence of this subspecies is most likely explained by their precarious survival in the spring’s outflow after they were eliminated by — transplanted largemouth bass in the spring pool, and their subsequent reestablishment throughout the spring system _ after the extirpation of the bass. Crystal Spring (= Big Spring of Miller 1948) is the type locality for the Ash Meadows pup- fish, Cyprinodon nevadensis mionectes Miller. Crystal Spring was chosen by Miller (1948) as the type locality because its pupfish population “has characters which very closely approach the average for the subspecies as determined by an analysis of all populations.” In recent years, however, the subspecific identity of the pupfish in Crystal Spring has been questioned. On 1 January 1966, J. E. Deacon, C. L. Hubbs, and R. R. Miller searched Crystal Spring for pupfish and found none ( J. E. Deacon, field notes; Miller 1969). However, at least 10 transplanted largemouth bass, Mi- cropterus salmoides, were seen in the main spring pool. The pupfish population “reap- peared” by early February 1975 (Liu and Soltz 1983) and was later described as in “fine shape’ with a population of approximately 1,500 pupfish (Hardy 1980). Two subspecies of Cyprinodon nevadensis occur in Ash Meadows. In addition to its pres- ence in Crystal Spring, C. n. mionectes occurs in a variety of lower-elevation springs (Miller 1948, Soltz and Naiman 1978). Among other springs, Cyprinodon n. mionectes occurs in Jack Rabbit, Point of Rocks, the Bradford Springs, and springs at the northern end of Ash Meadows that discharge water into the formerly vast Carson Slough area. Several of these springs, and an introduced population of C. n. mionectes at Collins Ranch (Baugh et 1U.S. Fish and Wildlife Service, 2800 Cottage Way, Room E-1823, Sacramento, California 95825. Department of Biological Sciences, University of Nevada, Las Vegas, Nevada 89154. al., in press), are within 3 km of Crystal Sante, Cyprinodon n. pectoralis also occurs | in nearby springs, such as Indian, Marsh, School, and Scruggs. The population of C. n. | pectoralis from Indian Springs was particu-— larly suspect as a source for the Crystal Spring | population because that spring’s outflow fre- | quently discharges into the outflow of Crystal | Spring. Both subspecies of Cyprinodon nevadensis are listed as endangered by the. U.S. Fish and Wildlife Service. The potential for surface water connection | among the various springs is compounded by periodic flash floods, which may distribute | pupfish some distance from their usual habi- tat, and by the formation of Crystal Spring Reservoir, which is fed by outflow water from Crystal Spring. Thus, at least three hypotheses can be em- ployed to explain the recurrence of pupfish in Crystal Spring: 1. pupfish from another spring reachec | Crystal Spring by surface water connec: tion, 2. pupfish from another spring were intro-. duced into Crystal Spring by man, or 3. the pupfish in Crystal Spring were noi — eliminated by the largemouth bass bu’ only reduced to such low numbers tha’ they appeared to be extirpated. | Because of the geographic proximity 0 | other springs, either of the first two hypothe: ses could explain the presence of either C. n | 220 is April 1986 mionectes or C. n. pectoralis in Crystal Spring. The latter hypothesis, of course, would only be appropriate for explaining the presence of C. n. mionectes. The purposes of this report are to determine the subspecific identity of C. nevadensis presently inhabiting Crystal Spring and to explain their occur- rence. MATERIALS AND METHODS Meristic and morphological characters were utilized to compare the unknown C. nevadensis from Crystal Spring to populations of C. n. mionectes from Point of Rocks Springs and C. n. pectoralis from Marsh Spring. The two previous taxonomic studies of C. nevadensis from Ash Meadows have docu- mented that the number of pectoral fin rays and number of preopercular pores in the cephalic lateral-line system are diagnostic in separating the two subspecies (Miller 1948, Miller and Deacon 1973). These two charac- ters plus the structure of the preopercular canal were used to determine the identity of the Crystal Spring population. Thirty specimens longer than 25 mm SL were analyzed from each of the following three collections of C. nevadensis made 17 January 1985 by T. M. Baugh and J. W. Pe- dretti: 1. UMMZ 213444. 59 Cyprinodon neva- densis ssp. (26.6-37.9 mm SL) from Crystal Spring. 2. UMMZ 213445. 60 Cyprinodon neva- densis mionectes (22.8-35.6 mm SL) _ from Point of Rocks Springs. 3. UMMZ 213446. 60 Cyprinodon neva- densis pectoralis (21.2-39.3 mm SL) from Marsh Spring. Samples from Point of Rocks and Marsh ~ Spring were chosen because of their geographic proximity to Crystal Spring. Also, Marsh Spring is adjacent to Indian Springs and should closely represent any C. n. pectoralis that may have entered Crystal. No meristic data are available i. it 7 \ i for C. n. pectoralis from Indian Springs. The “Indian Spring” referred to in Miller and Deacon (1973: Fig. 1 and text) clearly is Marsh Spring. The two spring systems are in close proximity. Marsh Spring is located more northerly and its outflow is partially impounded by a small reser- Voir. WILLIAMS, DEACON: AMARGOSA PUPFISH 22 The methods of Miller (1948) were em- ployed with the exception that only the left pectoral fin rays were counted. Counts of the left and right preopercular pores were sepa- rated from some analyses so that a count of 7-6, for example, refers to a specimen with 7 preopercular pores on the left side of the head and 6 preopercular pores on the right. Other- wise, counts on the left and right sides were combined. RESULTS AND DISCUSSION After all counts were completed, results from known samples of Cyprinodon n. mio- nectes and C. n. pectoralis were compared to previous results of Miller (1948) and Miller and Deacon (1973). Results from all studies were expected to be very similar because the methodology for performing the counts was nearly identical. Counts of the diagnostic meristic characters of the Point of Rocks Springs population of C. n. mionectes were very similar to those given by Miller (1948) (Tables 1 and 2). Pectoral fin rays were modal at 16 in both studies and averaged 15.68 for Miller (1948) and 15.73 (present study). Counts of preopercular pores were modal at 12 for both studies but aver- aged slightly higher in Miller (1948) than in this study (12.54 v. 12.17, respectively). Counts of the diagnostic meristic characters of the Marsh Spring population of C. n. pec- toralis also were very similar to the earlier studies (Tables 1 and 2). The number of pec- toral fin rays were modal at 17 and averaged 16.61 (Miller and Deacon 1973) and 16.50 (present study). Although the presence of 17 v. 16 pectoral fin rays is the primary character that separates pectoralis from mionectes , the Marsh Spring population of C. n. pectoralis has a higher frequency of individuals with 16 pectoral fin rays than other populations within the subspecies (Miller and Deacon 1973). Pre- vious data on preopercular pore counts for the Marsh Spring population were lacking, so data from this study were compared to preo- percular pore counts of the typical form of the subspecies from School Spring. Preopercular pore counts averaged 13.36 in the School Spring sample (= Lovell’s Spring of Miller 1948) and 13.30 in the Marsh Spring sample (present study). GREAT BASIN NATURALIST Vol. 46, No. 2 | TABLE 1. Comparison of pectoral fin-ray counts of Cyprinodon nevadensis from three springs in Ash Meadows, Nye. County, Nevada. 13 Crystal Spring Miller 1948: Table 16, 1942 present study, 1985 Point of Rocks Miller 1948: Table 16, 1942 1 present study, 1985 Miller and Deacon 1973:137 present study, 1985 Springs Marsh Spring | 14 15 16 17 18 x 8 102 112 11 15.54 1 10 18 1 15.63 3 65 112 12 15.68 2 8 16 4 15.73 24 30 2 16.61 1 13 16 16.50 TABLE 2. Comparison of preopercular pore counts of Cyprinodon nevadensis from four springs in Ash Meadows, Nye County, Nevada. 10 Miller 1948: Table 26, 1942 present study, 1985 Miller 1948: Table 26, 1942 1 present study, 1985 Miller 1948: Table 26, 1939 present study, 1985 Crystal Spring Point of Rocks Springs School Spring Marsh Spring A comparison of pectoral fin-ray numbers in the three 1985 collections indicates that the Crystal Spring population is C. n. mionectes (Table 1). A similar conclusion is reached when comparing the number and frequency of preopercular pore counts (Table 2). The Crystal Spring sample of C. nevadensis aver- aged 15.63 pectoral fin rays and 12.23 preop- ercular pores. Both values are within the re- sults expected for C. n. mionectes. The Crystal Spring sample of C. nevadensis was modal at 16 pectoral fin rays but included a relatively large percentage (33%) of individ- uals with 15 pectoral fin rays. This contrasts sharply with the Marsh Spring sample of C. n. pectoralis, which was modal at 17 pectoral fin rays and contained a relatively large percent- age (43%) of individuals with 16 rays and only one fish with 15. ; The Crystal Spring sample of C. nevadensis averaged 12.23 preopercular pores. This value is very similar to the average number of preopercular pores in the Point of Rocks Springs sample (12.17) but is quite distinct from the average in the Marsh Spring sample (13.30). The modal number of preopercular pores in the Crystal Spring sample is 12. This agrees with the value from the Point of Rocks sample but again contrasts with results from Marsh Spring, where those pores are modal at 13 (Table 2). Preopercular pore counts, with frequency of counts given parenthetically, are as follows for Point of Rocks: 5-6 (1), 6-5 (2), ll 12 13 14 15 16 x 1 39 18 8 12.50 3 18 8 l 12.23 2 55 20 18 12.54 | 3 21 5 0 1 12.17 l 24 18 54 1 9. 13.36mmI 1 7 10 8 2 2 13.30m a | 6-6 (21), 6-7 (3), 7 AS ), 8-7 (1); Crystal: 5-6 (2 | 6-571) 6 oe 7 (2), 7-6 (6), 8-6 (1); and Marsh: 5- 6 (1 \, 5 7(1 ), 6-6 (6), 6-7 (5), 7-6 (5), | 7-7 (8), 8-7 (2), and 8-8 @). | The structure of the preopercular canal also. varied significantly between the Point of Rocks Springs and Crystal Spring samples, i, and the C. n. pectoralis from Marsh Spring. | In the Marsh Spring sample, 13% of the fish | possessed at least one interrupted preopercu- | interrupted in one specimen. Although inter- a ruption of the canal results in a higher pore count, the primary causal factor in the higher | pore counts in the March Spring population — was the presence of a seventh pore on the anterior end of the preopercle. Most C. n. mionectes contain only 6 pores on each side. | : lar canal. Both the left and right canals were 4 : H No preopercular canal interruption was noted ~ in) C27: Crystal springs, although one specimen from| mionectes from Point of Rocks or | Crystal Spring possessed an open canal on one a side. Delays in completion of the cephalic cant system until adulthood have been noted for certain cyprinodonts (Hubbs and Miller 1965) and could possibly explain some of the abou ji differences noted between C. n. mionectes | and C. n. pectoralis. Differences in standard | length among the subsamples examined, however, are slight. Ranges and means ol standard length for the subsamples examined are: Point of Rocks Springs, 26.8—35.6 mm @ | | ih April 1986 = 30.98); Crystal Spring, 28.3-37.9 (33.26); and Marsh Spring, 25.3-39.3 (29.48). Finally, the meristic characters of the Crystal Spring sample were compared to data from the original study of the Crystal Spring population by Miller (1948). Very close agreement is seen in the average number of pectoral fin rays observed in the Crystal Spring sample taken in 1942 (Miller 1948) and in 1985 (present study) (Table 1). The 1942 sample averaged 15.54 rays, whereas the 1985 sample averaged 15.63. A sim- ilar agreement is seen in the number of preoper- cular pores from samples collected in 1942 and 1985 (Table 2). The 1942 sample included 59% of individuals with 12 preopercular pores and 27% with 13. The 1985 sample contained 60% of indi- viduals with 12 preopercular pores and 27% with 13. Based on the results of this study, the sub- specific identity of the pupfish from Crystal Spring is C. n. mionectes. It appears that the present C. n. mionectes population is descended from those individuals who withstood the pres- sure from largemouth bass. The question as to whether the present Crystal Spring population persists by natural occurrence, was introduced, or made its way by surface water connection from another C. n. mionectes population may be answered as follows. Although the predaceous largemouth bass apparently eliminated C. n. mionectes from the spring pool at Crystal, it may be that some pupfish survived in the spring out- flow. According to field notes of one of the au- thors (JED 66-1), only 100 yards of the outlet ditch were searched. Robert R. Miller (10 Octo- - ber 1985 in litt. to JEW) supported the hypothe- sis that C. n. mionectes may not have been elimi- nated from the entire spring system and stated, in part: to say unequivocally that Cyprinodon was gone from this system [Crystal] would be awfully hard to defend. In frigid midwater [ice on ground, | January 1966] you might - not expect to see pupfish, especially when they had been impacted for X-years by that dastardly predator Mi- cropterus salmoides. Granted that the pupfish could cer- tainly have been wiped out at the main spring pool. I think it is unjustified to assume they were eliminated throughout the long outlet ditch as well. | The causes of disappearance of largemouth bass from Crystal Spring are uncertain as well. 1 et NT ara WILLIAMS, DEACON: AMARGOSA PUPFISH 223 The warm water (30.9 C) of the spring pool may have inhibited reproduction, though such would not be the case throughout the cooler outlet ditch and reservoir. This, cou- pled with constant angling pressure by the workers in Ash Meadows, may have elimi- nated bass from the spring system. Regardless of the causes, it is quite reassuring to know that the type locality of Cyprinodon nevaden- sis mionectes still harbors that unique form of pupfish and that the spring has rid itself of the introduced bass. ACKNOWLEDGMENTS We appreciate the efforts of Tom M. Baugh and John W. Pedretti in collecting the pupfish used in this study. Reviews of this paper by Robert R. Miller, Cynthia D. Williams, and Donna L. Withers are gratefully acknowl- edged. Also, permission to quote from Miller's insightful missive is appreciated. LITERATURE CITED BauGu, T. M., J. E. WiLiaMs, D. A. BUCK AND J. E. DEa- CON. New distributional records for Cyprinodon nevadensis mionectes, an endangered pupfish from Ash Meadows, Nevada. Southwestern Natur. In press. Harpy, T. 1980. The Inter-basin area report 1979. Proc. Desert Fishes Council. 11(for 1979): 5-21. Husss, C. L., AND R. R. MILLER. 1965. Studies of cyprinodont fishes. XXII. Variation in Lucania parva, its establishment in western United States, and description of a new species from an interior basin in Coahuila, Mexico. Misc. Publ. Mus. Zool., Univ. Michigan. 127: 1-104. Liu, R. K., AND D. L. Sotz. 1983. [Letter to E. P. Pister dated 19 February 1975]. Proc. Desert Fishes Council. 7(for 1975): 209-210. MILLER, R. R. 1948. The cyprinodont fishes of the Death Valley system of eastern California and southwest- ern Nevada. Misc. Publ. Mus. Zool., Univ. Michi- gan. 68: 1-155. _ 1969. Conservation of fishes of the Death Valley system in California and Nevada. Cal-Neva Sec. Wildlife Soc. Trans. 1969: 107—122. MILLER, R. R., AND J. E. DEACON. 1973. New localities of the rare Warm Spring pupfish, Cyprinodon nevadensis pectoralis, from Ash Meadows, Ne- vada. Copeia. 1973: 137-140. Sotz. D. L., ANDR. J. NAIMAN. 1978. The natural history of fishes in the Death Valley system. Nat. Hist. Mus. Los Angeles County Sci. Ser. 30: 1-76. TWO ABERRANT KARYOTYPES IN THE SAGEBRUSH LIZARD : (SCELOPORUS GRACIOSUS): TRIPLOIDY AND A “SUPERNUMERARY’ ODDITY Pamela Thompson'* and Jack W. Sites, Jr.’ ABSTRACT. —Widespread karyotypic sampling in the lizard Sceloporus graciosus Baird & Girard has confirmed previous reports of chromosomal monotypy. Most individuals throughout the range have a diploid karyotype of 2N=30 | consisting of 12 biarmed macrochromosomes and 18 microchromosomes. A single female karyotyped from the vicinity of Riverside, California, was unmistakably triploid, showing 3N=45 with 18 macrochromosomes and 27 microchromo- somes. This female appeared phenotypically normal but appeared reproductively incompetent. A male from Zion National Park, Utah, showed an extra bivalent in some diakinesis arrays, which apparently represents a supernumerary chromosome. A major study is currently in progress to determine the population genetic structure of the chromosomally monotypic sagebrush lizard, Sceloporus graciosus Baird & Girard. In this study 112 individuals from seven west- ern states (AZ, CA, CO, ID, NV, UT, WY) were karyotyped to confirm chromosomal monotypy, and all but two individuals showed the previously reported 2N=30 normal kary- otype (Cole 1971, 1975a, Jackson and Hun- saker 1970). One showed a triploid karyotype, and another possessed an extra bivalent in diakinesis spreads. The former represents the third known case of unusual triploidy in a species of Sceloporus, and the latter may be an example of a supernumerary chromosome. METHODS Lizards were captured alive by noosing or by stunning with rubber bands. Karyotypes were prepared according to Baker et al. (1982) from testes, in the case of reproductively ac- tive males, or from bone marrow in the case of juveniles, females, or postreproductive males. Prior to karyotyping the lizards were injected with a yeast-sugar solution (Cole and Leavens, 1971) to increase the mitotic index. Treated suspensions of bone marrow or testes cells were dropped onto clean microscope slides and stained with a 6% Giemsa phos- phate solution (Patton 1967). At least five metaphase spreads were scored on most indi- viduals. ‘Department of Zoology, Brigham Young University, Provo, Utah 84602. Department of Biology, Idaho State University, Pocatello, Idaho 83209. bo RESULTS showed the normal 2N=30 karyotype, which consisted of 12 metacentric or submetacentric macrochromosomes (12M) and 18 microchro- mosomes (18m) (Fig. 1A). Macrochromosome pair two in metaphase cells with elongate chromosomes frequently showed terminal satellites (see arrow Fig. 1A) similar to those reported by Cole (1971) for this species. One adult female showed a triploid karyotype, 3N=45, consisting of 18 meta- or submeta- centric macrochromosomes, and 27 mi- crochromosomes. (Fig. 1B). Diakinesis arrays from testes of 90 repro-| ductively active males almost always con- sisted of six macrochromosomal and nine mi- — crochromosomal bivalents (Fig. 1C). One y! adult male, however, showed one extra mi-. a crochromosomal bivalent in seven of 44 cells. q examined (Fig. 1D). Seven other arrays — showed the more typical nine microchromo- | somal bivalents, while the remaining 30 | spreads were incomplete. | i, | Of the 112 lizards we karyotyped, 110 DISCUSSION Karyological work done on S. graciosus in- cludes 17 specimens examined by Cole (1971, * 1975a) from Arizona, California, Colorado, * New Mexico, and Utah, five specimens exam- ‘ ined by Jackson and Hunsaker (1970) from q California, and 112 specimens examined by us i April 1986 THOMPSON, SITES: LIZARD KARYOTYPES Fig. 1. Karyotypes of Sceloporus graciosus. Bar represents 10 ym for all karyotypes. A, Normal 2N=30 (12M + 18 m) karyotype with terminal satellites (see arrow) on pair two (juvenile, BYU 37620). B, Triploid 3N=45 (adult female, BYU 38201). C, Normal diakinesis array showing nine microchromosomal bivalents, (adult male, BYU 37975). D _ Diakinesis array showing extra “bivalent” (adult male, BYU 37608). from the above states as well as Idaho, Ne- vada, and Wyoming. Of 134 individuals kary- otyped thus far, only two individuals with the anomalies illustrated above (Fig. 1B, 1D) de- viate from the normal 2N=30, 12M + 18m pattern. Since both the anomalous individuals were single isolated cases within their respec- tive population samples (the triploid was one of seven individuals karyotyped from San Bernardino County, California, and the indi- vidual with an extra chromosomal bivalent “)was one of seven specimens karyotyped from ‘Zion National Park, Utah), it is likely that y these cytotypes are not widespread. _ Most reported cases of triploidy in lizards are associated with parthenogenetic popula- tions or species that are thought to have origi- nated by interspecific hybridization (Bickham 1984, Cole 1975b, 1984, Hall 1970). How- ever, isolated cases of triploidy have been found in nonparthenogenetic lizard species. A single aberrant triploid individual was re- ported by Witten (1978) in the Australian agamid eee nee nobbi. Additional cases of triploidy have been reported for two species of Sceloporus. From a sample of 1,300 lizards karyotyped, Hall (1973) scored four triploid individuals of the chromosomally variable S. grammicus complex. Three of these were rales of three different chromo- some “races” or cytotypes. The fourth, a fe- male discovered in a hybrid zone between two 226 cytotypes, possessed two chromosome sets from one parental cytotype and one from the other. The hybrid nature of this karyotype led ‘Hall to suggest that this female may represent an incipient parthenogen. The second exam- ple in the genus Sceloporus was a triploid adult female of S. occidentalis reported by Cole (1983) from a sample of 16 individuals karyotyped from the Pine Valley Mountains of southwestern Utah. At this locality S. occi- dentalis occurs sympatrically with three other species of Sceloporus, but there was no evi- dence that interspecific hybridization was in- volved in the production of this triploid. Fur- thermore, comparative study of the repro- ductive tracts of other adult S. graciosus col- lected the same day indicated that the triploid was sterile, having small unplaited oviducts compared to the broadly plaited oviducts and yolked follicles of the other females. The triploid example we report herein ap- pears also to be an aberrant individual of non- hybrid origin, as all three haploid sets are morphologically typical of S. graciosus (Fig. 1B). This individual appeared to be a pheno- typically normal adult female. The range of the four scale counts plus body length taken from all females collected from the San Bernardino County locality (N=5) are as fol- lows (triploid value in parentheses): (1) scales around midbody, 50-61 (50); (2) dorsal scale count, 48-53 (50); (3) total fourth toe lamellae, 46-50, (50); (4) total supraoculars, 13-15 (14), and (5) (snout vent length, 56-60 mm (56). A noticeable difference was observed, however, in the reproductive condition of these five females. Whereas the four diploids possessed visible paired oviducts and either a shelled egg (in the case of one female) or small clusters of yolked eggs, the triploid uniquely pos- sessed massive fat reserves and lacked visible paired ovaries and oviducts. We interpret this as sterility in the triploid female. We suggest that these three isolated in- stances of aberrant triploid individuals (in Sceloporus) be interpreted simply as repro- ductive accidents (the fusion of a reduced with an unreduced gamete), not as incipient parthenogenesis, unless there is clear evi- dence of reproductive viability and the thely- tokous production of all-female offspring. The male who possessed an extra bivalent in some of its diakinesis arrays is more difficult GREAT BASIN NATURALIST Vol. 46, No. 2 to interpret, as there are no metaphase | spreads or secondary spermatocytes available | for this individual. The simplest explanation is that this extra element represents a single | supernumerary or B-chromosome. It might — also represent an extra microchromosomal | bivalent, although the resolution on our slides — does not permit this distinction to be made. — The fact that not all cells possessed this extra element suggests that, if it is a B-chromosome | or extra bivalent, it is not being distributed to all cells after mitotic division. We stress the | very tentative nature of our interpretation and | recognize the need for additional meiotic and © C-band studies (see Patton, 1977, for a recent | example with rodents). | ACKNOWLEDGMENTS We thank Dr. Robert Bezy, Rusty Bishop, and Calvin Porter for their field assistance, | Dr. Duane Jeffery for reviewing the manu-_ script, Mark Rosenfeld for advice on kary- otypic matters, and Sonya Tracy for darkroom assistance. Funding for this study was pro- vided by the Associated Students of Brigham Young University, the Department of Zool- | ogy, and the Bean Endowment Fund of the M. L. Bean Life Science Museum. LITERATURE CITED BAKER, R. J., M. W. Harbuk, L. W. RopBins, A. CADENA, AND B F. Koop. 1982. Chromosomal studies of the South American bats and their implications. Spec. Pub. iy matuning Lab. Ecol. 6: 303-327. BICkHAM, J. W. 1984. Patterns and modes of chromosoma evolution in reptiles. Pages 13-40 in A. K. Sharma and A. Sharma, eds., Chromosome evolution in eu |— karyotic groups. CRC Press, Inc., Boca Raton, Flor. ida. CoLe, C. J. 1971. Karyotypes of the five monotypic specie: groups of the lizards in the genus Sceloporus. Amer Mus. Nov. 2450: 1-17. . 1975a. Karyotype and systematic status of the sanc dune lizard (Sceloporus graciosus arenicolus) of the American southwest. Herpetologica 31: 288-293. . 1975b. Evolution of parthenogenic species of reptiles ' Pages 340-355 in R. Reinboth, ed., Intersexuality i the animal kingdom. Springer-Verlag, New York. __ | . 1983. Specific status of the North American fenc’ lizards, Sceloporus undulatus and Sceloporus occi dentalis, with comments on chromosome variation Amer. Mus. Nov. 2768: 1-13. . 1984. Unisexual lizards. Sci. Amer. 250: 94-100. Cote, C. J., AND C. R. LEAVENS. 1971. Chromosome prepara, tions of amphibians and reptiles: an improved tech nique. Herp. Rev. 3(6) T-102. April 1986 THOMPSON, SITES: LIZARD KARYOTYPES 297 HALL, W. P. 1970. Three possible cases of parthenogenesis in lizards (Agamidae, Chamaeleontidae, Gekkonidae). Experientia 26: 1271-1272. _____. 1973. Comparative population cytogenetics, specia- tion, and evolution of the iguanid lizard genus Sceloporus. Unpublished dissertation, Harvard Uni- versity, Cambridge. ogy of sceloporine lizards (Sceloporus occidentalis and S. graciosus ). Experientia 26: 198-199. Jackson, L., AND D. HuNSAKER. 1970. Chromosome morphol- PaTToN, J. L. 1967. Chromosome studies of certain pocket mice, genus Perognathus (Rodentia: Heteromyi- dae). J. Mammal. 48: 27-37. ——_—. 1977. B-chromosome systems in the pocket mouse, Perognathus baileyi: meiosis and C-band studies. Chromosoma 60: 1-14. WITTEN. G. J. 1978. A triploid male individual Amphi- bolurus nobbi nobbi (Witten (Lacertilia: Agami- dae)). Aust. J. Zool. 19: 305-308. FOOD HABITS OF CLOUDED SALAMANDERS (ANEIDES FERREUS) IN CURRY COUNTY, OREGON (AMPHIBIA: CAUDATA: PLETHODONTIDAE) John O. Whitaker, Jr. "Chris Maser”, Robert M. Storm’, and Joseph J. Beatty® ABSTRACT. —Stomach contents of 650 clouded salamanders (Aneides ferreus ), collected monthly throughout the year from Curry County, Oregon, were examined. Samples from three age classes were involved: (1) 489 adults, (2) 131 juveniles, and (3) 30 hatchlings. Foods did not vary by sex, but did vary by age and by season. Hatchlings ate small items, particularly mites, springtails, flies, and small beetles. Juveniles fed mainly on flies, isopods (sowbugs), beetles, mites, and centipedes in winter; beetles, ants, and isopods in spring; ants and beetles in summer; and isopods, beetles, and ants in fall. Adults ate isopods and beetles as their major foods in winter, spring, and fall and isopods, ants, beetles, and earwigs in summer. Four species were exceedingly important as foods for these salamanders: an unidentified isopod, the snout beetle (Trachyphloeus bifoveatus), the European earwig (Forficula auricularia), and an ant (Lasius alienus). The clouded salamander (Aneides ferreus) is found from coastal northwestern California northward throughout western Oregon west of the Cascade Range to the Columbia River. Disjunct populations occur on Vancouver Is- land, British Columbia. In Oregon the clouded salamander is found from sea level to elevations of about 1525 m (Beatty 1979). Aneides ferreus is associated with two basic types of habitat throughout its range—talus and fallen trees (Beatty 1979). Large fallen trees are the product of old-growth forests (Franklin et al. 1981, Harris et al. 1982). In Oregon the fallen tree habitat used by A. fer- reus is essentially Douglas-fir (Pseudotsuga menziesii) in varying stages of decomposition (Beatty 1979, Maser and Trappe 1984, Phillips et al. 1981). There have been few food-habit studies of A. ferreus. Fitch (1936) examined the stom- ach of five specimens from the Rogue River Valley in southwestern Oregon. Storm and Aller (1947) examined 63 stomachs from A. ferreus found in decaying Douglas-fir logs in western Oregon. Bury and Martin (1973) compared the distribution and foods of four species of plethodontid salamanders, includ- ing A. ferreus, from the redwood region of northern California; they examined 29 stom- achs of this species. ‘Department of Life Sciences, Indiana State University, Terre Haute, Indiana 47809. 2U.S. Department of the Interior, Bureau of Land Management, Forestry Sciences Laboratory, 3200 Jefferson Way, Corvallis, Oregon 97331. 3Department of Zoology, Oregon State University, Corvallis, Oregon 97331. 298 th, Our study was undertaken to gain a more comprehensive idea of the food habits of A. ferreus throughout the year, by size groups, and by sex. All salamanders had been col-| lected and preserved in connection with a. previous study (Beatty 1979). i { STUDY AREA | | | | The study area, located 9.5 km south of the i Pistol River, Curry County, Oregon, is in the Sitka spruce (Picea sitchensis ) zone within the Klamath Mountains Province (Franklin anc. Dyrness 1973). The study area was burned ir ih 1937 and 1938 and was salvage logged in the. late 1950s and early 1960s. It has since beer, ; grazed continuously by domestic sheep ( Ovi aries) (Fig. 1). The area is open (low grass-forb vegetation with clumps of swordfern (Polystichum muni. ' tum) and trailing blackberry (Rubus ursinus) | The widely scattered trees include Douglas-fir, Sitka spruce, Pacific madrone (Arbutus men. ziesii), California laurel (Umbellularia califor nica), and clumps of red alder (Alnus rubra), Several large, charred Douglas-fir snags wer« still standing when the salamanders were col! | lected (from 1974 through 1977), and the lanc was strewn with many large, fallen, decompos) ing, Douglas-fir trees. These trees composed thi habitat of A. ferreus (Fig. 1). i \ April 1986 WHITAKER ET AL.: SALAMANDER FOOD HABITS Fig. 1. Study area, 9.5 km south of the Pistol River, Curry County, Oregon. The large, woody materials formed the ‘habitat of the clouded salamander. MATERIALS AND METHODS The salamanders were collected by dissect- ing fallen trees and were placed in refriger- iated containers to keep them cool and moist. ‘They were transported to the laboratory within 24 hours (except the May sample), \dge of the cloaca. The stomachs were opened later and the »reserved food items were removed. Sala- -nanders from every month of the year, except May, had food in their stomachs; the lack of ood in specimens taken in May was probably ‘he result of not preserving them soon enough ‘fter capture. Food items were examined in water with the aid of a 10-70 power zoom Hiissecting microscope. The food was identi- ‘ied as completely as possible, counted, and vercent volumes visually calculated for each ype of food in each stomach. Data were then ‘ummarized as mean percent volume, fre- quency of occurrence in stomachs, and per- cent frequency that each item composed of the total number of items. Data were sepa- rated by month, year, age class, and sex in the adult salamanders. RESULTS AND DISCUSSION Results are given in Tables 1-6. To deter- mine if there were any qualitative differences in feeding behaviors because of age, we sepa- rated the data into the following age classes: (1) adults (SVL > 45 mm), (2) juveniles (SVL 22-44 mm), and (3) hatchlings (SVL 14-19 mm). Adults (SVL 45+ mm) It appeared that foods of adult A. ferreus did not vary greatly by sex. To assess similari- ties and differences we used four of the larger individual samples: (1) 32 males, 54 females; 24 March 1976; (2) 15 males, 16 females; 11 April 1975; (3) 18 males, 13 females; 13 August 1976; (4) 15 males, 14 females; 30 November 1974. 230 TABLE 1. Food eaten in winter (December—February) by 55 adult Aneides ferreus (snout-vent length 45+ mm) from Curry County, Oregon. GREAT BASIN NATURALIST Vol. 46, No. 2 | Total percent Percent Percent of volume for Food item volume individuals major group IsorpoDa 44.0 39.4 (44.0) COLEOPTERA (22.9) Curculionidae (18.9) Trachyphloeus bifoveatus 16.1 17.1 Brachyrhinus rugosostriatus 1.8 0.6 Chaetechus setiger 0.7 1.8 Unidentified 0.3 0.6 Carabidae Amara sp. 0.7 0.6 Scarabaeidae Aphrodius cribratulus 2.6 1.8 Unidentified adult 0.7 0.6 DERMAPTERA (5.9) Forficula auricularia 10.8 5.9 DIPTERA (5.8) ' Sciaridae 2.9 §.2 | Mycetophilidae 9 1.8 Phoridae 0.1 0.6 : Unidentified larvae 0.9 ke ARANEIDA (4.2) 94 Gnaphosidae 21 es} | Autrodiaetus pacificus 1.0 0.6 | Ebo sp. probably pepinensis 0.2 12 Amaurobiidae 0.9 1.2 | Micryphantidae 0.04 . 0.6 | CHILOPODA LS 0.6 (1.8) 3% DieLoOPpoDA 1.8 0.6 (1.8) i HOMOPTERA (0.7) | Aphididae 0.3 1.2 Cercopidae 0.2 0.6 | Cicadellidae 0.2 0.6 HYMENOPTERA (0.7) Formicidae ! Aphaenogaster subterraneus 0.6 1.8 i Lasius alienus 0.1 0.6 Unidentified 0.04 0.6 NEUROPTERA (0.7) Hemerobiidae 0.7 0.6 HEMIPTERA (0.3) Tingidae 0.3 0.6 ACARINA 0.1 1.8 (0.1) COLLEMBOLA 0.04 3.5 (0.04) Vegetation 3.3 —_ | Unidentified insect St 2.4 | Shed skin ll i 100.0 99.9 J Percentages varied, perhaps because of The number of sowbugs eaten by female! | small sample sizes. The preferred foods were salamanders was greater than the number’ identical in each case—sowbugs (isopods) eaten by males in three of the four samples; for ranked highest in three of the samples, but example, females ate 23.8% more sowbugs in ants ranked highest in the 13 August sample. November than did the males. Spiders The next most abundant foods were snout formed 16.1% by volume of the male diet in) beetles (curculionids) in the first two samples, | November but were absent from the female ~ isopods in the third sample, and beetles diet. In the August sample, spiders composed (coleopterans) in the fourth sample. 11.5% by volume of the female salamandez) , April 1986 WHITAKER ET AL.: SALAMANDER FOOD Hairs 93] TABLE 2. Foods eaten in spring (March—May) by 212 adult Aneides ferreus (snout-vent length 45+ mm) from Curry County, Oregon. Total percent Percent Percent of volume for Food item volume individuals major group ISOPODA 61.4 41.7 (61.4) | COLEOPTERA (19.8) Curculionidae (14.6) Trachyphloeus bifoveatus iL 7 10.1 Chaetechus setiger 2.1 2.0 Rhyncolus sp. 0.6 0.4 Brachyrhinus ovatus 0.2 0.1 Carabidae (3.0) Calathus ruficollis Ne) 0.7 Harpalus sp. 0.5 0.2 Amara sp. 0.3 0.1 Brachyrhinus rugosostriatus 0.3 0.2 Unidentified 0.4 0.4 Coccinellidae Soymnus ardelio 0.5 0.4 Elateridae 0.2 0.1 _ Byrrhidae Lioon simplicipes 0.1 0.1 _ Tenebrionidae Tenebrio sp. 0.1 0.1 Cucujidae 0.02 0.1 Staphylinidae Tachyporus chrysomelinus 0.01 0.1 Unidentified Coleoptera 1.0 Unidentified larvae 0.3 0.1 HYMENOPTERA (5.2) Formicidae (5.0) Lasius alienus 1.7 GIS Lasius pallitarsus 1.0 1) Aphaenogaster subterraneus 0.7 BW Tapinoma sessile 0.02 0.1 Stenamma diecki 0.01 0.1 Leptothorax sp. 0.01 0.1 Unidentified 1.4 2.3 _ Cynipoidea 0.1 0.2 __ Vespidae 0.1 0.1 DEMAPTERA (5.0) Forficula auricularia 5.0 5.5 _ ARANEIDA (3.7) ) Linyphiidae 0.4 0.6 Antrodiaetus pacificus 0.4 0.1 __ Lycosidae 0.4 0.2 ' Gnaphosidae 0.2 0.1 _ Micryphantidae 0.1 0.1 iF Mimetus hesperus 0.01 0.1 _ Unidentified 9) IES I DIPTERA (2.5) _ Sciaridae 0.9 4.9 A _ Mycetophilidae 0.4 0.9 _ Chironomidae 0.1 0.7 L Chironomid larvae 0.02 0.1 | Phoridae 0.02 0.1 N | Dipterous larvae 0.1 0.2 | Unidentified 1.0 0.9 \ACARINA 0.11 4.7 (0. 11) “| SHILOPODA 0.6 0.6 (0.6) il) LEPIDOPTERA (0.3) (i Larvae 0.2 0.1 iia, Adults 0.1 0.1 232 GREAT BASIN NATURALIST Vol. 46, No. 2 Table 2 continued. Total percent Percent Percent of volume for Food item volume individuals major group HOMOPTERA (0.2) Aphididae 0.2 0.2 Cercopidae 0.04 0.1 ORTHOPTERA (0.2) Gryllidae 0.2 0.1 DIPLOPODA 0.1 0.2 (0.1) HEMIPTERA (0.1) Tingidae 0.1 0.5 COLLEMBOLA 0.06 3.3 (0.06) Unidentified insects 0.4 0.7 Vegetation 0.1 — 99.7 99.0 TABLE 3. Foods eaten in summer (June—August) by 94 adult Aneides ferreus (snout-vent length 45+ mm) from Curry County, Oregon. Total percent Percent Percent of volume for Food item volume individuals major group ISOPODA 25.3 8.9 (25.3) HYMENOPTERA (23.8) Formicidae ; (23.6) Lasius alienus 15.2 50.5 Tapinoma sessile 4.7 11.3 Lasius pallitarsus 2 9.92 Aphaenogaster subterraneus eS 0.5 Leptothorax nitens 0.1 0.2 Leptothorax crassipilis 0.1 0.2 Leptothorax andrei 0.1 0.2 Unidentified Hymenoptera 0.2 0.2 COLEOPTERA (22.4) Curculionidae (13.0) Trachyphloeus bifoveatus 6.4 2.4 Sitona hispidula 3.1 0.7 Brachyrhinus rugosostriatus 1.9 0.7 Rhyncolus sp. 1.2 0.3 Chaetechus setiger 0.4 0.7 Carabidae Calathus ruficollis 2.8 1.4 Tenebrionidae (2.0) Helops sp. 1.1 0.2 Phthora americanum 0.9 0.7 Elateridae Ctenicera sp. 1.1 0.2 Ostomidae Ostoma pippingskoldii 1.1 0.2 Throscidae Pactopus hornii 0.6 0.2 Staphylinidae (0.5) Quedius sp. 0.2 0.2 Stenus sp. 0.1 0.2 Byrrhidfae Lioon simplicipes 0.4 0.2 Anobiidae Coelostethus quadrulus 0.3 0.2 ate April 1986 Table 3 continued. WHITAKER ET AL.: SALAMANDER FOOD HABITS 233 Se ee eS ee Total percent Percent Percent of volume for Food item volume individuals major group Aleocharinae 0.1 0.2 Atheta sp. 0.04 0.2 Unidentified 0.02 0.2 Cucujidae Pediacus depressus 0.1 0.2 Scydmaenidae Lophioderus similis 0.1 0.2 Unidentified 0.4 0.3 DERMAPTERA (11.9) Forficula auricularia 11.9 4.1 HOMOPTERA (3.0) Cercopidae 1.6 0.7 Aphididae 0.02 0.2 Unidentified 1.4 0.3 LEPIDOPTERA (2.9) Larvae 2.3 1.0 Adults 0.6 0.2 ARANEIDA (1.9) Microphantidae Hed 0.2 Linyphiidae 0.6 0.3 Ebo probably pepinensis 0.2 0.2 CHILOPODA 1.8 0.5 (1.8) | HEMIPTERA (1.6) Lygaeidae 0.6 0.5 Unidentified 1.0 0.3 | NEUROPTERA (1.6) Hemerobiidae 1.6 0.7 DIPTERA (0.7) Sciaridae 0.02 0.2 Unidentified larvae 0.6 0.5 Unidentified 0.05 0.2 PHALANGIDA 0.7 0.5 (0.7) ORTHOPTERA (0.6) Gryllidae Gryllus sp. 0.6 0.2 ISOPTERA (0.4) Holotermitidae | Zootermopsis angusticollis 0.4 0.3 NEMERTINEA 0.2 0.2 (0.2) | COLLEMBOLA 0.1 0.2 (0.1) | Vegetation 1.5 = 100.4 100.5 diet but were absent from the stomachs of males. Thus, little or no preference for spiders was shown by either sex. When the same four samples were combined, females were found to eat more sowbugs, cur- culionid beetles, and spiders than did males, but imales did not eat appreciably more of any of the ofoods than did females. These data suggest that omales use a greater diversity of foods. If this is true, then the total volume of these main foods should be greater in females than in males, but such a major difference (67.5% in males versus ay 81.5% in females) was only found in the August sample. Neither does such a trend show up when the number of food categories listed for each sex is examined. For example, 11 categories of foods were eaten by males and females in April and 15 by each sex in November. In August 22 cate- gories were eaten by males and 12 by females; in March 24 categories were eaten by females and 14 by males. We combined the data for the sexes for seasonal comparisons because we detected no major differences in kinds of foods eaten by males versus females. 934 GREAT BASIN NATURALIST Vol. 46, No. 2 TaBLE 4. Foods eaten in fall (September—November) by 125 adult Aneides ferreus (snout-vent length 45+ mm) from Curry County, Oregon. Total percent Percent Percent of volume for Food item volume individuals major group ISOPODA 46.5 38.3 (46.5) COLEOPTERA (24.1) Curculionidae (16.0) Trachyphloeus bifoveatus 13.8 12.0 Chaetechus setiger 1.6 3.8 Sitona hispidulus 0.2 0.1 Unidentified 0.4 0.6 Carabidae (5.1) Calathus ruficollis 4.5 1.8 Unidentified 0.6 0.6 Tenebrionidae (0.9) Phthora americanum 0.5 1.0 Unidentified 0.4 0.1 Scarabaeidae (0.7) Aphrodius sp. 0.7 0.6 Staphylinidae (0.3) Atheta sp. 0.3 0.3 Pselaphidae (0.04) Batrisodes sp. 0.04 0.1 Unidentified 1.0 0.9 Unidentified larvae 0.1 0.1 HYMENOPTERA (6.1) Formicidae (5.5) Lasius alienus 2.1 9.0 Aphaeonogaster subterranea 1.6 1.8 Ant species #9 0.6 3.1 Tapinoma sessile 0.5 1.6 Camponotus vicinus 0.2 0.1 Stenamma diecki 0.2 0.6 Stenamma sp. 0.1 0.4 Ant species #10 0.02 0.1 Unidentified 0.2 0.7 Vespidae 0.3 0.1 Braconidae 0.1 0.1 Unidentified 0.2 0.3 DERMAPTERA (4.6) Forficula auricularia 4.6 12 ARANEIDA (4.3) Clubionidae Trachelus californicus 0.4 0.3 Gnaphosidae 0.4 0.1 Linyphiidae 0.3 1.2 Lycosidae 0.2 0.3 Ebo probably pepinensis 0.1 0.1 Micryphantidae 0.1 0.3 Unidentified 2.8 1.0 HEMIPTERA (3.7) Tingidae 1.4 1.5 Lygaeidae 1.2 0.6 Reduviidae 0.6 0.1 Nabidae 0.5 0.3 Unidentified 0.04 0.1 — LEPIDOPTERA (1.6) Adult 12, 0.6 Larvae 0.4 0.6 HOMOPTERA (1.5) Cercopidae 1.1 1.0 Aphididae 0.3 0.4 Cicadellidae 0.1 0.1 | | April 1986 Table 4 continued. WHITAKER ET AL.: SALAMANDER FOOD HABITS 235 Total percent Percent Percent of volume for Food item volume individuals major group DIPTERA (1.4) Tipulidae 0.9 0.9 Sciaridae 0.04 0.1 Mycetophilidae 0.04 0.1 Dipterous larvae 0.1 0.3 Unidentified 0.3 0.3 ISOPTERA (0.8) Holotermitidae Zootermopsis augusticollis 0.8 Ou) ORTHOPTERA (0.4) Gryllidae Gryllus sp. 0.4 0.1 PHALANGIDA 0.4 0.6 (0.4 ACARINA 0.3 2.4 (0.3) NEUROPTERA (0.3) Hemerobiidae 0.2 0.3 Larvae 0.1 0.1 COLLEMBOLA 0.2 1.3 (0.2) DIPLOPODA 0.1 0.1 (0.1) Vegetation and debris 2.0 — (2.0) Unidentified insect 1.6 0.9 (1.6) Unidentified insect larvae 0.04 0.1 (0.04) 100.0 99.1 WINTER.—We examined the stomachs of 55 adult salamanders taken during winter— December through February (Table 1). The major foods (by volume) were: (1) sowbugs, 44%; (2) beetles, 22.9% (primarily Curculion- idae, 18.9%); (3) earwigs, 5.9%; (4) flies, 5.8%; and (5) spiders, 4.2%. Ants composed 0.7% of the total volume in the sample. Three species were disproportionately im- portant: (1) sowbugs 44.0%, (2) snout beetles | (Trachyphloeus bifoveolatus) 16.1%, and (3) ‘European earwigs (Forficula auricularia) 5.9%. The three species totaled 66.0% by vol- ume of the total prey eaten. The same items occurred in 70.9%, 32.7%, and 16.4%, re- spectively, of the stomachs in the sample. They also composed 39.4%, 17.1%, and 5.9%, or a total of 62.4% of the total items eaten by ‘the 55 salamanders. Although 33 kinds of food »were eaten, the winter diet of A. ferreus in ‘this locality was rather simple. _ SpRING.—Stomachs of 212 salamanders jwere examined for spring diet—March ithrough May (Table 2). Sowbugs were the ‘major food, 61.4% by volume. Beetles, pri- marily curculionids, were second, 14.6% by volume. These items were followed by ants : | | ; and earwigs, each composing 5.0%, and spi- ders, which accounted for 3.7% of the vol- ume. The three species (sowbug, snout beetle, and European earwig) that formed 66% of the winter diet, by volume, formed 78.1% of the spring diet—61.4%, 11.7%, and 5.0%, re- spectively, by volume. Sowbugs occurred in 80.7%, snout beetles in 28.3%, and earwigs in 9.9% of the stomachs, whereas the three com- posed 41.7%, 10.1%, and 5.5%, respectively, for a total of 57.3% of the items eaten. Foods eaten in spring were relatively simi- lar to those eaten in winter. The major differ- ence was that flies decreased in use and ants increased. SUMMER.—We studied the contents of 94 stomachs from salamanders collected from June through August (Table 3). Major differ- ences in food habits among winter, spring, and summer were that in summer sowbugs and the snout beetles (Trachyphloeus bi- foveatus) were much less important, even though beetles, as a whole, were about the same. Ants became an important food in sum- mer and formed almost 25% of the diet. One ant (Lasius alienus), in fact, made up 55.5% of 236 GREAT BASIN NATURALIST Vol. 46, No. 2 TABLE 5. Foods of 131 juvenile Aneides ferreus (snout-vent length 20-45 mm from Curry County, Oregon. Presented as percent volume and percent of prey individuals. Winter Spring Summer Fall Volume Individ- Volume Individ- Volume _ Individ- Volume Individ- Food item % uals % % uals % % uals % % uals % ISOPODA 15.2 9.7 14.5 4.3 6.7 1.8 39.0 13.3 ACARINA 11.4 25.0 8.7 27.8 3.5 22.4 1.3 8.0 CHILOPODA 11.3 5.6 COLLEMBOLA 3.0 23.6 4.4 2252) 3.8 44.4 COLEOPTERA Curculionidae Trachyphloeus bifoveatus 8.0 2.8 5.8 1.3 4.8 1.8 13.6 6.6 Chaetechus setiger 20,1 7.4 Sitona hispidulus 0.4 1.4 Rhyncolus sp. 0.5 0.4 1.3 0.6 Carabidae Calathus ruficollis 1.0 1.4 11 0.6 1.9 0.3 Unidentified 2.4 0.4 Coccinellidae Scymnus ardelio 4.0 1.4 Staphylinidae Sitnao sp. 1.1 0.4 Unidentified D8 1.3 0.5 0.3 Tenebrionidae Phthora americana 0.6 0.3 Byrrhidae Listemus formosus Neil 0.4 : Pselaphidae 0.8 0.4 1.0 0.6 Cucujidae 3.3 0.6 Unidentified 3.3 0.6 1.4 1.0 Larvae 2.3 0.6 HYMENOPTERA Formicidae Lasius alienus 1.3 ES 24.3 38.2 9.2 5.6 Lasius pallitarsus 3.6 IES 2.3 12 Aphaenogaster subterranea 0.5 0.4 8.0 1957 Tapinoma sessile 0.3 0.4 9.5 7.3 Stenamma diecki 4.4 2.8 0.5 0.4 Unidentified 1.2 2.8 15.4 8.3 6.3 2.4 ToT 5.6 Unidentified 0.3 0.3 DIPTERA Sciaridae 4.0 4.2 2.5 1 0.1 0.3 Tipulidae 3.6 1.4 Culicidae 0.4 1.4 Phoridae 0.9 0.4 Syrphidae 1.7 0.6 Larvae 7.8 4.2 4.2 4.5 Unidentified 3.6 2.8 0.4 0.9 LEPIDOPTERA Adult 1.8 0.6 12 0.3 Larvae 2.6 0.3 DERMAPTERA Forficula auricularia 1.4 0.9 0.7 0.6 HOMOPTERA Cicadellidae 0.2 0.4 2.0 1:8 1.8 0.3 Cercopidae 0.9 9) 0.1 0.3 Aphididae 0.9 2) 0.1 0.3 Unidentified no results HEMIPTERA Lygaeidae 4.0 1.4 0.7 0.6 Tingidae 3.8 2.8 1.6 0.4 0.8 1.4 Unidentified 0.8 1.4 0.5 0.3 April 1986 Table 5 continued. WHITAKER ET AL.: SALAMANDER FOOD HABITS Winter Spring Summer Fall Volume Individ- Volume Individ- Volume Individ- Volume Individ- Food item % uals % % uals % % uals % % uals % NEUROPTERA Hemerobiidae 3.3 0.6 ARANEIDA Linyphiidae 4.0 1.4 Ebo pepinensis 0.6 0.7 Xysticus sp. 0.6 1.0 Unidentified 3.8 a7 0.3 0.6 1.4 ei PHALANGIDA 18} 0.3 DIPLOPODA 0.5 0.7 ISOPTERA Holotermitidae Zootermopsis augusticollis 5.3 2.6 Unidentified insect 5.6 2.8 1.6 1.3 RTI 0.6 0.5 0.7 Insect larvae 0.5 0.4 3.2 0.7 Vegetation and debris 4.3 — ' Shed skin 0.6 — 100.0 100.3 99.9 99.6 99.8 99.8 100.1 99.5 allitems eaten and 15.0% ofthe volume. Ants, had its lowest relative use in summer. as a whole, composed 70.2% of all organisms in the sample. The three species that had been so impor- tant in winter and spring composed 43.6% of the summer diet: sowbugs, 25.3%; snout beetles, 6.4%; and European earwigs, 11.9%. There was nearly as great a diversity of foods represented in summer (52 categories) from a _ sample of 94 stomachs as in spring (55 cate- gories) with a sample of 212 stomachs. FaLL.—We examined 125 salamander stomachs collected during fall—September through November (Table 4). Data for fall _resembled those for spring. Major fall foods were sowbugs (46.5% by volume), beetles (24.1%, including 16.0% curculionids), ants (5.5%), European earwigs (4.6%), and spiders (4.3%). Lasius alienus was again the most im- portant ant (61 individuals) and formed 2.1% of the volume and 9.0% of the total items in stomachs. The three species that have been the most important foods, the sowbug, snout beetle, and European earwig, formed, respectively, 446.5%, 13.8%, and 4.6% of the volume, or »64.9% total volume; they also composed, re- ‘spectively, 38.3%, 12.0%, and 1.2%, ora total of 51.5% of the organisms in the sample. ANNUAL.—The sowbug was the major food ‘item of A. ferreus throughout the year, but it | Beetles, collectively, and curculionid beetles in particular were clearly second by volume except in summer when ants were eaten in slightly greater amounts. European earwigs and/or ants were generally in third position by total volume. The earwigs were third in win- ter. Earwigs and ants were used about equally by volume in fall. Ants were second by vol- ume in summer, followed by beetles, then earwigs, even though earwigs contributed their greatest volume (11.9%) during this time. In terms of percent of prey individuals eaten, sowbugs, followed by beetles (mainly curculionids), were highest in winter, spring, and fall. The third and fourth most used food items were flies (11.8%) and earwigs (5.9%) in winter, ants (13.7%) and flies (7.9%) in spring, and ants (17.6%) and spiders (3.4%) in fall. Ants came first in summer; 70.0% of the or- ganisms eaten were ants, followed by beetles (9.4%), sowbugs (8.9%), and earwigs (4.1%). Juveniles (SVL 20-45 mm) WINTER. —The major foods of juveniles in winter (Table 5) were, in order of decreasing volume, flies (19.4%), sowbugs (15.2%), beetles (13.4%), mites (11.4%), centipedes (11.3%), hemipterans (8.6%), ants (5.67), and 238 GREAT BASIN NATURALIST Vol. 46, No. 2 TABLE 6. Foods of 30 hatchling Aneides ferreus (snout-vent length 14-19 mm) from Curry County, Oregon. Fall (n = 17) Winter (n = 13) Volume Frequency Individuals = Volume — Frequency Individuals % % % % % % ACARINA 30.3 76.5 28.0 10.4 30.8 15.0 COLLEMBOLA 27.9 70.6 62.3 16.5 38.4 56.3 COLEOPTERA Trachyphloeus bifoveatus 5.8 oul 1E3 Unidentified 14.7 35.3 3.4 Toll Tot 3.8 HYMENOPTERA Formicidae 5.3 11.8 1.0 4.6 Holl 3.8 HOMOPTERA Cicadellidae Coll UoU 1.3 HEMIPTERA Tingidae 3.8 11.8 1.0 DIPTERA Sciaridae 15.0 15.4 10.0 Chironomidae etl Uo 1.3 Culicidae 3.8 15.4 3.8 Mycetophilidae 1.2 Uo! 1.3 Larvae 4.4 17.6 2.9 DIPLOPODA 2.9 5.9 0.5 ARANEIDA 0.9 5.9 0.5 el Toll 1.3 Insect larvae 5.9 5.9 0.5 Unidentified insect 6.5 Toll 1.3 Unidentified material 3.8 17.6 — Vegetation Be). |: hee — 99.9 100.1 100.4 100.5 springtails (5.5%). Thus, no foods were domi- nant. Rather, the important foods of hatch- lings—flies, mites, and springtails—were still important but in lesser amounts, whereas some of the important foods of adults—sow- bugs and snout beetles—were becoming im- portant but much less so than in adults. This sample suggests a transition from foods of hatchlings to those of adults. The transition is apparently based on size of prey, from small to large, as a salamander grows. SPRING.—By spring the juveniles could contend with larger prey items as follows, in decreasing volume: beetles (34.1%)—again primarily snout beetles; ants (21.6%); sow- bugs (14.5%); mites (8.7%): termites (5.3%): and springtails (4.4%). There was consider- able change from the foods eaten in winter. Flies, centipedes, and bugs decreased, and beetles and ants increased. The decrease in flies in the diet in spring, summer, and fall may be explained by availability. Flies are present and may be relatively inactive in fallen trees in winter, but in summer they either may not be present in large numbers or may be more active and caught less often. Ants may also be more abundant in spring, summer, and fall. Beetles are eaten more of- ten, probably because the more mature sala- manders have an easier time capturing them. We do not have a good explanation for the decrease in centipedes and bugs. These or- ganisms only compose a major food source in winter, and even then only four centipedes | and four bugs were involved, but each ac- — counted for a large part of the respective stom- ach contents. SUMMER.—The most important summer food was ants (50.4% by volume, 61.8% of the» total prey). Ants were followed by beetles — (17.2%), flies (4.2%), and mites (3.5%). No > springtails occurred, and sowbugs—a major | food in all other seasons—formed only 1.8% of the volume. FALL.—The major items by volume were sowbugs (39.0%), beetles (18.0%), and ants | (16.9%). Many or most of the salamanders classified as juveniles have progressed past the size | when the main foods were springtails, mites, and flies. This progress is best illustrated in a_ single collection of 38 subadult salamanders — taken with full stomachs during 29-30. November 1974. Subadults separated nicely | April 1986 into two size classes: (1) 17 individuals with a snout-vent length of 14-18 mm, x — 16.9, and (2) 21 individuals with a snout-vent length of 24-36 mm, x = 29.3. Stomachs of hatchlings contained mites (30.3%) and _ springtails (27.9%); these two foods totaled 58.2% of the diet. Stomachs of juveniles contained only 7.0% springtails and 2.5% mites for a total of 9.5% for these two items. Hatchlings (SVL 14-19 mm) Among the smallest salamanders were six nestlings attended by both an adult male and female. The nest was found 13 September 1976. Five of the nestlings had stomachs that contained 100% yellowish material, thought to be yolk. The sixth had a stomach that con- tained three mites (5% by volume), an ant (Tapinoma sessile) (15%), a mycetophilid fly (30%), and a linyphyid spider (50%). Some feeding may therefore occur while the young are still in the nest. With the exception of the above young and one collected in April, hatchlings (<20 mm SVL) were found from November through February. Of these, 17 were taken in fall and 13 in winter (Table 6). As expected, food items were small. Major foods in the fall sample, in order of decreasing volume, were mites, springtails, and very small beetles. These three groups composed 72.9% of the total vol- ume. The mites and springtails occurred in about 70% to 75% of the stomachs, and the three together accounted for 93.7% of the prey items in the sample. No flies were found. Small food items also dominated the winter sample. Springtails formed the most abun- dant prey items (56.3% of the total), but flies—primarily dark-winged fungus gnats, Sciaridae—made up the greatest collective volume (27.7%). In order of importance, by decreasing volume, were flies, springtails, very small beetles, mites, and spiders. CONCLUSIONS A newly hatched clouded salamander eats ‘small food items (mites and springtails). As an ‘individual matures, the size of prey increases \to larger items, such as beetles and earwigs. ‘All the prey listed in this paper can be found in vand around large, fallen, decomposing Dou- ‘glas-fir trees. Large, rotting Douglas-fir trees Tq WHITAKER ET AL.: SALAMANDER FOOD HABITS 239 concentrate the salamanders’ food; for exam- ple, some beetles and flies, eaten by the sala- manders, depend on these trees for part or all of their life cycles (Deyrup 1975, 1976). Fur- ther, large, decomposing Douglas-fir trees re- main moist inside during summer drought (Maser and Trappe 1984) and thereby main- tain quality habitat not only for the clouded salamander but also for its food supply. ACKNOWLEDGMENTS D. Grayson, D. Metter, and J. Trappe criti- cally read and improved this paper. D. Bel- navis, H. Hanlin, W. Hoffman, S$. Howe, and R. Pietruszka helped with the fieldwork. G. Peters, A. Moldenke, and J. Munsee identi- fied some of the food items. B. Crook allowed us to collect the salamanders on his property. Fieldwork was supported by grants to JJB and RMS from the Theodore Roosevelt Memorial Fund, the American Museum of Natural His- tory, and the Society of the Sigma Xi; identifi- cation of food items was supported by a grant to JOW from the USDA Forest Service, Pacific Northwest Research Station (Coopera- tive Agreement PNW-82-232). We are sin- cerely grateful for the help. This paper represents a partial contribution (no. 3) of the project entitled “The Fallen Tree—An Extension of the Live Tree.” The project is cooperative among the U.S. De- partment of the Interior, Bureau of Land Management; U.S. Department of Agricul- ture, Forest Service, Pacific Northwest Re- search Station; Oregon State University, De- partment of Forest Science; U.S. Department of Agriculture, Agricultural Research Service; and Oregon Department of Fish and Wildlife. LITERATURE CITED Beatty, J. J. 1979. Morphological variation in the clouded salamander, Aneides ferreus (Cope) (Amphibia: Caudata: Plethodontidae). Unpublished disserta- tion, Oregon State University, Corvallis. 94 pp. Bury, R. B., AND M. Martin. 1973. Comparative studies on the distribution and foods of plethodontid sala- manders in the redwood region of northern Cali- fornia. J. Herpetol. 7: 331-335. Deyrup, M. A. 1975. The insect community of dead and dying Douglas-fir. 1: The Hymenoptera. Conifer- ous For. Biome, Ecosyst. Anal. Stud. Bull. 6. 104 pp. 240 _____. 1976. The insect community of dead and dying Douglas-fir: Diptera, Coleoptera, and Neu- roptera. Unpublished dissertation, University of Washington, Seattle. 504 pp. Fitcu, H. S. 1936. Amphibians and reptiles of the Rogue River Basin, Oregon. Amer. Midl. Nat. 17(3): 634-652. FRANKLIN, J. F., AND C. T. DyrNEss. 1973. Natural vegeta- tion of Oregon and Washington. USDA For. Serv., Pacific Northwest For. and Range Exp. Stn., Portland, Oregon, Gen. Tech. Rep., PNW- 8. 417 pp. FRANKLIN, J. F., K. CROMACK, JR., W. DENISON, A. MCKEE, C. MASER, J. SEDELL, F. SWANSON, AND G. JUDAY. 1981. Ecological characteristics of old-growth Douglas-fir forests. USDA For. Serv., Pacific Northwest For. and Range Exp. Stn., Portland, Oregon. Gen. Tech. Rep., PNW-118. 48 pp. GREAT BASIN NATURALIST Vol. 46, No. 2 Harris, L. D.,C. MASER, AND A. MCKEE. 1982. Patterns of _ old growth harvest and implications for Cascades wildlife. N. Amer. Wild. and Nat. Resour. Conf. 47: 374-392. MASER, C., AND J. M. TRAPPE. 1984. The seen and unseen world of the fallen tree. USDA For. Serv., Pacific Northwest For. and Range Exp. Stn., Portland, Oregon. Gen. Tech. Rep., PNW-133. 56 p. PHILLIPS, C., H. CHRostowski, A. MCMILLAN, M. WAL- | TER, R. WILSON, AND M. ELtzrotu. 1981. Wildlife — habitats and species management relationships program, Oregon Coast Range. Vol. II: Amphibi- ans and reptiles. USDA For. Serv., Pacific North- west Region, Corvallis, Oregon, Siuslaw Nat. For., 57 pp. Storm, R. M., AND A. R. ALLER. 1947. Food habits of Aneides ferreus. Herpetol. 4(2): 59-60. WINTERING BATS OF THE UPPER SNAKE RIVER PLAIN: OCCURRENCE IN LAVA-TUBE CAVES David L. Genter! ABSTRACT.—Distribution and habitat selection of hibernating bats at the Idaho National Engineering Laboratory (INEL) and adjacent area are reported. Exploration of over 30 lava-tube caves revealed that two species, Myotis leibii and Plecotus townsendii, hibernate in the upper Snake River Plain. Five species, M. lucifugus, M. evotis, Eptesicus fuscus, Lasionycteris noctivagans, and Lasiurus cinereus are considered migratory. Myotis leibii and P. townsendii _ hibernate throughout much of the area, occasionally in mixed-species groups. Myotis leibii uses the dark and protected regions of the cave, usually wedged into tiny pockets and crevices near or at the highest portion of the ceiling. Individuals of P. townsendii may be found at any height or depth in the cave. Temperature appears to be the primary limiting factor in habitat selection. Myotis leibii was found in significantly cooler air temperatures than P. townsendii. Neither species tolerated continuous temperatures below 1.5 C. Relative humidity does not seem to be a significant | factor in the distribution or habitat selection of the two species in lava-tube caves. Field studies in the northern Rocky Moun- ‘tains have provided fairly comprehensive records of bats for discontinuous geographic locations (Fenton et al. 1983, Genter 1985b, Negus and Findley 1959, Swenson and Shanks 1979). However, information regard- ing wintering species and their ecological re- quirements in the region is minimal. An intensive survey of potential bat hiber- »nacula, population size, and species composi- “tion was conducted during the winter of 1984-85 at the Idaho National Engineering Laboratory (INEL). Additionally, distribu- ‘tion and microhabitat preference of each spe- cies were investigated. STUDY AREA AND METHODS The INEL is on the upper Snake River /Plain near the southeastern ends of the Lemhi ‘and Lost River Mountain ranges. The eleva- _ tion of this predominantly level plain averages 1,525 m. Two volcanic buttes near the south- ‘ern boundary rise to 1,993 m. The vegetation is typical of the cold-desert in the Great Basin -egion. Recent basalt flows cover the south- ‘western portion of the site, extending chroughout the adjacent Craters of the Moon ‘National Monument (CROM). The northeast- erm area is a complex mixture of volcanic de- Epsition, glacial debris, and Lake Terreton 515 E. 6th Avenue, Helena, Montana 59620. Playas. All caves studied were lava tubes re- stricted to the basalt flows (Fig. 1). Temperatures were recorded with a Taylor bulb thermometer. Relative humidity was de- termined using a sling psychrometer (Taylor Instrument Co., Rochester, New York). Ob- servations were made from 12 December 1984 to 27 January 1985, between 1000 and 1600 MST. Temperature and relative humid- ity were measured within each cave at the entrance and throughout the cave at approxi- mately 15 m intervals regardless of the pres- ence of bats and at sites where bats were hibernating. Voucher specimens are in the Zoological Museum of the University of Mon- tana and in the vertebrate collection of the Radiological and Environmental Sciences Laboratory at the INEL. RESULTS Hibernating populations of Myotis leibii and Plecotus townsendii were found. Nine of the 31 caves investigated contained a total of 185 P. townsendii and 78 M. leibii (Table 1). Successive counts in each of several caves re- vealed that the number of individuals of each species remained constant within any cave. Plecotus townsendii were observed to move about within caves, but M. leibii apparently did not. Temperature within caves containing \° Department of Zoology, University of Montana, Missoula, Montana 59812. Present address: Montana Natural Heritage Program, State Library Building, 241 242 02 46 8 =o Kilometers 012345 _— _ = _ <=} Miles eum |NEL boundary Paved road ----- Gravel road Craters of the Moon GREAT BASIN NATURALIST Big Lost River Vol. 46, No. 2 To SALMON Antelope Butte ri mi [XS > TERRETON | Se ea a) To REXBURG ; 4 Circular Butte “Cinder Butte To IDAHO FALLS {20 | © 0 East Butte @ Middle Butte | a © To BLACKFOOT Xs Fig. 1. Bat hibernacula of the INEL basalt flow. 1, EBR II Cave 3. 2, EBR II Cave 4. 3, Rattlesnake Caves. 4, | Moonshiner’s Cave. 5, Middle Butte Cave. 6, Catscat Cave. 7, Sixteen Mile Cave. 8, Arco Tunnel. bats ranged from —1.2 to 7.0 C. Temperatures of hibernacula were more limited. Myotis leibii was found in air temperatures from 1.5 to 5.5 C (X = 2.4 + 0.82). Plecotus townsendii inhabited significantly warmer regions of the caves (t = 2.91, p < .05), with temperatures ranging from 2.2to7.0C (X = 4.8 + 0.96; see Table 1). Caves containing only M. leibii were colder overall than those containing only P. townsendii (X = 1.8 + 1.38 and X = 4.7 + 0.51, respectively; t = 2.49, p < .05). Myotis leibii would commonly be found in the warmer regions of cold caves (sub- freezing), whereas P. townsendii was never found in caves with extensive areas of subfreez- ing temperatures. Relative humidity varied widely among | caves (< 43% to 100%). Hibernacula humidity | | ranged from 65% to 80% for M. leibii and from 68% to 100% for P. townsendii but was not | significantly different between species (t =| 1.22: p >.1). | Each species used different types of roosts | | within the caves. Myotis leibii was closely as- |. sociated with textured and rimatious sub-| strates. Eighty percent wedged themselves in crevices facing outward. All but 2 of the re- ) maining 15 bats were in hollow depressions. In contrast, P. townsendii hung in exposed, | open areas of the cave. M. leibii hibernated’ ) with the ventrum against the cave surface. , i | April 1986 GENTER: WINTERING BATS 243 TaBLE 1. Mean temperature and relative humidity with standard deviation for each cave. Number of each species counted with mean temperature for their roost sites. Myotis Plecotus Cave Temperature (C) Relative Humidity (%) leibii townsendii Arco Tunnel 0.9 + 1.12 Of 28 &83 8) (eo) 0 (-) ‘| Catscat Cave 2.0 + 0.89 67.4 + 11.3 4 (2.1 (0) (=) Rattlesnake Cave 2.4 + 0.82 78.4 +-9.8 4 (2.4 0 (—) (East Tunnel) | Rattlesnake Cave 3.9 + 1.51 Helh se IBY 41 (4.1 132 (5.0) | Sixteen Mile Cave 4.2 + 1.13 84.54 4.7 16 (4.0) 8 (4.4) | Middle Butte Cave 4.7 + 1.36 G26 as M17 9 (4.1 15 (5.5) ) EBRII- Cave 4 4.4 + 0.59 80.3+ 6.7 0 (-) 7 (4.7) EBR II - Cave 3 4.8 + 0.24 Sho as 1/50) 0 (—) 11 (5.0) Moonshiner’s Cave 4.7 + 0.49 100.0 + 0.0 0 (-) 12 (4.8) Plecotus townsendii hung from the substrate. |In approximately 94% of the P. townsendii | both ears were curled in a ram’s horn fashion. - The other 9 individuals had either one or both ears erect. All the latter were found in warmer _ (above 5 C) regions of the cave. Four bats with erect ears were fully aroused and occasionally ‘took flight, whereas bats with curled ears never did so. DISCUSSION The length of time that a bat can hibernate has been estimated for several species (Stud- ier and O Farrell 1976, Hill and Smith 1984). In general, larger bats are capable of longer periods of hibernation and may undergo more frequent episodes of activity. Individuals of M. leibii were found in hibernacula with low air temperatures that varied little temporally or spatially. The narrow range of tempera- tures occupied illustrates habitat selection re- _ flecting their need to conserve energy (Mc- ~Nab 1969, O'Farrell et al. 1971, Studier and O'Farrell 1976). Higher ambient tempera- _ tures induce a higher basal metabolic rate, and the ranges of temperatures chosen by bats keep them within species-specific energy _ budgets necessary for prolonged hibernation (Davis 1970, Hill and Smith 1984). The ten- _ dency of M. leibii to use crevices and shel- ' tered areas likely subjected them to less envi- _ ronmental fluctuation. i}? Plecotus townsendii was the most abundant | species found hibernating in the area. This | Fimay reflect a higher population density or lack | of suitable wintering habitat for other species. oo townsendii hangs from ceilings at varying heights and occupies nearly all depths within the caves. I did not observe them close to cave openings as did Twente (1955, 1960). Nearly all individuals of P. townsendii hiber- nated with their ears in ram’s horn fashion, which may serve to decrease heat loss by radi- ation. The apparent absence of other cavernous species of bats hibernating is not readily ex- plained. Summer field studies indicated the few M. lucifugus at the INEL were closely associated with buildings. Nine individuals of M. leibii were recovered from within and around buildings late in September, suggest- ing these structures serve as hibernacula. Eptesicus left the caves used as summer roosts in late September. As is the case over much of its range, little is known on the win- tering habits of M. evotis. Likely both M. lucifugus and E. fuscus winter in southeastern Idaho; these species commonly hibernate near their summer roost areas (Barbour and Davis 1969). Their absence in this study sug- gests that neither species finds lava-tube caves suitable for hibernation. More extensive surveys are needed to gain a better understanding of the habitat selection and ecology of wintering bats in the northern Great Plains. Investigating a greater diversity of cavernous structures over a wider physio- graphic region should prove fruitful. The win- tering ecology of bats is an important aspect of their life history and one that deserves further study. ACKNOWLEDGMENTS I thank L. H. Metzgar and O. D. Markham for reviewing a previous draft of this paper. This research was funded by the Office of 244 Health and Environmental Research, U.S. De- partment of Energy, and is a contribution of the INEL Radioecology and Ecology Program. Technical and logistical support were provided by many individuals at the Radiological and En- vironmental Sciences Laboratory at the INEL. LITERATURE CITED BARBOUR, R. W., AND W. H. Davis. 1969. Bats of America. University of Kentucky Press, Lexington. 286 pp. Davis, W. H. 1970. Hibernation: ecology and physiologi- cal ecology. Pages 265-300 in W. A. Wimsatt, ed., Biology of bats. FENTON, M. B., H. G. MeRRIAM, AND G. L. HOLROYD. 1983. Bats of Kootenay, Glacier and Revelstroke national parks in Canada: identification by echolo- cation calls, distribution and biology. Canadian J. Zool. 61: 2503-2508. GENTER, D. L. Grand Teton National Park. Spec. Publ. Grand Teton Natl. Hist. Assoc., Moose, Wyoming. 12 pp. ____. 1985b. Bats of the Idaho National Engineering Laboratory: distribution, feeding behavior, and their role in radionuclide transfer. Report to U.S. DOE Radiol. Envir. Sci. Lab. HALForb, D. K., AND J. B. MILLARD. 1978. Vertebrate fauna of a radioactive leaching pond complex in southeastern Idaho. Great Basin Nat. 38: 64-70. HENSHAW, R. E. 1970. Thermoregulation in bats. Pages 188-222 in B. H. Slaughter and D. W. Walton, eds., About bats: a chiropteran symposium. Southern Methodist University Press. GREAT BASIN NATURALIST 1985a. Annotated checklist of bats of Vol. 46, No. 2 HI, J. E., AND J. D. Smiru. 1984. Bats: a natural history. University of Texas Press, Austin. 243 pp. MARKHAM, O, D. 1973. National reactor testing station environmentally related publications. Idaho Op- erations Office, Idaho Falls. 11 pp. . L978. Ecological studies on the INEL site. 1978 progress report. U.S. DOE, Idaho Operations OF fice, Idaho Falls. 371 pp. . 1983. INEL radioecology and ecology programs. 1983 progress report. U.S. DOE, Idaho Opera- tions Office, Idaho Falls. 434 pp. McNas, B. kK. 1969. The economics of temperature regu- lation in neotropical bats. Comp. Biochem. Phys- iol. 31; 227-268. Necus, N.C., AND J. S. FINDLEY. 1959. Mammals of Jack- son Hole, Wyoming. J. Mammal. 40: 371-381. O’FARRELL, M. J., E. H. STUDIER, AND W. G. Ewinc. 1971. Energy utilization and water requirements of cap- tive Myotis thysanodes and Myotis lucifugus (Chi- roptera). Comp. Biochem. Physiol. 39: 549-552. STUDIER, E. H., AND M. J. O’FARRELL. 1976. Biology of Myotis thysanodes, M. lucifugus (Chiroptera; Ves- pertilionidae)—II. Metabolism, heart rate, breathing rate, evaporative water loss and general energetics. Comp. Biochem. Physiol. 54: 423-432. SWENSON, J. E., AND G. Y. SHANKS, JR. 1979. Noteworthy records of bats from northeastern Montana. J. Mammal. 60: 650-652. TWENTE, J. W., JR. 1955. Some aspects of habitat selection and other behavior of cavern-dwelling bats. Ecol- ogy 36: 706-732. 1960. Environmental problems involving the hi- bernation of bats in Utah. Utah Acad. Sci. Proe. 37: 67-71. | | iH i i { \ { GROWTH RATES OF MULE DEER FETUSES UNDER DIFFERENT WINTER CONDITIONS Richard M. r Bartmann ABSTRACT. —Based on forehead-rump length, growth rates of mule deer (Odocoileus hemionus) fetuses in Piceance Fetal forehead-rump measurements have | displayed strong correlations with age in mule ‘deer (Odocoileus hemionus) (Hudson and | Browman 1959) and white-tailed deer (O. vir- / ginianus) (Cheatum and Morton 1946, Short 1970). These relationships were developed \ with captive deer maintained on artificial ra- tions and provide no indication of variability ‘that may occur in fetal growth rates under stress in natural environments. Verme (1963) reported slower fetal growth in captive white- ‘tailed does in poor condition during the last ‘third of gestation, anda possible relationship between poor doe condition and slower fetal - growth was reported for white-tailed deer in ‘New York (Jackson and Hesselton 1973). The 1971 and 1972 winters in Piceance Basin, northwestern Colorado, were rela- tively moderate compared to harsh conditions in 1973, when an estimated 40% of the deer population perished (Bartmann and Bowden 1984). The severity of the latter winter al- lowed examining the effects of undue stress on fetal growth rates of mule deer. STUDY AREA Piceance Basin includes about 1,722 km? of pinyon-juniper (Pinus edulis—Juniperus os- ‘teosperma ) winter range for mule deer at ele- vations between 1,675 and 2,285 m (Bart- ‘mann and Steinert 1981). Deer begin migrating to winter range in early October, with return migration to summer range in ‘April and May. Deer movement across sev- eral major roads, particularly during late win- iter and spring, often results in a high inci- dence of road-killed deer. Basin, Colorado, were slower during severe winters than during moderate ones. Growth rates in both situations were _ slower than reported for both mule deer and white-tailed deer (Odocoileus virginianus) fetuses from captive does. METHODS Fetuses were acquired from 83 mule deer does, aged 1'/2 to 10+ years, and included 58 road-kills, 18 trapping mortalities, 6 fence- kills, and 1 predator kill. Deaths occurred 25 March-13 May 1971 (N = 13), 21 January—14 April 1972 (N = 19), and 19 Janwary 216 May 1973 (N = 51). Date of death was known for 54 does and known to within a maximum of two days for the remainder. Fetal data were taken only from does with the abdominal cavity in- tact and all fetuses undamaged. Forehead- rump measurements were made as described by Armstrong (1950). Measurements for twins were averaged to provide a single value. Separate linear regressions of fetal fore- head-rump length (mm) (Y) on collection date (Julian day) (X) were calculated for each year's data. Separate linear regressions were also calculated from data for known-age mule deer (Hudson and Browman 1959) and white-tailed deer (Cheatum and Morton 1946) fetuses 48—174 days old. Fetal ages in this study were estimated to be within this age range, and Short (1970) indicated a linear relationship adequately reflected this period of fetal growth. Regression line slopes for Piceance deer were compared to those for known-age mule deer and white-tailed deer fetuses to test for differences in growth rates. I emphasize that the X variable in this study is collection date, whereas for the known-age fetuses it is fetal age. Collection date for a given deer can be written as age in days minus an unknown con- stant plus an error. If the population of deer fetuses was completely measured on a given ‘Colorado Division of Wildlife Research ¢ Jenter, 317 West Prospect, Fort Collins, Colorado 80526 2 | 45 246 400 (b] 300 200 £ E — 100 Go = S . 12=0.94 . 12=0.89 TT} ¥=14.2 + 2.74x ¥=361+ 2.30X — o (0) 100 200 0 100 200 = COLLECTION DATE (Julian Day) oc fa) i 400 [d] x= WwW oc fe) re 300 200 100 r2=0.998 r2=0.996 ¥=-124.6 + 3.21X Y¥=-122.04+3.21X 100 200 O 200 0) 100 FETAL AGE (Days) Fig. 1. Forehead-rump length as a function of collec- tion date for mule deer fetuses collected in Piceance Basin, Colorado, January—May (a) 1971-1972 and (b) 1973. Forehead-rump length as a function of age for known-age (c) mule deer (Hudson and Browman 1959) and (d) white-tailed deer (Cheatum and Morton 1946) fetuses. collection date, then the corresponding errors are assumed to have a mean of zero. Given the approach of Berkson (1950), the slope of the regression line of fetal length on collection date is then the same as the slope of the re- gression line of fetal length on age. Additional analyses were performed to test for differ- ences in growth rates for single and twin fe- tuses and for size differences in male and fe- male fetuses within the same twin set. GREAT BASIN NATURALIST Vol. 46, No. 2. RESULTS AND DISCUSSION Sample sizes for 1971 and 1972 were small compared to 1973, and fetal collections in 1971 occurred over only a 1'/:-month period. Linear regression lines for these two years were similar _ (P = 0.91) (1971, Y =3.1 + 2.81X; 1972; Y = S70 + 2.87X); but the line for 1972 was different (P = _ 0.02) from that for 1973. Since winter conditions | in 1971 and 1972 were also similar, data from’ these two years were pooled and a new regres-_ sion calculated. ; ( 4 The fetal growth rate in 1973 was slower (P =' 0.02) than in 1971-1972 (Fig. la, 1b), and it was also slower (P < 0.002) than growth rates foi’ known-age mule deer and white-tailed deer fe, tuses (Fig. lc, 1d). The rate in 1971-1972 wa: also slower than the rate for known-age fetuses ! but differences were not quite significant (P =) 0.06). i Growth rates were similar (P = 0. 87) for fe tuses collected January-March (Y = 21.3 4 2.59X, N = 26) and April-May (Y = 10.7 -| 2.50X, N = 25) in 1973. This suggests that i. natural environments doe condition affects fet:' growth sooner than the last third of gestation ¢ reported by Verme (1964). Variability in fetal development due to sex an. litter size is expected, but its effect on age-lengt relationships is unclear. Strong correlations be’ tween age and length of known-age mule dee and white-tailed deer fetuses (Fig. lc, 1d) re! sulted despite not only small sample sizes (4 an | 10, respectively) but also, with white-taile deer, a mix of single and twin fetuses of bot’ sexes. Differences in forehead-rump length of) | sets of mixed gender twins in this study range’ from 0-32 mm. Females averaged 3 mm long¢ | than males, but this difference was not signi! ‘ cant (P = 0.09). In contrast, Jackson and Hesse * ton (1973) found male fetuses were general larger than female fetuses at any given agi Growth rates for single and twin fetuses we compared separately for 1971-1972 and 197. Again, there was no difference (P = 0.31) | either case. A common use of fetal growth rates is to est mate breeding dates. The growth rate for mu‘ deer fetuses derived from data of Hudson ar ‘ Browman (1959) was applied to the 1971-19" and 1973 Piceance data. Linear regressions | estimated breeding date on collection date we then calculated (Fig 2a, 2b). Both regression lit; Ss = = “Ulan slopes were >0 (P < 0.001). Thus, the later |}, April 1986 BARTMANN: MULE DEER 9AT 10 ° = oq . 2=0.32 r2=0.56 3604 Y—322.0+0.15Xx [al e ¥=314.9 + 0.29X Ib) 340 320 r2—0.05 [c] 360 ¥=327.1- 0.06 x ESTIMATED BREEDING DATE (Julian Day] 340 O ee e : e e ee e e 320 i ° 3.. e e ie e 300 : COLLECTION DATE r2=0.10 (d} ¥ =318.4 + 0.11% 0 50 100 150 (Julian Day] Fig. 2. Change in estimated breeding date with change in collection date of mule deer fetuses in Piceance Basin, | Colorado, (a) 1971-1972 and (b) 1973 using the fetal growth rate calculated from data of Hudson and Browman (1959) _ and (c) 1971-1972 and (d) 1973 using the fetal growth rate of Thomas (1970). / gestation fetal data are collected, the later the _ breeding date estimate. Thomas (1970) derived a fetal growth rate for Columbian black-tailed deer (O. h. colum- bianus) (Y = 2.57X — 83.6) based on fetuses from wild does plus known-age fetuses from _ two captives. This growth rate better charac- _ terized fetal growth in 1971-1972 because the _ regression line slope, although negative, was not different from 0 (P = 0.23) (Fig. 2c). How- { ever, slower fetal growth in 1973 still pro- j, Bncedia slope >0 (P = 0.02) (Fig. 2d). _ Results of this study indicated fetal growth _ can vary significantly with winter conditions, and published growth rates do not always fit _ wild populations. Thus, it is important that _ fetuses be obtained over the range of gestation _to enable evaluating applicability of any fetal é ‘growth rate to be used. | ACKNOWLEDGMENTS » Ithank D. C. Bowden, D. J. Freddy, N. T. Ss F. B. Samson, and G. C. White for manuscript review and S. F. Steinert and J. J. Klein, Jr., for field assistance. D. C. Bowden also provided statistical assistance. Research was supported by Colorado Federal Aid to Wildlife Restoration Project W-38-R. LITERATURE CITED ARMSTRONG, R. A. 1950. Fetal development of the northern white-tailed deer (Odocoileus virginianus borealis Miller). Amer. Mid]. Nat. 43: 650-666. BARTMANN, R. M., AND D. C. BowbeEN. 1984. Predicting mule deer mortality from weather data in Colorado. Wildl. Soc. Bull. 12: 246-248. BARTMANN, R. M., AND S. F. STEINERT. 1981. Distribution and movements of mule deer in the White River drainage, Colorado. Colorado Div. Wildl. Spec. Rep. 51. 12 pp. BERKSON, J. 1950. Are there two regressions? J. Amer. Stat. Assoc. 45: 164-180. CHEATUM, E. L., ANDG. H. Morton. 1946. Breeding season of white-tailed deer in New York. J. Wild]. Manage. 10: 249-263. Hupson, P., AND L. G. BrowMaN. 1959. Embryonic and fetal development of the mule deer. J. Wildl. Manage. 23: 295-304. 248 GREAT BASIN NATURALIST Vol. 46, No. 2. Tuomas, D. C. 1970. The ovary, reproduction, and pro- | ductivity of female Columbian black-tailed deer. | Unpublished dissertation, University of British | Jackson, L. W., AND W. T. HESSELTON. 1973. Breeding and parturition dates of white-tailed deer in New York. New York Fish and Game J. 20: 40-47. ; Columbia, Vancouver. 211 pp. SHoRT, C. 1970. Morphological development and aging of — VgeRue L. ]. 1963. Effect of nutrition on growth of white- | mule and white-tailed deer fetuses. J. Wildl. Man- tailed deer fawns. Trans. N. Amer. Wildl. and age. 34: 383-388. Nat. Resour. Conf. 32: 405-420. DENNING HABITAT AND DIET OF THE SWIFT FOX IN WESTERN SOUTH DAKOTA Daniel W. Uresk' and Jon C. Sharps” ABSTRACT.—Swift fox (Vulpes velox) were investigated in western South Dakota to determine food habits and denning site characteristics. Over a three-year period food habits consisted of mammals (49%), followed by insects (27%), plants (13%), and birds (6%). Dens were located near hilltops within two habitat types, shortgrass and midgrass prairie; each type is characterized by differing plant species. Soil type was not a selective factor for den sites of swift fox. Management considerations for enhancing swift fox populations are presented. The swift fox (Vulpes velox), a threatened species in South Dakota, was reported to be abundant on the Great Plains when settlers arrived (Egoscue 1979). With the increasing settlement of the northern High Plains, popu- lations of swift fox declined, and by 1900 the species was rare in its northern range (Beck 1958, Soper 1964, Egoscue 1979). In South Dakota swift fox sightings were not reported between 1914 and 1966 (Hillman and Sharps 1978). The decline of the swift fox population has been attributed to the loss of natural prairie habitat, predator and rodent control programs, excessive trapping, and hunting ((Egoscue 1979). Since 1975 a small population of swift fox has been present in South Dakota, which pro- vided an opportunity for observation of food habits and habitat characteristics around den- ning sites (Hillman and Sharps 1978). Other ‘investigations on food habits and habitat of ‘denning sites have been reported by Kilgore (1969) in Oklahoma and Cutter (1958a, 1958b) ‘in Texas. However, very little information is available on swift fox food habits and den site characteristics within the northern range of its distribution. The objectives of this study were to determine food habits of the swift fox and habitat character- istics at denning sites in western South Dakota. STUDY AREA _ The study areas selected were known to ‘aave viable swift fox populations. The first was located on the Pine Ridge Indian Reservation, Shannon County, South Dakota, approxi- mately 23 km north of Oglala. This area is a broad flood plain with gently sloping to undu- lating upland prairie, bordered by the White River to the north. Badland outcroppings, which are found throughout the Pine Ridge area, are typified by bare soil; soil types are primarily clayey to sandy-clay-loam. Annual precipitation averages 41 cm, with an annual snowfall of 79 cm. Dominant vegetation con- sists of buffalograss (Buchloe dactyloides), needleleaf sedge (Carex eleocharis), blue grama (Bouteloua gracilis), and western wheatgrass (Agropyron smithii). Livestock graze throughout the area. The second study area was located in Haakon County, 40 km north of Philip. This area is char- acterized by gently undulating hills with numer- ous livestock watering ponds. The soil type is primarily clay to clay-loam. Annual precipitation averages 43 cm, with an annual snowfall of 30 cm. Dominant vegetation of this area includes western wheatgrass, buffalograss, and blue grama. Livestock graze the area and, in addition, there is some farming. METHODS Swift fox were located by spotlighting within each of the study areas. On the Pine Ridge study site, three natal dens were stud- ied during 1977 and 1978 and four in 1979. One natal den was studied north of Philip during 1978 and 1979. non ! TUSDA Forest Service, Rocky Mountain Forest and Range Experiment Station, Rapid City, South Dakota 57701. >South Dakota Department of Game, Fish, and Parks, Rapid City, South Dakota 57701. 249 250 TABLE l. Major plant species characterizing den sites of swift fox at two study areas (frequency of occurrence = 5%) in South Dakota. Species GRASS AND GRASSLIKE Western wheatgrass (Agropyron smithii) Little bluestem (Schizachyrium scoparium) Red threeawn (Aristida longiseta) Blue grama (Bouteloua gracilis) Japanese brome (Bromus japonicus ) Cheatgrass brome (Bromus tectorum) Buffalograss (Buchloe dactyloides ) Sixweeks fescue (Vulpia octoflora) Needleandthread (Stipa comata) Needleleaf sedge (Carex eleocharis ) FORBS Onion (Allium spp.) Pinnate tansy mustard (Descurania pinnata ) Curlycup gumweed (Grindelia squarrosa) Prairie sunflower (Helianthus petiolaris ) Stickseed (Lappula redowskii) Prairie pepperweed (Lepidium densiflorum) Virginia pepperweed (Lepidium virginicum) Dotted gayfeather (Liatris punctata) Rush skeletonplant (Lygodesmia juncea) Black medic (Medicago lupilina) Yellow sweetclover (Melilotus officinalis ) Musineon (Musineon divaricatum) Hoods phlox (Phlox hoodii) Scarlet globemallow (Sphaeralcea coccinea) Field pennycress (Thlaspi arvense ) Vetch (Vicia spp.) SHRUBS Silky wormwood (Artemisia dracunculoides) Vegetation analyses were conducted during June and early September of 1978 and 1979. Frequency of occurrence was estimated along two 30m line transects located at each den site, and estimates were made by reading 30 2 broadly ovate to ovate-oblong or elliptic, basally rounded to truncate or subcordate, — irregularly denticulate blade 4—6 x 2.5-5 cm; — herbage glandular-puberulent; flowers axil- lary to small bracts on the lateral branches and — toward end of the main stem, in bud crowded, | erect, and nearly sessile, but becoming more _ widely spaced and evidently pedicellate at anthesis, fairly showy and adapted to cross- pollination; petals purple, 7-14 mm long, somewhat asymmetrically (adaxially) dis- posed; stamens 4 + 4, unequal, declined; an-_ thers ca 1.5-2 mm long; fruits 12-25 mm_ long, spreading-deflexed, on spreading-de- flexed pedicels 3-5 mm long; seeds biseriate | in each locule, plumply obovate from a sub- stipitate base, 1.5-1.8 mm long, very finely cellular-reticulate. This species is known only by several cole lections by Duane Atwood from the type lo-, cality on open slopes on the southwest side of | the Kaiparowits Plateau, ca 27 km by road northeast of Glen Canyon City, Kane Co., Utah. Flowering is from August to October. | Camissonia boothii var. condensata, (Munz) Crong. comb. nov. [based on:) Oenothera decorticans var. condensata Munz | Bot. Gaz. 85: 247. 1928]. | Camissonia boothii var. villosa (Wats.)) Crong. comb. nov. [based on: Oenothera. alyssoides var. villosa Wats., Proc. Amer. Acad. | Arts 8: 591. 1873; not Oe. villosa Thunb. 1794:;, typification will be discussed in the forthcoming treatment in the Intermountain Flora. ]. | Camissonia clavaeformis (Torr. & Frem.) Raven var. purpurascens (Wats.) Crong. comb. nov. [based on: Oenothera scapoidea var. pur- | purascens Wats. Proc. Amer. Acad. 8: 595. if 1973]. 4 Camissonia scapoidea (T. & G) Raven var. utahensis (Raven) Welsh comb. nov. [based on: Oenothera scapoidea ssp. utahensis Raven Univ. California Publ. Bot. 34: 96. 1962]. April 1986 Oenothera _caespitosa Nutt. — var. macroglottis (Rydb.) Crongq. stat. nov. [based on: Pachylophus macroglottis Rydb. Bull. Torrey Bot. Club 30: 259. 1903]. Oenothera caespitosa Nutt. var. navajoen- sis (Wagner, Stockhouse, & Klein) Cronq. stat. nov. [based on: Oenothera caespitosa ssp. navajoensis Wagner, Stockhouse, & Klein Monogr. Syst. Bot. Missouri Bot Gard. In press 1985]. Oenothera flava (A. Nels.) Garrett var. acutissima (W. L. Wagner) Welsh comb. nov. [based on: Oe. acutissima W. L. Wagner Syst. Bot. 6: 153. 1981, type from Flaming Gorge vicinity, Daggett County, Utah]. Oenothera primiveris Gray var. bufonis (Jones) Crong. comb. nov. [based on: Oe. bufonis Jones, Contr. W. Bot. 8: 28. 1898]. ORCHIDACEAE Habenaria zothecina Higgins & Welsh sp. nov. Planta Habenaria sparsiflora affinis et similis sed in calcari labium 1.5 longiore et floribus paucioribus differt. Type: USA: Utah. Grand Co., ca 1.6 km N of Moab, T25S, R21E, $25, at ca 1,281 m, ina hanging garden community, 10 July 1985, S. L. Welsh & L. C. Higgins 23629 (Holotype BRY; 4 isotypes to be distributed. ADDITIONAL COLLECTIONS: Utah. Grand Co., Arches National Monument, 28 Sept. 1963, S. L. Welsh & G. Moore 2735; ibid., Negro Bill Canyon vicinity, 3 August 1984, B. Franklin 1108. San Juan Co., Natural Bridges _ National Monument, 13 August 1963, S. L. Welsh & G. Moore 2410; ibid., 15 August / 1963, S. L. Welsh & G. Moore 2503; ibid., The Neck vicinity, 31 Oct. 1964, S. L. Welsh & G. Moore 3840 (all BRY). This peculiar, few-flowered bog orchid with its very long spurs has been known for many years. It has always been placed previously within an expanded version of H. sparsiflora but differs as outlined in the diagnosis and in other salient features. The leaves are broadly rounded initially, becoming obtuse and finally “acute upward. The orchid grows in the vege- {tative assemblages known as hanging gardens salong the canyons of the Colorado River in southeastern Utah. They occur there with »maidenhair fern, sheathed death camus, and other shade-tolerant mesophytes. They occur qi WELSH: NEW PLANT TAXA AND COMBINATIONS 259 mainly along the margin of the detrital slope adjacent to the back wall of the alcoves. Addi- tional work is necessary to determine the total distribution. PAPAVERACEAE Papaver radicatum Rottb. var. pygmaeum (Rydb.) Welsh stat nov. [based on: Papaver pygmaeum Rydb. Bull. Torrey Bot Club 29: 159. 1902]. PRIMULACEAE Dodecatheon pulchellum (Raf.) Merr. var. zionense (Eastw.) Welsh stat. nov. [based on: Dodecatheon zionense Eastw. Leafl. W. Bot. 2: 37. 1937, type from Zion Canyon]. RANUNCULACEAE Aquilegia flavescens Wats. var. rubicunda (Tidestr.) Welsh stat. nov. [based on: Aquile- gia rubicunda Tidestr. Amer. Midl. Nat. 1: 168. 1910]. A peculiar specimen of Aquilegia taken by Dr. Robert Foster from Zion Canyon in 1977 requires description. It has the habit and flower color of A. formosa but is glandular throughout and both petal spur and blade are longer than in that taxon. A search of the herbarium at Zion National Park yielded two specimens of the same taxon taken many years ago by Dr. Angus Woodbury. Initial efforts to relocate the area of collection of the type have been fruitless. There are many areas to be investigated, however, and the Hidden Canyon locality of the Woodbury specimens has not been searched recently. Possibly the specimens result from introgression between A. formosa and A. chrysantha Gray, both abundant in the canyon. The longer spurs and petal blades suggest such a possibility, but neither of the potential parents are glandular below the inflorescence. The plant is named after Dr. Robert Foster, enthusiastic collector and plant geographer. Aquilegia formosa Fisch. in DC. var. fos- teri Welsh var. nov. A var. formosa in petali laminis et calcari longioribus et herba glandu- losis diversa. Typr: USA: Utah. Washington Co., T41S, R1OW, S21, W of Tunnel, 1,373 m, N slope of 260 Bridge Mt., mountain brush community, on Wingate detritis, 25 May 1977, R. & R. Foster 3939 (Holotype BRY). ADDITIONAL SPECIMENS: Utah. Washington Co., Zion National Park, Hidden Canyon, in 1924, A. Woodbury s.n. (ZNP herbarium, 2 specimens). Delphinium andersonii Gray var. scapo- sum (Greene) Welsh stat. nov. [based on: Del- phinium scaposum Greene Bot. Gaz. 6: 156. 1881]. Delphinium occidentalis (Wats.) Wats. var. barbeyi (Huth) Welsh comb. nov. [based on: Delphinium exaltatum var. barbeyi Huth He- lios 10: 35. 1892]. Ranunculus andersonii Gray var. juniperi- nus (Jones) Welsh stat. nov. [based on: Ra- nunculus juniperinus Jones Proc. Calif. Acad. II. 5: 616. 1895]. Ranunculus acris L. var. aestivalis (L. Ben- son) Welsh comb. nov. [based on: Ranunculus acriformis var. aestivalis L. Benson Amer Midl. Naturalist 40: 43. 1948], type from 8.3 mi N of Panguitch]. GREAT BASIN NATURALIST Vol. 46, No. 2 | ROSACEAE Purshia mexicana (D. Don) Welsh comb. | nov. [based on: Cowania mexicana D. Don Trans. Linnaean Soc. 14: 575. 1825]. Purshia mexicana (D. Don) Welsh var. stansburyi (Torr.) Welsh stat. nov. [based on: | Cowania stansburiana Torr. in Stansbury Expl. Surv. Utah 386. 1852]. RUBIACEAE Galium mexicanum H.B.K. var. asperri- | mum (Gray) Higgins & Welsh stat. nov. | [based on: Galium asperrimum Gray Mem. © Amer. Acad. Arts II. 4: 60. 1849]. SCROPHULARIACEAE Castilleja parvula Rydb. var. revealii (N. Holmgren) N. D. Atwood stat. nov. [based i on: Castilleja revealii N. Holmgren Bull. tom rey Bot. Club 100: 87. 1973]. Castilleja rhexifolia Rydb. var. sulphurell | (Rydb.) N. D. Atwood stat. nov. Based on: | Castilleja sulphurea Rydb. Mem. New York | Bot. Gard. 1: 359. 1900]. NEW TAXA IN MISCELLANEOUS FAMILIES FROM UTAH Stanley L. Welsh! ABSTRACT.—Named are the following: Astragalus limnocharis Barneby var. tabulaeus Welsh var. nov., from the pass between Boulder Mountain and the Table Cliff Plateau, Garfield County, Utah; A. eremiticus Sheldon var. ampularioides Welsh var. nov. from Washington County, Utah; Lupinus argenteus Pursh var. moabensis Welsh var. nov., from southeastern Utah, validated by inclusion of a Latin diagnosis; Erigeron zothecinus Welsh sp. nov. described from moist alcoves along Lake Powell, eastern Kane County, Utah; Cleomella palmerana Jones var. goodrichii Welsh var. nov. described from Uintah County, Utah; Arabis vivariensis Welsh sp. nov. named from northeastern Uintah County, Utah; Draba kassii Welsh sp. nov. described from material taken in the Deep Creek Mountains, western Tooele County, Utah. A curious, small-flowered Astragalus was dis- covered by Sherel Goodrich and me immedi- ately north of the pass between Boulder Moun- tain and Table Cliff Plateau 20 June 1981. The material was first taken to be an extension of the similar A. montii Welsh from much farther north on the Wasatch Plateau, primarily on the basis of the pink-purple flowers. However, the wing tips are not white, and the flowers average smaller than in that taxon. Placement with the geograph- ically nearer and morphologically more similar A. limnocharis Barneby became evident with additional study. The flower size, shape of petals, and pod size and conformation fit well within the range for A. limnocharis. The strongly soboliferous habit and pink-purple flowers are notably different, however. The soboliferous habit is an adaptation that allows occupation of the steep slopes where the plants grow, a habitat not usually occupied by A. lim- nocharis proper. The presence of sobols might represent merely an ecological response to the ‘creeping mantle on the slopes, but it is readily \apparent both in the field and in herbarium spec- imens. The plants from adjacent to the Table Cliff Plateau are named as follows: Astragalus limnocharis Barneby var. tabu- aeus Welsh var. nov. Planta persimilis Astra- galo limnochari Barneby in floribus et fructus sed in caudicibus soboliferis et floribus pur- oureis differt. Type: USA: Utah. Garfield Co., T34S, R1W, BE/SW S22, ca 2,930 m in a Pinus longaeva | community, on the White Limestone Member of the Wasatch Formation, on a60%-—70% south- facing slope, 20 June 1981, S. L. Welsh 20666 (Holotype BRY; 2 isotypes distributed previ- ously as A. montii). ADDITIONAL SPECIMENS: Utah. Garfield Co., same approximate locality and date as the holo- type, S. L. Welsh 20667, 20667a, and S. Goodrich 15662, 15669 (all BRY). Growing on the Chinle Formation west of the Gunlock intersection at Shem, Washington County, Utah, is a second more or less distinc- tive phase of Astragalus eremiticus Sheldon. The plants simulate A. ampullarius in having subterranean caudices and short, broad, long- stipitate pods. The pods are smaller than in A. ampullarius, but the stems recline as in that species. The elongate many-flowered racemes are similar to those of typical A. eremiticus , which occurs elsewhere in the county, but the flowers are more numerous (up to 45), the pe- duncles more elongate (up to 21 cm), the raceme very lax in fruit (up to 25 cm), and the pods are tumid and truncate to obtuse basally and 8-15 (18) mm long and 6-12 mm wide. Pods of some plants from elsewhere in Washington County are tumid but not as abruptly broadened or as broad as in the Shivwits plants. And, when the pods are tumid, the other features are as in typi- cal E. eremiticus. The habitat consists of barren silty clays of the Chinle Formation, a stratum supporting A. ampullarius in all its known locali- ties. Because of the similarities to A. ampullar- ius, this distinctive phase is designated as fol- lows: | 1Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. | 261 262 Astragalus eremiticus Sheldon var. am- pullarioides Welsh var. nov. Planta similis A. eremitico var. eremitico in racemis et floribus sed in leguminibus brevioribus et latioribus et caudicibus subterraneis differt, et similis A. ampullario in leguminibus formis et cau- dicibus subterraneis sed in racemis elongatis et floribus plus numerosis et coloris flavis dif- fert. Type: USA: Utah. Washington Co., T41S, RI7W, N of Highway 91 at Shivwits, 1050 m elev., 21 Apr. 1982, S. L. Welsh and N. D. Atwood 21049 (Holotype BRY; an isotype dis- tributed previously as A. eremiticus Sheld.). ADDITIONAL SPECIMENS: Utah. Washington o., ibid., 4 May 1976, N. D. Atwood 6586 (BRY); ibid., 9 June 1983, L. C. Higgins and N. D. Atwood 13683 (BRY); ibid., May 1985, S. L. Welsh 23456 (BRY). The plants are evidently relished by live- stock, and in most years the inflorescences are eaten. The extent of the population is un- known. The presumption was made by Welsh (1978, p. 326) that a name proposed by Dunn and Harmon for the large flowered phase of Lupinus argenteus in southeastern Utah had been or was about to be effectively and validly published. That name, it turns out, was never published by Dunn and Harmon and was ef- fectively but not validly published as a nomen nudum by Welsh. To correct that imperfec- tion the name is here published with a Latin diagnosis: Lupinus argenteus var. moabensis Welsh var. nov. [L. argenteus var. moabensis Welsh, nom. nud.; L. argenteus ssp. moabensis Dunn & Harmon ex Welsh, nom. nud.]. A var. ar- genteo caeterius persimili floribus maximus, alis 12-14 (nec 8-12) mm longis differt. Type: USA: Utah. Grand Co.: Professor Valley Ranch, T24S, R23E, S27, 1,342 m elev., sandy soil, 28 April 1977, S. L. Welsh and K. Taylor (Mastin) 14632 (Holotype BRY; Isotype NY, and 8 others distributed previ- ously). ADDITIONAL SPECIMENS: Utah. Grand Co., 5 km W of Deadhorse Point, 26 May 1950, W. P. Cottam 12107; ibid., 2 km S of Landscape Arch, 19 May 1949, B. F. Harrison 11414; ibid., 3 June 1963, S. L. Welsh & G. Moore 2028: ibid, Between Moab and Castleton on the Colorado River, 13 May 1933, W. P. Cot- GREAT BASIN NATURALIST Vol. 46, No. 2 | tam 5620; ibid., T25S, R24E, S29, Castle Val- | ley, 18 May 1982, S. L. & E. R. Welsh 21154; ibid., T24S, R23E, S26, Professor Valley | Ranch, S. L. Welsh & K. Taylor (Mastin) 14631; ibid., T25S, R23E, $9, ca 1 km east of | Castle Rock, 28 Apr. 1984, S. L. Welsh & D. | Trotter 22719. Emery Co., T22S, R8E, S6, ca 15 km E of Emery, 13 May 1981, S. L. Welsh 20462; ibid., T24S, R9E, S18, San Rafael Swell, 22 May 1980, J. G. Harris 731; ibid., Rock Springs Bench, NE of Cathedral Valle | Jct., 10 June 1973, H. K. Harrison 1026. San Juan Co., Road to Upheaval Dome, Island in the Sky, 9 July 1964, G. Moore 200, 213; ibid., Island in the Sky, 16 May 1965, G. | Moore 392 (all BRY). This is the early flowering, large flowered | phase of L. argenteus that occurs at low eleva- - tions, mainly along sandy washes in mixed and warm desert shrub communities. During a study of hanging gardens in the} Glen Canyon National Recreation Area, a cu- rious daisy was discovered in a moist alcovell near the confluence of Glen Canyon and Es- | calante Canyon. The seepy alcove faces to the’ south and extends along a horizontal bedding plane for several hundred feet, forming a se- | ries of minor alcoves. Traditional hanging gar-| den plants, such as Primula specuicola, are present in the garden community. The daisy occurred within the hanging garden assem-_ blage mainly at the base of the face wall, | where some detritis has accumulated. Only a! few species of the enormous genus Erigeron. occur in this habitat in Utah. This one seems. to be specialized for growth in the moist, evi- dently saline substrate that is constantly re-. newed by addition of sand from the alcove. face. It is similar in size of flower heads and. general aspect with Erigeron abajoensis, Crongq., a species of montane sites in southern. Utah. The alcove daisy differs from its mon- tane counterpart in having linear to narrowly, oblanceolate basal and cauline leaves, fewer | pappus bristles, and glands in addition to strigose to spreading stiff hairs on the involu- cral bracts. The species is named and de- scribed as follows. Erigeron zothecinus Welsh sp. nov. Planta similis Erigeronte abajoensi Cronq. in capit- ) ulis et bracers sed in foliis angustioribus, pappo paucioribus, et bracteis glandulosis am fert. April 1986 Type: USA: Utah. Kane Co., T40S, R91/2E, $36, GCNRA, Lake Powell, N Escalante hanging gardens, ca 1,140 m elev., Navajo Sandstone, 29 May 1983, S. L. Welsh 22115 (Holotype BRY; 5 isotypes to be distributed). ADDITIONAL SPECIMENS: Utah. Kane Co., same locations as the type, 29 May 1983, S. L. Welsh 22128; ibid, 24 May 1984, S. L. Welsh 22860, both BRY. In 1979 Sherel Goodrich discovered a pop- ulation of Cleomella palmerana Jones north of Split Mountain in Uintah County, Utah. The material was routinely assigned to the species as it was understood in eastern Utah. How- ever, the habit of growth and fruit characters differ from the body of the species. The raceme stands above the foliage, and the fruit is distinctly horned, resulting in fruit 8-9 mm wide, not 3-5 mm wide as in the material from south of the Uinta Basin. Because of these differences the plants are here designated as follows: Cleomella palmerana Jones var. goodrichii ‘Welsh var. nov. Ab. Cleomella palmerana var. palmerana in fructu latioribus et cornuto differt, et similis Cleomella plocasperma Wats. in fructu ambitu sed in foliis latioribus differt. Type: USA: Uintah Co., T3S, R24E, $25, » Rainbow Draw, 1,647 m elev, Morrison For- mation, eroded slopes of heavy raw, vertisol- like clay, soil violently effervescent with 10% HCl, with Machaeranthera venusta, Phacelia _demissa, Astragalus flavus, and Atriplex cor- rugata, 26 May 1979, S. Goodrich 12312 (Holotype BRY; isotypes distributed previ- ously). On 8 May 1955, I collected a peculiar Ara- bis specimen in Little Rainbow Park, Uintah County, while doing field work for my first attempt at ecological and, as it turned out, taxonomic studies. The plant was sent to a specialist in the genus for determination prior to completion of the project in 1957. It was dentified initially as A. microphylla Nutt. ex I. &G. Subsequently, in 1976, the plant was sent again to the specialist, and this time it was determined as A. microphylla, with some fea- tures of A. fernaldiana Rollins. In May 1979, blants of a similar nature were discovered in ones Hole, east of the initial find. The plants _ aave been compared to both A. microphylla nd A. fernaldiana, of which abundant mate- WELSH: NEW UTAH PLANT TAXA 263 rial is now at hand for comparison. The simi- larity to A. microphylla is superficial indeed, but it is very much like the material of the type variety of A. fernaldiana. The specimens from Rainbow Park and Jones Hole differ from the Nevada material of the type variety in having smaller flowers and from the species in having narrower siliques and shorter styles. The plants in question are separated from A. fernaldiana geographically by the width of Utah. They are named as follows: Arabis vivariensis Welsh sp. nov. Planta persimilis Arabe fernaldiana Rollins sensu lato, differt in stylis brevioribus (0.5 nec 1 mm), floribus parvioribus [prater var. sty- losam (Wats.) Rollins], et siliquis angus- tioribus (1-1.5 nec 1.5-2 mm). Plants perennial, forming mats or carpets to 1 m wide or more, the caudex branches bear- ing marcescent leaf bases, the branches of several seasons evident back from the branch ends, horizontally spreading to decumbent, finally erect and bearing flowering stems of the season or terminating in leafy rosettes, the flowering stems mainly 8-32 cm tall, puberu- lent with minute dendritic trichomes or glabrous above; basal leaves and those of the innovations 0.7-3 cm long, 1.2-4 mm wide, oblanceolate to elliptic, the blade tapering toa long, slender petiole, green to gray, pubescent overall with minute dendritic hairs, acute; cauline leaves 3-13 mm long, 1-2.5 mm wide, oblong to lanceolate or lance- subulate, puberulent to glabrous, much re- duced upward; pedicels ascending to erect, 5-15 mm long in fruit, glabrous or minutely puberulent; sepals 2.5-4.5 mm long, the outer pair gibbous at the base, the inner ones less so, commonly purplish, glabrous to puberu- lent; petals 7-9 mm long, tapering to a basal claw, purplish; siliques 3-7 cm long, 1-1.5 mm wide, glabrous, nerved at the base, erect-as- cending, typically curved or contorted, the style to 0.5 mm long; seeds uniseriate, ca 1.2 mm long, narrowly winged apically. Type: USA: Utah. Uintah Co., T3S, R25E, S1, Jones Hole, National Fish Hatchery, 1,830 m, sandy calcareous gravel, Morgan Formation, 16 May 1979, S. L. Welsh & E. C. Neese 18341 (Holotype BRY; 10 isotypes to be distributed). ADDITIONAL SPECIMENS: Utah. Uintah County, Little Rainbow Park, Dinosaur Na- 264 tional Monument, Navajo Sandstone, sandy soil, juniper association, at 1,525 m, 8 May 1855, S. L. Welsh 152; same locality as the type, 20 June 1980, E. Neese & S. L. Welsh 8978; ibid., 24 May 1982, N. D. Atwood 8822 (all BRY). During the spring of 1981, anew Draba was discovered by Ronald J. Kass, growing in crevices of granite cliffs in Goshute Canyon, Deep Creek Mountains, Tooele County, Utah. The plants appear to be shade requiring mesophytes of north-facing outcrops. An ex- tensive field search of other canyons did not yield evidence of other specimens of the spe- cies. It is easily separated from other scapose and subscapose perennial drabas by its slen- derly petiolate leaves, definite caudices clothed with persistent, filiform leaf bases, and long persistent scapes. The species does not appear to have close allies among our nu- merous taxa, but does share certain morpho- logical features with D. asprella. The strongly branched caudex, with persistent, marces- cent leaf bases and narrowly oblanceolate to spatulate leaves is diagnostic from D. as- prella. The species is named in honor of its discoverer, as follows: Draba kassii Welsh sp. nov. Planta similis Draba asprella generalis sed in caudicibus vestitis petiolis marcescentibus valde, foliis angustioribus et glabis supra et interdum in- fra, et pilis simplicibus vel furcatis differt. Perennial, caespitose, from a definite, branching, subligneus caudex, this clothed with persistent, filiform, threadlike, leaf bases; stems 2-13 cm tall, glabrous or spar- ingly hirsute with mixed simple and forked to dendritic hairs; leaves all basal, rarely with 1 GREAT BASIN NATURALIST cauline, 1.8-4.8 cm long, 2-6 mm wide, nar- rowly oblanceolate to spatulate, entire or ob- Vol. 46, No. 2 | scurely and sparingly denticulate, green, the | surfaces glabrous, sparingly ciliate with sim- | ple or forked hairs; racemes simple, 2- to 9- | flowered, elongating in fruit; pedicels 2-10 | (15) mm long, ascending, glabrous; sepals 1.5- | 2.4 mm long, greenish, sparingly hairy, with © simple or forked hairs; petals 4.6-5.9 mm long, yellow, obovate-spatulate, rounded; sil- | icles 3-10 (14) mm long, 0.8-2.5 mm wide, elliptic to oblong, glabrous; styles 1-2 mm long; seeds 2-14. Type: USA: Utah, Tooele Co., T10S, | RI8W, SW1/4 S36, Deep Creek Mtns., | Goshute Canyon, granite cliff, where soil ac- | cumulates in cracks, at 2,135 m elev., on _ north exposure, with Juniperus osteosperma, © Pinus monophylla, Lomatium grayii, etc., 8 June 1981, R. J. Kass, with Herrick 330 (Holo- | type BRY; 5 isotypes to be distributed). ADDITIONAL SPECIMENS: Utah, Tooele Co., ibid, 20 May 1981, R. J. Kass 284 (BRY); ibid, 23 April 1981, R. J. Kass, with Alan Taye 243, (BRY). The plants begin to flower while snow is still on the ground in mid-April and continue to, flower to early June. They occur on granite at 2,135 to 2,500 m elevation. LITERATURE CITED WELSH, S. L. 1978. Utah flora: Fabaceae (Leguminosae). | Great Basin Nat. 38:225-367. Basin Nat. 43: 179-357. WELSH, S. L. AND J. L. REVEAL. 1977. Utah flora: Brassi- | caceae (Cruciferae). Great Basin Nat. 279-365. a 37 | _____. 1983. Utah flora: Compositae (Asteraceae). Great. | NEW SYNONYMY AND NEW SPECIES OF AMERICAN BARK BEETLES (COLEOPTERA: SCOLYTIDAE), PART XI Stephen L. Wood! ABSTRACT.—The following new generic synonymy is proposed: Coptodryas Hopkins (=Microperus Wood), Cyrto- genius Strohmeyer (=Carpophloeus Schedl, Taphroborus Nunberg), Glostatus Schedl (=Ctonocryphus Schedl, Rhopalocryphus Nunberg), Hylurgops LeConte (=Hylescerites Schedl), Hypothenemus Westwood (=Ernophloeus Nunberg), Monarthrum Kirsch (=Eupteroxylon Eggers), Terminalinus Hopkins (=Kelantanius Nunberg), Xylechinus Chapuis (=Pruniphagus Murayama), Xylocleptes Ferrari (=Hylonius Nunberg). New combinations include: Pityoph- thorus anticus Sched] is transferred to Araptus; Hylesinus machilus Sched] is transferred to Phloeosinus; Phloeophtho- rus acaciae Lea is transferred to Phloeotribus; Blastophagus squamosus Schedl is transferred to Polygraphus; Chramesus semibrunneus Eggers is transferred to Pseudochramesus; Dacryophthorus capensis Sched is transferred to Xylechinus; Pseudochramesus imperialis Sched] is transferred to Xylechinus; and Hoplitontus abyssinicus Sched is transferred to Xylocleptes. New specific synonymy includes: Hypothenemus sundaensis (Eggers) (=Ernophloeus costalimai Nunberg). A note on the South American Hylesinus antipodius Sched is included. New names are proposed as follows: Hylesinopsis kenyae for africanus (Sched] 1963) (from Alniphagus ) and Hylesinopsis ugandae for africanus (Schedl 1965) (from Hylesinus ). Species described as new to science include: Ambrosiodmus ferus (Mexico), Ambro- siodomus paucus (Costa Rica), Carphoborus bicornis (USA), Chaetophloeus pouteriae (Mexico), Cnemonyx euphor- biae (Mexico), Corthylus convexifrons (Venezuela), Corthylus senticosus (Mexico), Corthylus sentosus (Mexico), Cryptocarenus pubescens (Brazil), Cryptocarenus spatulatus (Mexico), Dendrocranulus mexicanus Mexico), Hylesi- nus caseariae (Mexico), Pityophthorus levis (USA), and Trischidias exigua (Mexico). On the following pages are recorded syn- onymy and nomenclatural changes that affect new synonymy of 11 genera and one species and new combinations for eight species. Two new names are proposed for new junior homonyms that were created by the transfer of species from one genus to another. In addi- tion to these changes, 14 species are de- scribed as new to science in the genera Am- brosiodmus (2), Carphoborus (1), Chaeto- phloeus (1), Cnemonyx (1), Corthylus (3), Cryptocarenus (2), Dendrocranulus (1), Hylesinus (1), Pityophthorus (1), and Tri- schidias (1). These species are from the USA (2), Mexico (9), Costa Rica (1), Venezuela (1), and Brazil (1). Under each heading the species are listed alphabetically by genus and species. GENERIC SYNONYMY Coptodryas Hopkins Coptodryas Hopkins, 1915, U.S. Dept. Agric. Rept. 99:10, 54 (Type-species: Coptodryas confusa Hopkins, original designation) Microperus Wood 1980, Great Basin Nat. 40:94 (Type- species: Xyleborus theae Eggers, original designation). New synonymy. The unique female holotype of Coptodryas confusa Hopkins was examined and compared to more than 20 species previously placed by me in Microperus Wood. Notes from my pre- vious examination of this type that were dated 1955 indicated that it was a true Xyleborus. However, the current revision of the tribe places it elsewhere. In my collection under the name Xyleborus cryphaloides Eggers, as determined by F. G. Browne, are two differ- ent species, the larger of which is identical to the type of C. confusa. I have not attempted to locate the type of cryphaloides to review its specific status, but, regardless of the outcome of such a review, the name Microperus Wood must be placed in synonymy as indicated above. Cyrtogenius Strohmeyer Cyrtogenius Strohmeyer, 1910, Ent. Blatt. 6:127 (Type- species: Cyrtogenius bicolor Strohmeyer, monobasic) Carpophloeus Schedl, 1959, Tijdschr. Ent. 101:143 (Type-species: Carpophloeus rugipennis Schedl, mon- obasic). New synonymy Taphroborus Nunberg, 1961, Ann. Mag. Nat. Hist. (13)3:617 (Taphroborus vaticae Nunberg, original des- ignation). New synonymy life Science Museum and Department of Zoology, Brigham Young University, Provo, Utah 84602. 265 266 Schedl named the monobasic Carpophloeus for rugipennis Sched on the basis of the 3-seg- mented antennal funicle. Because the number of funicular segments is variable among the smaller species of Cyrtogenius, there is no way to distin- guish Schedl’s genus; consequently, it must be placed in synonymy as indicated above. The spe- cies rugipennis was based on two male and one female syntypes. Sched (1979, Katalog der wis- senschaftlichen Sammlungen des Naturhis- torisches Museums in Wien 3 (Heft 2):216) des- ignated the male in his collection as the lectotype of this species. The lectotype was examined. Nunberg named Taphroborus vaticae from four specimens of undesignated sex. A “holotype” and paratype (both mounted on the same card on one pin) were deposited in the British Museum (Natural History). Be- cause there is no way to tell which specimen is the type, both must be regarded as syntypes. The specimen that seems to fit the description most completely is a female and is. still mounted on the card. The other specimen is a male and has subsequently been dislodged from the card and is missing the abdomen and ventral parts of the thorax. The female syn- type is here designated as the lectotype and the male the allotype of Nunberg’s species. This species falls well within the genus Cyrto- genius. For this reason, Taphroborus is placed in synonymy as indicated above. Glostatus Schedl Glostatus Schedl, 1939, Rev. Zool. Bot. Afr. 32:386 (Type-species: Glostatus declividepressus , monobasic) Ctonocryphus Schedl, 1941, Rev. Zool. Bot. Afr. 34:398 (Type-species: Ctonocryphus xyloctonus Sched, monobasic) Rhopalocryphus Nunberg, 1967, Rev. Zool. Bot. Afr. 76:320 (Type-species: Rhopalocryphus seydeli Nun- berg, monobasic). New synonymy Following my examination of the types of most of the species of Glostatus, Ctonocry- phus xyloctonus Schedl, and Rhopalocryphus seydeli Nunberg, I see only one moderately variable genus. The bisinuate basal margin of the pronotum and deeply impressed elytral striae of Ctonocryphus intergrade through Rhopalocryphus to Glostatus to such an ex- tent that there is no possibility of making a generic division within the group. Both Ctonocryphus and Rhopalocryphus are placed in synonymy as indicated above. GREAT BASIN NATURALIST Vol. 46, No. 2 Hylurgops LeConte Hylurgops LeConte, 1876, Proc. Amer. Philos. Soc. 15:389 (Type-species: Hylastes pinifex Fitch =Hylurgops rugipennis pinifex (Fitch), subsequent designation by Hopkins 1914:123). Hylescerites Schedl, 1947, Zentralbl. (Type-species: Hylescerites monobasic). New synonymy Ges. granulatus Ent. 2:29 Schedl, Sched] named the monobasic fossil genus Hylescerites based on H. granulatus Sched] (1947:30) from Baltic amber. Neither the de- scriptions nor the photograph of the holotype | indicate any characters that distinguish this — genus and species from Hylurgops. In the | absence of distinguishing generic characters, | Hylescerites is placed in synonymy under the | older name as indicated above. Hypothenemus Westwood Hypothenemus Westwood, 1836, Trans. Ent. Soc. Lon- | don 1:34 (Type-species: Hypothenemus eruditus West- | wood, monobasic) Ernophloeus Nunberg, 1958, Acta Zool. Cracoviensia | 2:484 (Type-species: Ernophloeus costalimai Nunberg =Stephanoderes sundaensis Eggers, original designa- | tion). New synonymy | The female holotype and two female paratypes of Ernophloeus costalimae Nun- berg were examined and compared directly to. my female homotypes of Stephanoderes sun- | daensis Eggers. Because they are quite clearly congeneric with Hypothenemus erudi- tus Westwood, Nunberg’ss genus must be — placed in synonymy under Hypothenemus | and his species under sundaensis as indicated | above. i | | i Monarthrum Kirsch Monarthrum Kirsch, 1866, Berliner Ent. Zeitschr. 10:213 (Type-species: Monarthrum chapuisii Kirsch, | monobasic) Eupteroxylon Eggers, 1936, Rev. de Ent. 6:392 (Type- | species: Eupteroxylon comatum Eggers, monobasic) | The female holotype of Eupteroxylon coma-. | | | I } t | i) | it tum Eggers is in the laterale Eichhoff species: i group of Monarthrum. The holotype of coma-. tum was compared to a series of females all | (Cosmocorynus ) latus Sched] from Colombia. The Eggers species is smaller but has almost, identical antennae, and they share the same | general sculptural destan of frons and elytra. ie They are obviously congeneric. For this rea-\ — son, Eupteroxylon is placed in synonymy as indicated above. | April 1986 Terminalinus Hopkins Terminalinus Hopkins, 1915, U.S. Dept. Agric. Rept. 99:10, 57 (Type-species: Terminalinus terminaliae Hopkins, original designation) Kelantanius Nunberg, 1961, Ann. Mag. Nat. Hist. (13)3:621 (Type-species: Xyleborus punctatopilosus Schedl, original designation). New synonymy The unique female holotype of Terminali- nus terminaliae Hopkins was examined and compared to material in my collection. It is in the same species group with Xyleborus major Stebbing, X. latus Eggers, X. siclus Schedl, X. pseudopilifer Schedl, X. postecipilosus Schedl, pilifer Eggers, pseudomajor Sched], xanthophyllus Schedl, and macropterus Schedl. Because this species group makes up _part of the genus previously known as Kelan- tanius Nunberg, Nunberg’s name must be placed in synonymy under the older name Terminalinus as indicated above. Xylechinus Chapuis Xylechinus Chapuis, 1869, Synopsis des Scolytides, p. 36 _ (Type-species: Hylesinus (Dendroctonus) pilosus Ratzeburg, monobasic) Pruniphagus Murayama, 1958, Bull. Fac. Agric. Ya- maguti Univ. 9:930 (Type-species: Pruniphagus gum- mensis Murayama, original designation) The “holotype” and “allotype” of Prunipha- gus gummensis Murayama are mounted on _ the same pin. Because the description is com- posite, both specimens are female, and there is no clue in the description as to which is the WoobD: AMERICAN BARK BEETLES 267 cleptes bispinus (Duftschmidt); consequently, Hylonius must be placed in synonymy as indi- cated. The status of Nunberg’s species was not investigated. NEW COMBINATIONS Araptus anticus (Schedl), n. comb. Pityophthorus anticus Schedl, 1976, Ent. Abh. Mus. Tierk. Dresden 41:66 (Holotype, female; Rio Negro, Brasil; Wien Nat. Mus.) In the original treatment of Pityophthorus anticus Schedl, the sexes were reversed. This species is a member of the genus Araptus and is here transferred to that genus. Phloeosinus machiius (Schedl), n. comb. Hylesinus machilus Schedl, 1959, Indian For. Rec., n.s., Ent. 9(8):173 (Paratype; Chachpur, Chakrata, Uttar Pradesh, India; Wien Nat. Mus.; holotype lost, if it ever existed) None of the type specimens, including the holotype of this species, that were sent by the Forest Research Institute to Schedl for study were ever returned to the FRI. After examin- ing the loan sheets at FRI, the nontype mate- rial returned by Schedl, and the FRI speci- mens retained by Schedl from that loaned material, I suspect that the holotypes cited by Sched] in the descriptions of FRI species never existed. A consequence of that action is seen in the description of Hylesinus machilus Schedl. The head and prothorax of Schedl’s paratype of this species are missing. As a re- sult, Schedl named this species in the wrong type, I here designate the upper female that \ has one unbroken antenna (the club is lost on | the other side of the type and on both sides in the lower specimen) as the lectotype of this species. This species is very closely allied to | Xylechinus padi Wood but it is distinct. These two species are members of Xylechinus, al- ‘though the scales on the pronotum and elytra are rather small and slender and the setae on the metepisternum are palmately divided. This placement requires that Pruniphagus be placed in synonymy as indicated above. Xylocleptes Ferrari Xylocleptes Ferrari, 1867, Die Forst- and Baumsucht- schadlichen Borkenkafer, p. 37 (Type-species: Bostrichus bispinus Duftschmidt) 'Hylonius Nunberg, 1973, Exploration du Parc Nacional des Virunga (2)23:16 (Type-species: Hylonius brunneus Nunberg, original designation). New synonymy | __ A paratype of Hylonius brunneus Nunberg was examined. It is congeneric with Xylo- | genus even though it had been sent to him under the clearly marked manuscript designa- tion of Phloeosinus machili Beeson (nomen nudum). I have examined more than 50 specimens of this species from Uttar Pradesh, several of which bear data identical to that published by Schedl. As indicated above, it must be trans- ferred to Phloeosinus. Phloeotribus acaciae (Lea), n. comb. Phloeophthorus acaciae Lea, 1910, Proc. Roy. Soc. Victo- ria, n.s., 22:146 (Syntypes; Tasmania) A series of this species was found in the Sched] Collection (Wien Nat. Mus.). Because the genus is unknown in the area from south- ern Asia to Australia, except for this species, it is of special interest. The three terminal seg- 268 ments of the antenna are no wider than those of the funicle, smaller than in rhododactylus (Marsham) of Europe. This and other primi- tive characters suggest that this species was derived from South American stock prior to the Tertiary and has been preserved with lit- tle modification. The scutellum is not visible. Sched] (1938, Proc. Linn. Soc. N.S.W. 83:216) erroneously placed the species in Xylechinus, a genus quite unrelated to the one to which it belongs. Polygraphus squamosus (Schedl), n. comb. Blastophagus squamosus Schedl, 1975, Ent. Basil. 1:384 (Holotype; Bhutan, Dorjula; Nat. Mus. Basel) The species named Blastophagus squamo- sus Sched] is represented in the Sched] mate- rial at Wien by one paratype. This specimen fits the description of the species, but it is a member of Polygraphus and must be trans- ferred to that genus. Pseudochramesus semibrunneus (Eggers), n. comb. Chramesus semibrunneus Eggers, 1950, Ent. Blatt. 45—46:145 (Holotype, male; Brasil; Wien Nat. Mus.) Schedl had the sexes reversed in this genus. The male holotype of Chramesus semibrun- neus Eggers was examined. It is a member of the genus Pseudochramesus and is here trans- ferred to the genus as indicated above. Xylechinus capensis (Schedl), n. comb. Dacryophthorus capensis Sched], 1971, Opusc. Zool. Munchen 119:6 (Holotype, female; Cape Prov., South Africa; Wien Nat. Mus.) The female holotype of Dacryophthorus capensis Schedl belongs to the genus Xylechi- nus and is here transferred to that genus. Xylechinus imperialis (Schedl), n. comb. Pseudochramesus imperialis Schedl, 1958, Acta Zool. Lil- loana 16:39 (Lectotype, male; Wien Nat. Mus., present designation) The original description of Pseudochrame- sus imperialis Schedl is composite. The “holotype” cited by Sched] (1979:122) and so labeled in his collection is here designated as the lectotype of this species. An abundance of characters indicates that it belongs to the genus Xylechinus. A second male in the Sched] Collection is labeled as the “holotype” GREAT BASIN NATURALIST Vol. 46, No. 2 of Xylechinus calvus Schedl. Because I have © found no description associated with this | name, itis presumed tobe anomen nudum. Xylocleptes abyssinicus (Schedl), n. comb. Hoplitontus abyssinicus Sched], 1965, Rev. Ent. Mogambique 8:364 (Holotype; Abyssinien; Wien Nat. Mus.) The holotype of Hoplitontus abyssinicus — Sched] is almost totally covered by glue. It definitely is a member of the Dryocoetini and _ probably is in Xylocleptes. A more precise identification must await a time when the glue — can be dissolved to expose additional charac- | ters. NEW NAMES Hylesinopsis kenyae, n. n. Alniphagus africanus Schedl, 1963, Ent. Abh. Ber. Mus. | Tierk. Dresden 28:259 (Holotype; Riff Valley, Kenya; | Wien Nat. Mus.) Preoccupied | The species named Alniphagus africanus Schedl is a member of-the genus Hylesinop- sis. Because the transfer of this species causes — it to become a junior homonym of africanus (Eggers 1933), areplacement name is needed. | The new name kenyae is proposed as a re- | placement for the Sched] species. | | Hylesinus africanus Schedl, 1965, Nova Taxa Ent. 38:4 | (Holotype; Mpanga, Uganda; British Mus. Nat. Hist.). | Preoccupied Hylesinopsis ugandae, n. n. The transfer of Hylesinus africanus Schedl to Hylesinopsis makes this species a junior) secondary homonym of africanus (Eggers. 1933). The new name ugandae is proposed as. a replacement for the Sched] species. SPECIAL NOTE Hylesinus antipodius Sched] | : Hylesinus antipodius Schedl, 1951, Rev. Chil. Ent. 1:17, (Syntypes; Rengo, Chile; Wien Nat. Mus. and Museo Nacional de Historia Natural, Santiago) Hylesinus antipodius Sched] is the ol | known true member of this genus in America |. south of Guatemala. The elytral scales are more slender than in any North American, species. It appears to be more closely allied to H. cordipennis Lea, from Australia, than to April 1986 any other species known to me. If this is cor- rect, then H. antipodius would probably have been derived from Australian stock prior to the Tertiary when island hopping was still possible between these separating land masses. NEw TAXA Ambrosiodmus ferus , n. sp. This species is clearly allied to divexulus Wood, although only one suture is on the posterior face of the antennal club. It is distin- guished from divexulus by the larger size, by the more gradual, more finely punctured ely- tral declivity, and by other characters de- scribed below. FEMALE.—Length 2.6 mm _ (paratypes 2.5-2.7 mm), 2.4 times as long as wide; color black. _ Frons about as in divexulus except a weak median carina present on upper half. _ Pronotum as in divexulus except summit -more distinct, asperities behind summit slightly larger and closer. Elytra similar to divexulus except discal strial punctures slightly larger, not as deep, interstrial punctures smaller, not as deep; de- _clivity more gradual, particularly on upper half, interstriae 1 not elevated; vestiture finer, slightly larger. TYPE LOCALITY.—Jalapa, Veracruz, Mex- ‘ico. TYPE MATERIAL.—The female holotype and \ five female paratypes were taken at the type locality on 16-VITI-1983, FANM-33, from Quercus, by Felipe A. Noguera. _ The holotype and paratypes are in my col- ‘lection. ) Ambrosiodmus paucus, n. sp. This species is distinguished from divexulus ‘Wood by characters described below. FEMALE.—Length 1.9 mm _ (paratypes 1.8-1.9 mm), 2.2 times as long as wide; color very dark brown. _ Frons about as in divexulus except central ' three-fourths without reticulation, shining. ' _ Pronotum similar to divexulus except retic- " \ulation absent, asperities on posterior half dis- sinctly larger, closer. | Elytra similar to divexulus except discal ' »ounctures of medium size, confused, striae rn Woop: AMERICAN BARK BEETLES 269 not indicated; declivity as in divexulus except strial and interstrial punctures distinctly smaller, interstriae 1 armed as on 3: declivital vestitute stouter, of more uniform length. TYPE LOCALITY.—Isla del Coco, Costa Rica. TYPE MATERIAL.—The female holotype and three female paratypes were taken at the type locality in April 1980 by George Stevens. The holotype and paratypes are in my col- lection. Carphoborus bicornis, n. sp. This species is distinguished from bifurcus Eichhoff by the frontal and declivital charac- ters that are described below. FEMALE.—Length 1.5 mm _ (paratypes 1.3-1.6 mm), 2.5 times as long as wide; color dark brown, vestiture pale. Frons convex, flattened on lower two- thirds on median half, surface reticulate; a pair of rather widely spaced, conspicuous hornlike spines just below upper level of eyes on median two-thirds, each two or more times as high as basal width in Alabama series, about as high as wide and blunt in Florida series, ventral surface of each spine with a few scales. Pronotum and elytra as in bifurcus except declivital interstriae 3 with crest of elevation wider, denticles more numerous and much more strongly confused. MALE.—Similar to female except frons as in male bifurcus. TYPE LOCALITY.—Fayette, Alabama. TYPE MATERIAL.—The female holotype, male allotype, and two paratypes were taken at the type locality V-159-10037, from Pinus, by Walter Grimes. Fourteen paratypes were taken at Archibold Biological Station, Lake Placid, Florida, 7 March 1984, Pinus clausa, by Mark Deyrup. The holotype, allotype, and paratypes are in my collection. Chaetophloeus pouteriae, n. sp. This species is distinguished from insularis (Blackman) as indicated by characters de- scribed below. This is the third species in the group lacking submarginal crenulations be- hind the marginal row at the base of the elytra. MALE.—Length 1.2 mm (paratypes 1.1— 1.2 mm), 1.8 times as long as wide; color dark brown, vestiture pale. 270 Frons as in insularis except impression less extensive above eyes. Pronotum as in insularis except punctures much smaller, more definite, spaces between punctures almost smooth; scales shorter, broader, those on anterior margin conspicu- ously longer. Elytraas in insularis except strial punctures deeper, very slightly larger; scales in ground cover much stouter. FEMALE.—Similar to male except frons al- most flat on lower half, convex above. TYPE LOCALITY.—Campo Experimental. INIF, Escarcega, Campeche, Mexico. TYPE MATERIAL.—The male holotype, fe- male allotype, and six paratypes were taken at the type locality on 9-I-1984, AEV-85, Poute- ria campechana, by A. Estrada V. GREAT BASIN NATURALIST Vol. 46, No. 2 Corthylus convexifrons, n. sp. This species is unique and does not fit into any known species group. The female frons is convex and glabrous, the general pronotal and elytral features are much as in the larger Cor- thylocurus species. FEMALE.—Length 3.4 mm _ (paratypes 2.9-3.6 mm), 2.4 times as long as wide; rather light reddish brown. Frons evenly, strongly convex, median fourth with a slight elevation, a small median tubercle on elevation; surface finely reticu- late, punctures minute, sparse; glabrous. An- tennal club slightly asymmetrical, two feebly © procurved sutures present; posterior face | bearing a small tuft of hair arising from lateral - half of all three segments, extending shanty beyond tip of club. Pronotum 1.1 times as long as wide; side | almost straight and subparallel on basal half, _ broadly rounded in front; anterior margin The holotype, allotype, and paratypes are in my collection. Cnemonyx euphorbiae, n. sp. This species is distinguished from splen- dens Wood by characters described below. FEMALE.—Length 2.5 mm (male paratypes 2.3 mm), 2.04 times as long as wide; color reddish brown. Frons with median two-thirds from epis- toma to upper level of eyes rather abruptly, concavely impressed, impressed area dense- ly, rather coarsely punctured and ornamented by numerous, erect, rather stout setae of uni- form length; epistoma not subcarinate as in splendens. Antenna about as in splendens. Pronotum as in splendens except anterior constriction more distinct, punctures coarser and closer. Elytra about as in splendens except inter- strial punctures confused, declivital inters- triae less strongly elevated, declivital inters- trial setae much smaller, almost obsolete, hairlike. MALE.—Similar to female except frons con- vex, glabrous, punctures less dense. TYPE LOCALITY.—Canon de Lobos, Yaute- pec, Morelos, Mexico. TYPE MATERIAL.—The female holotype, male allotype, and two male paratypes were taken at the type locality on 14 Marzo 1984, 1,400 m, SM-247, by Edgar Martinez F. The holotype, allotype, and paratypes are in my collection. subcostate, with about eight weak serrations | indicated; posterior half reticulate, punctures | sparse, minute. Glabrous. | Elytra 1.36 times as long as wide, 1. 4A) times as long as pronotum; sides almost! straight and parallel on basal three-fourths, — very broadly rounded behind; disc subreticu- | late, punctures small, shallow, distinct, con- fused. Declivity very steep, shallowly sulcate | on median third; lateral margins on upper half. armed by two pairs of small, blunt tubercles; a’ weak ventrolateral margin indicated on me- dian third. Almost glabrous. MALE.—Similar to female except epistomal elevation poorly formed, anterior margin ol pronotum with two coarse serrations, declivi- | tal impression slightly stronger. TYPE LOCALITY.—La Mucuy, 20 km west 0: Merida, Merida, Venezuela. TYPE MATERIAL.—The female holotyatl male allotype, and 13 paratypes were taken at | the type locality on 12-XI-1969, 2,500 m, No. 131, from an unidentified tree branch, by me. ii Paratypes include: 6 from the type locality, taken 22-XII-1969, No. 207, from a tree | seedling; 19 from La Carbonera Experimenta | Forest, 50 km west of Merida, Merida) } Venezuela, 9-XII-1969, 2,500 m, No. 174) from Neciandne branches; all were taken by i me. The holotype, allotype, and paratypes are in my collection. \ April 1986 Corthylus senticosus , n. sp. This species is distinguished from sentus Wood by characters of the frons, antenna, and elytral declivity as described below. FEMALE.—Length 1.5 mm (allotype 1.8 mm), 2.1 times as long as wide; color very dark brown. Frons extensively excavated as in sentus except vestiture on vertex slightly shorter, lateral spongy areas slightly larger, a large longitudinal, cylindrical, subcarinate eleva- tion extending from just above epistoma to just below middle of frons; excavated area glabrous except for fringe of long hair on ver- tex. Antennal club much less asymmetrical. Pronotum as in sentus. Elytra resembling sentus except disc mostly subreticulate, punctures mostly obso- lete; declivity steeper, interstriae 2 on upper third rather strongly, narrowly elevated and armed by small, pointed denticles to middle of declivity, sutural interstriae not elevated or armed. MALE.—Similar to female except frons con- vex, reticulate, subglabrous; anterior margin or pronotum armed by two serrations. TYPE LOCALITY.—Jalapa, Veracruz, Mex- ico. TYPE MATERIAL. —The female holotype and male allotype were taken at the type locality on 23-X-1983, FANM-77, from Psitacanthus schiedeanus, by Felipe A. Noguera. The holotype and allotype are in my collec- tion. Corthylus sentosus, n. sp. _ This species is distinguished from sentus ~ Wood by characters of the frons, antenna, and elytral declivity as described below. FEMALE.—Length 1.9 mm (allotype and paratype 2.0 mm), 2.3 times as long as wide; color very dark brown. Frons as in sentus except spongy areas more widely separated below but extending well above upper level of eyes laterally, their inner margins on upper half each bearing a row of about six long, coarse setae, setae on vertex Woob: AMERICAN BARK BEETLES Dial ulate, interstriae 2 on middle half distinctly, rather weakly elevated, crest of this elevation armed by a row of four fine denticles. MALE. —Similar to female except frons con- vex, reticulate, subglabrous; anterior margin of pronotum armed by two small serrations. TYPE LOCALITy.—Km 32 on Carretera Patzcuaro-Ario de Rosales, Michoacan, Mex- ico. TYPE MATERIAL.—The female holotype, male allotype, and one female paratype were taken on 31-X-1980, 2,360 m, S-130, from Psiticanthus sp., by T. H., Atkinson and Ar- mando Equihua. The holotype, allotype, and paratype are in my collection. Cryptocarenus pubescens, n. sp. This unique species is the largest, stoutest, most pubescent member of the genus. FEMALE.—Length 3.1 mm, 2.3 times as long as wide; color reddish brown. Frons broadly convex; coarsely, closely, subrugosely punctured from epistoma to ver- tex; median line from upper level of eyes with an impunctate, transversely strigose, low, subcarinate elevation; vestiture fine, rather long, moderately abundant. Antennal sutures more strongly procurved than usual for this genus. Pronotum as long as wide; sides weakly ar- cuate and subparallel on basal half, somewhat narrowly rounded in front, anterior margin armed by 14 serrations; anterior slope armed by numerous asperities of moderate size; pos- terior areas finely punctured, punctures on disc finely granulate. Vestiture of fine, erect, rather abundant hair. Elytra 1.4 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; surface almost smooth and shining, punctures rather small, shallow, con- fused, rather close. Declivity rather steep, convex; surface obscurely reticulate, punc- tures as on disc. TYPE LOCALITY.—Sixty-nine km north of Manaus, Brazil. TYPE MATERIAL. —The unique female holo- type was taken at the type locality on 7-XII- 1979, by George Stevens. The holotype is in my collection. imuch shorter. Antennal club about as in senti- pcosus Wood. _ Pronotum as in sentus except disc without | crenulations. _ Elytra as in sentus except devoid of punc- | tures, uniformly subreticulate; declivity retic- | Cryptocarenus spatulatus, n. sp. This species is distinguished from lepidus Wood by characters described below. FEMALE.—Length 1.8 mm (Paratypes 1.8 mm), 2.5 times as long as wide; color dark reddish brown. Frons as in lepidus except subparallel acicu- lations on upper two-thirds of frons deeper. Pronotum about as in lepidus except asperi- ties much coarser, resembling Hypothene- mus; anterior margin armed by 6-8 serra- tions. Elytra as in lepidus except declivity steeper; strial punctures more deeply im- pressed, interstriae more regularly punc- tured; vestiture extending to base on at least odd-numbered interstriae, regular on poste- rior half of elytra on all interstriae, much more closely spaced than in lepidus, each seta erect, strongly flattened on its distal half, about twice as wide as in lepidus. TYPE LOCALITY.—Sta. Maria Chimalpa, Oaxaca, Mexico. TYPE MATERIAL. —The female holotype and seven paratypes were taken at the type local- ity on 10-II-1984, 250 m, S-977, Stru- thanthus, by Armando Equihua. One para- type is labeled 5 mi N. Mazatlan, Sinaloa, Mexico, 24-VII-1964, H. F. Howden. A spec- imen, not designated as a paratype, is labeled Peru, 12-IX-1963, E. M. Jones, in derus plant intercepted at Miami. The holotype and seven paratypes are in my collection; one paratype is in the Canadian National Collection. Dendrocranulus mexicanus, n. sp. This species is distinguished from the allied costaricensis Eggers by the characters de- scribed below. FEMALE.—Length 2.5 mm (paratypes 2.2— 2.5 mm), 2.7 times as long as wide; color very dark brown. Frons as in costaricensis except punctures mostly finely granulate; vestiture equal in abundance but conspicuously shorter and of darker color. Pronotum as in costaricensis except punc- tured area smaller, asperities slightly larger and more extensive in posterolateral. areas; punctures in discal area mostly with fine gran- ule on lateral margin. GREAT BASIN NATURALIST Vol. 46, No. 2 Elytra as in costaricensis except surface on basal fourth of disc more wrinkled; declivity not quite as steep, strial punctures smaller, fine interstrial granules replace punctures; | vestiture similar but finer on disc and decliv- ity. MALE.—Similar to female except frons more strongly convex above, punctures ob- | scure except laterally, vestiture sparse, incon- spicuous; elytral declivity similar to costari- censis except not as steep, more strongly impressed on interstriae 2, 3 higher, granules on 2 not evident; setae a bit more slender. TYPE LOCALITY.—Naolinco, Veracruz, Mexico. | TYPE MATERIAL.—The female holotype, | male allotype, and six paratypes were taken at | the type locality on 28-I-1984, FANM-120, from Sechium edulis, F. A. Noguera M. The holotype, allotype, and paratypes are | in my collection. Hylesinus caseariae, n. sp. | This species is distinguished from californi- cus Swaine by the more deeply, more broadly impressed male frons, by the less distinct, | slightly longer female frontal carina, by the. less strongly impressed declivital interstriae 2 in both sexes, and by the smaller, stouter scales on pronotum and elytra. | MALE.—Length 2.5 mm (paratypes: male | 2.8 mm, females 2.6 mm), 1.8 times as long as_ wide; color pattern as in californicus except with more dark brown scales, fewer pale’ scales. Frons resembling californicus except im- pression deeper, extending higher on vertex, — much broader, lateral margins higher, more | abruptly rounded to upper level of eyes; me- dian line feebly elevated. Pronotum about as in californicus except rugose-reticulation stronger, asperities ap- parently smaller and more numerous, scales smaller, usually stouter. | Eyer similar to californicus except inter-. striae 2 very slightly less strongly impressed, tubercles on 1 and 3 and base of 2 smaller, — closer, more definite; erect scales on 1 more | slender, each 2.5—3.0 times as long as wide, those on 3-7 distinctly smaller, slender setae , on 8, 9, and basal half of 7 much more slender, hairlike; scales in ground cover smaller, stouter. April 1986 FEMALE.—Similar to male except frons basi- cally convex as in californicus except carina lower and slightly longer; declivital interstriae 3 not impressed, resembling | and 3, erect scales _ normal, not enlarged. TYPE LOCALITY.—Acajete, Veracruz, Mexico. TYPE MATERIAL.—The male holotype, female _ allotype, and one male and one female paratype were taken at the type locality on 22-XI-1983, FANM-92, from Casearia (Flacourtiaceae), by Felipe A. Noguera. The holotype, allotype, and paratypes are in my collection. Pityophthorus levis, n. sp. This species is distinguished from the boycei \ Swaine and the comosus Blackman groups of » species by the virtually impunctate, subglabrous | frons that is not sexually dimorphic and by the elytral declivity that resembles neither group. FEMALE.—Length 2.5 mm (paratypes 2.2—2.7 mm), 2.7 times as long as wide; color dark brown, elytra often reddish brown. Frons broadly convex; surface smooth, shin- ing, a few minute punctures; subglabrous, a few ‘fine, short setae usually present. Antennal club resembling comosus except sutures almost straight. Pronotum resembling comosus except sum- ‘mit not as high, punctures on posterior areas ‘considerably smaller, shallow, much more »widely spaced. Elytra resembling boycei except discal punc- {tures slightly larger and less numerous, con- ‘fused, declivity steeper, lateral convexities more jabrupt, interstriae 2 narrower on lower half, tu- bercles (about 6) conspicuously larger. Vestiture fine, rather long on declivity, absent on inter- striae 2, very short on 1. Woop: AMERICAN BARK BEETLES 273 TYPE LOCALITY.—Fifteen miles northwest of Flagstaff, Arizona (Hart’s Prairie Road). TYPE MATERIAL.—The female holotype, male allotype, and 13 paratypes were taken at the type locality on 18-IX-1984, from lateral shoots of Pinus ponderosa, by M. R. Wagner. Triscnidias exigua, n. sp. This is the fourth species named in this genus. It is distinguished from the closely allied atoma (Hopkins) as described below. FEMALE.—Length 0.8 mm _ (paratypes 0.8—0.9 mm), 2.1 times as long as wide; color very dark brown, almost black. Frons as atoma except without a median impression. Pronotum resembling atoma except ante- rior margin armed by four serrations, asperi- ties higher and apparently less numerous, surface smooth, shining, without any indica- tion of reticulation. Vestiture mostly hairlike, a few stouter setae on posterior areas. Elytra about as in atoma except strial punc- tures much smaller, weakly impressed, inter- strial granules much smaller (half as large), erect interstrial scales of equal length but only half as wide, each about four times as long as wide. TYPE LOCALITY.—Campo Experimental, INIF, Escarcega, Campeche, Mexico. TYPE MATERIAL.—The female holotype and six paratypes were taken at the type locality on 15-II-1984, AEY-111 from Belotia campbelli, by A. Estrada V. After the above description was prepared, a series from southern Florida was examined. These are not part of the type series. The holotype and paratypes are in my col- lection. ENERGY AND PROTEIN CONTENT OF COYOTE PREY IN SOUTHEASTERN IDAHO James G. MacCracken'* and Richard M. Hansen! ABSTRACT.—Gross energy, digestible energy, crude protein, and digestible crude protein were estimated for two leporids and five rodents that were the primary prey of coyotes (Canis latrans) in southeastern Idaho. Digestible protein estimates differed (38%—54%) more than digestible energy (3.5—4.4 kcal), in the prey examined. Information on the energy and nutrient content of small mammals that are the food of coyotes (Canis latrans) is necessary to evalu- ate prey selection (Pyke et al. 1977). In addi- tion, those data are valuable in ecological studies of other predators, and for research on nutrient cycling and energy flow through ecosystems (Golley 1960, Odum et al. 1962, Weigert 1965, Fleharty et al. 1973). Research has shown that energy composition of some small mammal bodies varies seasonally and geographically (Gorecki 1965, Fleharty et al. 1973, Cameron et al. 1979), which indicates that the use of data from the immediate study area may be necessary. The purpose of this study was to estimate the gross and digestible energy and crude protein and digestible crude protein of small mammals in conjunction with a study of coy- ote feeding strategies (Johnson and Hanson 1979, MacCracken and Hansen 1982). Large differences in prey body composition might influence coyote food selection. METHODS Small mammals were collected from the Idaho National Engineering Laboratory (INEL) Site in southeastern Idaho (~113°00'W, 44°00'N). The INEL Site occu- pies about 231,500 ha of the Upper Snake River Plain. The dominant vegetation of the study area was a_ sagebrush/bunchgrass (Artemisia/Agropyron) shrubsteppe. Dr. B. L. Keller of Idaho State University supplied five specimens each of the deer mouse (Per- omyscus maniculatus), Townsend's ground squirrel (Spermophilus townsendii), Ord’s 'Department of Range Science, Colorado State University, Fort Collins, Colorado 80523. kangaroo rat (Dipodomys ordii), and least chipmunk (Eutamias minimus). All speci- mens were trapped during the summer of — 1982 and frozen. Additionally, five specimens each of the black-tailed jackrabbit (Lepus cali- | fornicus ), Nuttall cottontails (Sylvilagus nut-_ tallii), and montane vole (Microtus mon-— tanus) were collected during July 1983 and — frozen. | Frozen specimens were chopped into ap- | proximately 1 cm” pieces, oven-dried at 60 C _ for 72 h, then ground in a Willey Mill to pass © through a 2-mm mesh screen. Samples were | then submitted to the Nutrient Analysis Lab, | Colorado State University, to determine gross _ energy (kcal/g dry matter) by bomb calorime- | try and crude protein (% dry matter) by Kjel- | dahl nitrogen ( 6.25), in duplicate for each | individual animal. The digestible fraction of each species was. estimated using the data of Johnson (1978). | Litvaitis and Mautz (1980) reported similar results from feeding trials with captive coy-. otes. Using Johnson's estimates, digestible | energy and protein were calculated for each. species on a dry weight basis. if Analysis of variance, followed by Tukey's. mean separation procedure, was used to test’ for differences in mean gross energy and crude protein among the species examined (P = 0.05). Adequacy of sample size (N = 5) for’ each species for gross energy and crude protein estimates was assessed using a stan-) dard formula based on the t distribution (Giles 1971:158). Adequate sample size (N,.) preci- sion levels were set so as to permit estimates) within 10% of the mean with 95% confidence. Present address: Institute of Northern Forestry, 308 Tanana Drive, Fairbanks, Alaska 99775-5500. 274 April 1986 MaACCRACKEN, HANSEN: COYOTE PREY 975 TABLE 1. Mean (SE) gross energy (kcal/g dry matter), digestible gross energy’, crude protein (% dry matter), and digestible crude protein of coyote prey on the Idaho National Engineering Laboratory in southeastern Idaho. N, is estimated adequate sample size” for gross energy and crude protein values. Energy Protein Species Gross N. Digestible Crude N. Digestible Lepus californicus 4.8(0. 1) 2 3.8(0.2 64.0(1.3) 2 50.6(1.3) Sylvilagus nuttallii 5.0(0.2) 4 4.1(0.2 65.4(2.5) 5 53.6(2.5) Microtus montanus 5.0(0.2) 4 3.9(0.2 65.4(2.5) 5 51.0(2.5) Peromyscus maniculatus 5. 1(0.1) 1 3.9(0.2 62.5(1.9) _ 3 47.5(1.9) Spermophilus townsendii 5.6(0.4) 19 4.4(0.4 48.5(4.1) 26 38.3(4.1) Dipodomys ordii 4.6(0.1) 2 3.5(0.1 62.7(1.4) 2 47.7(1.4) Eutamias minimus 4.9(0. 1) 2 3.9(0.1 60.4(1.9) 4 48.3(1.9) lEstimates of the digestible fraction (% dry matter) of each species were reported by Johnson (1978). Adequate sample size was determined using a standard formula based on the t distribution (Giles 1971: 158). RESULTS AND DISCUSSION Mean gross energy estimates ranged from 4.6 to 5.6 kcal/g and were greatest (P < 0.03) for the Townsend's ground squirrel (Table 1). The other small mammals examined were similar in energy content except Ord’s Kanga- » roo rat, which was the lowest (P < 0.03). Mean digestible gross energy ranged from 3.5 to 4.1 kcal/g being greatest (P > 0.05) for the Townsend's ground squirrel, and lowest (P > 0.05) for Ord’s kangaroo rat (Table 1). Mean crude protein estimates ranged from 48.5 to 65.4% (Table 1). Crude protein was _ lowest (P < 0.04) in Townsend's ground squir- rel and greatest (P > 0.05) for the Nuttall cottontail and montane vole. Digestible protein ranged from 38.3% for Townsend's ground squirrel to 53.6% for Nuttall cotton- tail. Five individuals of each species were ade- quate to estimate energy and protein content, except for Townsend's ground squirrel (Table 1). The large variation in energy and protein content for Townsend's ground squirrel ap- peared to be related to fat content because some individuals had considerable fat, whereas other did not. This was probably at- ‘tributable to specimens originating from dif- ferent cohorts and/or captured at different _ stages in the annual fat cycle. Our gross energy estimates were similar to _ those of other published studies which exam- tined the same species or genera. Gorecki _ (1965) reported a mean (+ SD) summer esti- “umate of 5.1 (+ 0.5) kcal/g for Microtus ar- valis, which is 0.1 keal/g greater than our estimate for montane voles. Fleharty et al. - examined energy content of four ro- dents, including the deer mouse and prairie vole (M. ochrogaster). They reported sea- sonal extremes ranging from 5.05 to 5.14 kcal/g for the deer mouse and 4.91 to 5.01 keal/g for the prairie vole. The maximum val- ues of Fleharty et al. (1973) are similar to our data for the deer mouse and montane vole. Gorecki (1965) stated that energy values were greatest in summer in his study. Davison et al. (1978) examined energy content of the snow- shoe hare (Lepus americanus ), meadow vole (M. pennsylvanicus ), and white-footed mouse (P. leucopus) collected from October through January. Their data are similar to ours for species of the same genus. Litvaitis and Mautz (1980) presented energy estimates for the snowshoe hare (4.97 kcal/g) and laboratory mouse (Mus musculus) (6.00 kcal/g), but col- lection dates were not given. Both of those estimates are higher than we observed for the black-tailed jackrabbits and deer mouse. Few studies have examined crude protein levels of wild mammals (Davison et al. 1978, Litvaitis and Mautz 1980). Energy is typically the currency used in modeling foraging the- ory and in experiments testing those models (Pyke et al. 1977). However, Davies (1977) suggested that nutrients as well as energy may be important in prey selection, and Pulliam (1974) cited protein as having potential impor- tance. Mean crude protein levels for the black-tailed jackrabbit estimated in our study were 6% to 8% lower than those reported for the snowshoe hare by Davison et al. (1978) and Litvaitis and Mautz (1980). Protein levels for the deer mouse reported here were 3% to 9% greater than those reported for the white- footed mouse (Davison et al. 1978) and labora- tory mouse (Litvaitis and Mautz 1980). 276 Our results and those of Davison et al. (1978) and Litvaitis and Mautz (1980) indicate that percent crude protein varies among spe- cies more than does gross energy. This obser- vation suggests that generalizations about protein content of animal bodies across spe- cies lines are of limited value. However, our conclusions and those of other studies cited herein suggest that gross energy levels are similar among species and locales. To what degree coyotes are able to detect differences in prey body composition is un- known. Digestible energy ranged from 3.5 to 4.4 kcal/g dry matter, or a 25% difference. Apparently, prey abundance, body size, and/ or defensive strategies may be more impor- tant in coyote prey selection. However, di- gestible protein estimates ranged from 38% to 54%, or a 42% difference. Protein could be important in prey selection. Furthermore, other nutrients and trace elements should be studied. ACKNOWLEDGMENTS We thank O. D. Markham, W. J. Arthur, and D. K. Halford for their assistance in this study. R. J. Gates and M. P. Stafford helped collect leporids. B L. Oskroba analyzed the specimens. Dr. B. L. Keller is thanked for the use of rodents captured during his studies on the INEL site. The Agricultural Experiment Station, University of Alaska, Palmer, Alaska, kindly provided facilities for the completion of this research. This study was funded by the U.S. Department of Energy, Idaho Opera- tions Office, INEL Ecology Program. GREAT BASIN NATURALIST Vol. 46, No. 2 | LITERATURE CITED CAMERON, G. H., E. D. FLEHARTY, AND H. A. Watts. 1979. | Geographic variation in the energy content of cot- | ton rats. J. Mammal. 60: 817-820. | Davies, N. B. 1977. Prey selection and the search strategy of the spotted flycatcher (Musciapa striata): afield study of optimal foraging. Anim. Behav. 25: — 1016-1033. Davison, R. P., W. W. Mautz, H. H. Hayes, AND J. B. | HOLTER. 1978. The efficiency of food utilization — and energy requirements of captive female fish- — ers. J. Wildl. Manage. 42: 811-821. FLEHARTY, E. D., M. E. KRAUSE, AND D. P. STINNETT. 1973. Body composition energy content, and lipid cycles of four species of rodents. J. Mammal. 54: 426-438. | GILES, R. H., JR. 1971. Wildlife management techniques. | Wildl. Soc., Washington, D. C. 633 pp. GorECKI, A. 1965. Energy values of body in small mam- mals. Acta Theriol. 23: 333-352. GOLLEY, F. B. 1960. Energy dynamics ofa food chain of an | old-field community. Ecol. Monogr. 30: 187-206. | JOHNSON, M. K. 1978. Food habits of coyotes in southcen- tral Idaho. Unpublished dissertation, Colorado State University, Fort Collins. 77 pp. JOHNSON, M. K., AND R. M. HANSEN. 1979. Food habits of | coyotes on the Idaho National Engineering Labo- | ratory. J. Wildl. Manage. 43: 951-955. Litvaltis, J. A.. AND W. W. Mautz. 1980. Food and energy © use by captive coyotes. J. Wildl. Manage. 44: | 56-61. | MACCRACKEN, J. G., AND R. M. HANSEN. 1982. Seasonal | foods of coyotes in southeastern Idaho: a multivari- __ ate analysis. Great Basin Nat. 42: 45-49. OpvuM, E. P.,C. E. CONNELL, AND L. B. DAVENPORT. 1962. | Population energy flow of three primary consumer | components of old field ecosystems. Ecology 43: | 88-96. PULLIAM, H. R. 1974. On the theory of optimal diets | Amer. Nat. 108: 59-74. Pyke, G. H., H. R. PULLIAM, AND E. L. CHaRNov. 1977 | Optimal foraging: a selective review of theory and tests. Quart. Rev. Biol. 52: 137-154. WEIGERT, R. G. 1965. Energy dynamics of the grasshop- | per populations in old field and alfalfa field ecosys- | tems. Oikos 16: 161-176. | { h ESTIMATES OF SITE POTENTIAL FOR DOUGLAS-FIR BASED ON SITE INDEX FOR SEVERAL SOUTHWESTERN HABITAT TYPES Robert L. Mathiasen', Elizabeth A. Blake!, and Carleton B. Edminster” ABSTRACT. —Estimates of site potential for Douglas-fir based on measured site indexes in 450 stands are compared between 10 southwestern habitat types. Significant differences in site potential are found between the habitat types studied. Site index is currently the most widely used method of evaluating site quality or potential productivity of forest lands in the United States (Jones 1969, Husch et al. 1972, -Daubenmire 1976). Site index is based on the average heights of dominant and codominant trees at a specified index age (usually 50 or 100 years). Because stands of the index age are seldom encountered, site index curves are constructed to allow for estimation of site in- dex for stands older or younger than the index age by interpolation between the curves. Site index curves describe the height growth of hypothetical trees of specified site indexes. The use of habitat types (Daubenmire 1952) to classify forest vegetation is gaining accep- ‘tance by land managers and researchers in the swestern United States (Layser 1974, Pfister 1976, Pfister and Arno 1980). One of the pri- mary uses of habitat types is in timber man- )agement. Habitat types are used to compare regeneration success, succession patterns, cutting methods, and timber productivity and ‘to develop guidelines for collecting seed and planting nursery stock (Pfister and Arno 1980). The use of habitat types to predict forest site productivity potential is proposed by sev- eral investigators. Differences in the rate of neight growth by habitat type are demon- strated for several tree species (Daubenmire 1961, Deitschman and Greene 1965, Stanek 1966, Stage 1975, Hoffman 1976). Significant differences between site indexes are also ‘hhown for habitat types (Stanek 1966, Stage \975, Hoffman 1976). Pfister et al. (1971, 4977) and Steele et al. (1981) use site index | ') Northern Arizona University, School of Forestry, Flagstaff, Arizona 86011. curves and normal yield tables to estimate yield capability for habitat types in Montana and Idaho. Southwestern forests are becoming more intensively managed for timber production than in the past. However, growth and pro- ductivity data are presently limited (Gottfried 1978). Habitat type classifications are recog- nized for these forests, but little information is available on the timber productivity potential for these habitat types (Moir and Ludwig 1979, Hanks et al. 1983, Alexander et al. 1984). Jones (1974) provides a summary of the silviculture of southwestern mixed conifer forests and emphasizes a need for improving their management based on the application of habitat types or stand types. This study pro- vides additional quantitative data on site po- tential based on site index measurements for Douglas-fir for several recognized southwest- ern forest habitat types. METHODS Total height and age at diameter breast height were measured for two to six vigor- ously growing dominant or codominant Dou- glas-firs in 450 uneven-aged southwestern spruce-fir (31) or mixed conifer (419) stands from 1979 to 1985. Trees with visible signs of abiotic, insect, or disease damage were not selected as site trees in the stands. The follow- ing information was recorded for each stand: national forest, location (township, range, and section), elevation (nearest 100 feet), aspect (four cardinal directions), slope (nearest 5%), slope position (flat, bottom, ridge, slope) and || USDA Forest Service, Rocky Mountain Forest and Range Experiment Station, Fort Collins, Colorado 80526. ET 278 GREAT BASIN NATURALIST TABLE 1. Southwestern spruce-fir and mixed conifer habitat types sampled. Vol. 46, No. 2 Spruce-fir Habitat Types ABLA/LIBO: ABLA/ EREX: Abies lasiocarpa/Linnaea borealis” (Abies lasiocarpa/Vaccinium scoparium— —Linnaea borealis*) Abies lasiocarpa/Erigeron eximius~ (Abies lasiocarpa/Erigeron superbus’) Mixed Conifer Habitat Types PIEN/SECA: Picea engelmannii/Senecio cardamine® (Picea pungens—Picea engelmannii/Senecio cardamine’*) PIEN/EREX: Picea engelmannii/Erigeron eximius® (Picea pungens—Picea engelmannii/Erigeron superbus’) PIPU/EREX: Picea pungens/Erigeron eximius~”* (Picea pungens—Pseudotsuga menziesii) ABCO/ACGL: Abies concolor/Acer glabrum‘* (Abies Eencolarperuderace menziesulAcer glabrum’) ABCO/FEAR: Abies concolor/Festuca arizonica* (Abies concolor—Pseudotsuga menz siesti/Poa fendleriana =) ABCO/QUGA: Abies concolor/Quercus g porabelit pee (Abies concolor—Pseudotsuga menziesii/Quercus gambelii*) ABCO/BERE: _ Abies concolor/Berberis repens°® (Abies concolor—Pseudotsuga menziesii/[sparse]*) ABCO/EREX: Abies concolor/Erigeron eximius”~” | (Abies concolor—Pseudotsuga menziesii/Erigeron superbus'*) TALEXANDER, B. G., JR., F. RONCO, JR., E. L. FrtzHuGH, AND J. A. Lupwic. 1984. A classification of forest habitat types of the Lincoln National Forest, Nev | | Mexico. USDA Forest Service, Gen. Tech. Rep. RM-104, 29 pp. } *DeVELICE, R. L., J. A. Lupwic, W. H. Morr, AND F. RONCO, Jr. In preparation. A classification of forest habitat types of northern New Mexico and southerr | Colorado. USDA Forest Service, Gen. Tech. Rep. RM. 5Frrznucu, E. L., W. H. Morr, J. A. Lupwic, AND F. Ronco. Forest habitat types in the Apache, Gila, and part of the Cibola national forests, Arizona and Nor ; Mexico. In preparation. ‘Morr, W. H., AND J. A. Lupwic. 1979. A classification of spruce-fir and mixed conifer habitat types of Arizona and New Mexico. USDA Forest Service, Res | Pap. RM-207. 47 pp. The Abies concolor—Pseudotsuga menziesii/Poa fendleriana habitat type originally described by Moir and Ludwig (1979) is now considered to represent | phase of the Abies concolor/Festuca arizonica habitat type described by Moir and Ludwig (1979). Personal communication with W. B. Moir, 1985. YOUNGBLOOD, A. P. In press. Coniferous forest habitat types of central and southern Utah..USDA Forest Service, Gen. Tech. Rep. INT. TABLE 2. Mean Douglas-fir site indexes and standard Forests, Arizona; the Carson (49 stands), Gili deviations by habitat type. (9 stands), Lincoln (50 stands), and Santa Fu (58 stands) National Forests, New Mexico) Number Site Habitat of potential and the San Juan National Forest, Colorad: type stands Mean class (36 stands). | PIEN/SECA 39 91.3+ 9.6 A High Site indexes were determined from averag: ABCO/ACGL 69 89.5+114A High height and age data for each stand using th ABCO/FEAR = 250 87.5 + 8.2 A High Douglas-fir site index curves developed b’ PIPU/EREX 2 ete S 226 =s 103298 Moderate Edminster and J (1976). Mean site inde ABCO/EREX 52 81.2+10.8B Moderate Lee ee Ue EE NO) ee ABCO/QUGA 92 769+ 10.7B Moderate and standard deviation were calculated fo: PIEN/EREX 28 76.2+17.0B Moderate — each habitat type. A one-way analysis of vari) ABCO/BERE 87 14.59 = 956 B Moderate ance, with p 0.10, was used to compar | ALENT a2 12 73.6 + 10.7 BC Moderate ean site indexes among habitat types. Th’ ABLA/LIBO 19 67.3 + 14.4 C Low d l lied tl TOTAL 450 Student-Newman-Kuels test was applie the analysis to show where significant differ / Oneway AOV, p = 0.10, Student-Newman-Kuels. Means followed by ences occurred. different letters are significantly different. habitat type (Moir and Ludwig 1979, Alexan- RESTS | der et al. 1984, Fitzhugh et al. unpublished, | DeVelice et al. unpublished, Youngblood un- published). A total of 10 habitat types were sampled (Table 1). Stands sampled were lo- cated in the Apache (216 stands), Coconino (9 stands), and Kaibab (23 stands) National The 10 habitat types are divided into thre’ site potential classes (high, moderate, an_ low) based on statistically significant differ ences in mean site indexes (Table 2). Th’ PIEN/SECA (mean-91.3), ABCO/ACG| \ April 1986 (89.5), and ABCO/FEAR (87.6) habitat types are classified as high site potential habitat types for Douglas-fir. The ABLA/LIBO (mean-67.7) is classified as the low potential | habitat type. The moderate site potential class _ includes the remaining six habitat types stud- | ied with mean site indexes ranging from 82.6 | (PIPU-EREX) to 73.6 (ABLA/EREX). Mean ; site indexes for the ABLA/EREX and ABLA/ | LIBO habitat types were not significantly dif- | ferent, but they were classified into different . site potential classes because the mean for the -ABLA/LIBO habitat type was below 70 feet. DISCUSSION Site index is currently the most widely used method of evaluating site quality in the United States (Jones 1969, Husch et al. 1972). Several investigators note significant differ- ences in site index between habitat types for «several tree species (Stanek 1966, Roe 1967, Hoffman 1976). However, Daubenmire (1961) rejects the use of ponderosa pine site index curves for predicting potential produc- _ tivity of habitat types in eastern Washington. Our results indicate that the southwestern \spruce-fir and mixed conifer habitat types sampled in this study can be grouped into ‘three significantly different site quality classes ‘for Douglas-fir. Hoffman (1976) also demon- ‘strates significant differences in Douglas-fir site index between three habitat types in cen- tral Idaho. _ In their descriptions of southwestern spruce-fir and mixed conifer habitat types, Moir and Ludwig (1979) give estimates of site _/ quality for individual tree species for several _ aabitat types. Their estimates are based on _ site index and height-age data. Our site index _ data supports Moir and Ludwig's site quality estimates for Douglas-fir in the PIEN/SECA high potential), PIEN/EREX and PIPU/ . 2ZREX (moderate potential) and the ABCO/ 3ERE (usually moderate potential) habitat ypes. Our data also confirm their suggestion hat the ABCO/ACGL habitat type has high Douglas-fir height growth potential. » Moir and Ludwig (1979) interpret heights _ £75-100 feet in 100 years (breast height age) } or the fastest growing Douglas-firs in the j 1BCO/QUGA habitat type to represent poor (| 9 moderate site quality, whereas we inter- \ } ha MATHIASEN ET AL.: DOUGLAS-FIR ECOLOGY 219 pret these data as representing moderate to high site quality for the Southwest. Our site index data for this habitat type basically corre- spond to that of Moir and Ludwig’s, except the range of site indexes is wider in our study. Moir and Ludwig (1979) do not present Douglas-fir site quality estimates for three of the habitat types sampled in this study. Based on our site index data, site quality for Dou- glas-fir is moderate for the ABCO/EREX and ABLA/EREX habitat types and high for the ABCO/FEAR habitat type. Several investigators discuss the difficulties of using site index for estimating site potential in uneven-aged stands (Stage 1963, Jones 1969, Curtis 1976, Daubenmire 1976). Steele et al. (1981, 1983) use site indexes and normal yield tables to estimate productivity potential for habitat types in Montana and Idaho. How- ever, normal yield tables for Douglas-fir in the Southwest are not available, and we do not feel the use of yield tables from other regions would be valid for the Southwest. The devel- opment of separate site curves for different habitat types should improve the accuracy of site index as an estimate of site quality. How- ever, this approach may not solve the prob- lems related to using site index in uneven- aged stands such as early suppression of shade tolerant species (Vincent 1961, Curtis 1976). We agree with the suggestion of Steele et al. (1983) that the development and subsequent validation of growth and yield simulation models using growth coefficients based on habitat types (Stage 1973, 1975) will improve productivity estimates for habitat types. LITERATURE CITED ALEXANDER, B. G., F. Ronco, L. FITZHUGH, AND J. LuD- wic. 1984. A classification of forest habitat types of the Lincoln National Forest, New Mexico. USDA Forest Service, Gen. Tech. Rep. RM-104. 29 pp. Curtis, R. O. 1976. Uneven-aged silviculture and man- agement in the United States. USDA Forest Ser- vice, Proceedings of In-Service Workshops, Mor- gantown, West Virginia, and Redding, California, Timber Management Research, Washington, DAC: DAUBENMIRE, R. 1952. Forest vegetation of northern Idaho and adjacent Washington, and its bearing on the concepts of vegetation classification. Ecol. Monogr. 22: 301-330. 1961. Vegetative indicators of rate of height growth in ponderosa pine. For. Sci. 7: 24—34. _ 1976. The use of vegetation in assessing the pro- ductivity of forest lands. Bot. Review 42: 115-143. 280 DEITSCHMAN, G. H., AND A. W. GREENE. 1965. Relations between western white pine site index and tree height of several associated species. USDA Forest Service, Res. Pap. INT-22. 27 pp. DEVELICE, R. L., J. A. Lupbwic, W. H. Morr, AND F. RONCO, JR. In preparation. A classification of forest habitat types of northern New Mexico and south- ern Colorado. USDA Forest Service, Gen. Tech. Rep. RM. EDMINSTER, C. B., AND L. H. Jump. 1976. Site index curves for Douglas-fir in New Mexico. USDA Forest Ser- vice, Res. Note RM-326. 3 pp. FirzHuGcu, E. L., W. H. Morr, J. A. LuDWwIc, AND F. RONCoO. In preparation. Forest habitat types in the Apache, Gila, and part of the Cibola national forests, Arizona and New Mexico. In preparation. GOTTFRIED, G. J. 1978. Five-year growth and develop- ment in a virgin Arizona mixed conifer stand. USDA Forest Service, Res. Pap. RM-203. 22 pp. HANKS, J. P., L. FirzHuGu, AND S. R. HANKS. 1983. A habi- tat type classification system for ponderosa pine forests of northern Arizona. USDA Forest Ser- vice, Gen. Tech. Rep. RM-97. 22 pp. HoFrMaN, L. J. 1976. Height growth of Douglas-fir in relation to habitat types in northern Idaho. Un- published thesis, University of Idaho, Moscow, Idaho. 30 pp. Huscu, B., C. MILLER, AND T. W. BEERS. 1972. Forest mensuration. Ronald Press Company, New York. 410 pp. JONES, J. R. 1969. Review and comparison of site evalua- tion methods. USDA Forest Service, Res. Pap. RM-75. 19 pp. _____. 1974. Silviculture of southwestern mixed conifer and aspen: the status of our knowledge. USDA Forest Service, Res. Pap. RM-122. 44 pp. LaysER, E. F. 1974. Vegetation classification: its applica- tion to forestry in the northern Rocky Mountains. J. For. 72: 354-357. Morr, W., AND J. Lupwic. 1979. Classification of spruce- fir and mixed conifer habitat types of Arizona and New Mexico. USDA Forest Service, Res. Pap. RM-207. 47 pp. GREAT BASIN NATURALIST Vol. 46, No. 2 PFISTER, R. D. 1976. Land capability assessment by habi- tat types. Pages 312-335 in America’s renewable resource potential—1975: the turning point. Pro- ceedings of the 1975 National Convention Society of American Foresters, Washington, D.C. PFISTER, R. D., ANDS. ARNO. 1980. Classifying forest habi- tat types based on potential climax vegetation. For. Sci. 26: 52-70. PFISTER, R. D., B. KOVALCHIK, S. ARNO, AND R. PRESBY. 1977. Forest habitat types of Montana. USDA Forest Service, Gen. Tech. Rep. INT-34. 174 pp. PFISTER, R. D., J. SCHMAUTZ, D. ON, AND C. BRown. 1971. Management implication by habitat types. USDA Forest Service, Northern Region Training Man- ual, Mimeograph. 30 pp. Rok, A. L. 1967. Productivity indicators in western larch forests. USDA Forest Service, Res. Note INT-59. 4 pp. STAGE, A. R. 1963. A mathematical approach to the poly- morphic site index curves for grand fir. For Sci. 9: 167-180. _____. 1973. Prognosis model for stand development. USDA Forest Service, Res. Pap. INT-137. 32 pp. _____. 1975. Prediction of height increments for models for forest growth. USDA Forest Service, Res. Pap. INT-164. 20 pp. STANEK, W. 1966. Relative quality of the major forest association of the southern British Columbia inte- rior for growth of lodgepole pine, Engelmann spruce, Douglas-fir, and alpine fir. For. Chron. _ 42: 306-313. STEELE, R., S. V. Cooper, D. M. ONDov, D. W. ROBERTS, AND R. D. PFISTER. 1983. Forest habitat types of eastern Idaho—western Wyoming. USDA Forest _ Service, Gen. Tech. Rep. INT-144. 122 pp. STEELE, R., R. D. PFISTER, R. A. RYKER, AND J. A. KITTAMS. 1981. Forest habitat types of central Idaho. USDA Forest Service, Gen. Tech. Rep. INT-114. 138 pp. | VINCENT, A. B. 1961. Is height/age a reliable index of site? For. Chron. 37: 144-150. YOUNGBLOOD, A. P. In press. Coniferous forest habitat types of central and southern Utah. USDA Forest Service, Gen. Tech. Rep. INT. WINTERING MULE DEER PREFERENCE FOR 21 ACCESSIONS OF BIG SAGEBRUSH Bruce L. Welch and E. Durant McArthur! ABSTRACT. —Wintering mule deer showed differential browsing preference among 21 accessions of big sagebrush (Artemisia tridentata) grown on gardens on three different mule deer herd ranges. The Hobble Creek accession of big sagebrush was significantly preferred over the other 20 accessions across all three sites and for all three years. Accessional preference means for the study period for all sites combined ranged from 28.3 to 57.5%. The data collected support the planned release of the Hobble Creek accession as a superior cultivar of big sagebrush for use on mule deer winter ranges. Plant coumarin content was primarily under genetic control, but site factors also had an effect. Assay for coumarin compounds is useful in determining subspecies of A. tridentata but not for precise prediction of mule deer browsing preference. During in vivo digestion trials Smith (1950) observed that mule deer showed definite aversion to individual big sagebrush (Artemisia tridentata) plants. Since then sev- eral field workers have observed differential preference of wintering mule deer not only for individual plants but also for populations of big sagebrush (Plummer et al. 1968, Winward 1970, Brunner 1972, Hanks et al. 1973, Stevens and McArthur 1974, Winward and Tisdale 1977, McArthur et al. 1979, Willms et al. 1979). We reported (Welch et al. 1981) that wintering mule deer showed differential pref- erence for 10 accessions of big sagebrush grown on a uniform garden. This study is an effort to include 11 more accessions (21 total), different sites, different mule deer herds, and the role of coumarin compounds in the prefer- ence of wintering mule deer for accessions of big sagebrush. Coumarin compounds have been shown to be positively correlated with mule deer preference of some sagebrush taxa (Stevens and McArthur 1974). MATERIALS AND METHODS Three outplanting sites were selected within the elevational range of most Utah mule deer winter ranges (1,370—2,140 m): Springville, Utah; 3 km east of Nephi, Utah (Salt Creek Canyon); and 15 km west/south- west of Helper, Utah (Gordon Creek Wildlife _. Management Area). } The Springville site at 1,402 m is a basin big sagebrush habitat type. Soil is of the Pleasant Grove series (Pleasant Grove gravelly loam, 6-10% slope), consisting of deep, well- drained, gravelly or cobbley soils, alluvial fans. Weathered limestone is the parent ma- terial. Soil permeability is moderately rapid. Roots can penetrate to a depth of 1.5 m or more. About 9 cm of available water is held by this soil to a depth of 1.5 m. Soil pH ranges from 7.4 to 7.9. Average annual precipitation is 35-45 cm. The mean annual temperature is 8.9-11.1 C, and the frost-free period is 150-170 days (Swenson et al. 1972). The Salt Creek Canyon site (1,676 m) is a basin big sagebrush habitat type. Soil is a Rofiss gravelly clay loam 4—15% slope. It is a deep, well-drained alluvial soil. Parent mate- rial is Arpien shale. Soil permeability is mod- erately slow. Effective rooting depth is 1.5 m or more. Soil pH ranges from 8.2 to 8.6. Aver- age annual precipitation is 20-35 cm. The mean annual temperature is 7.2-8.3 C, and the frost-free period is 100-120 days (Soil Conservation Service 1980). The Gordon Creek Wildlife Management Area, at 2,130 m, is a Wyoming big sagebrush habitat type. Soil is of the Atrac series (Atrac is a very fine sandy loam 1-6% slopes). This series consists of deep, well-drained soils. Parent material is sandstone. Effective root- ing depth is 1.5 m or more. Soil pH ranges Mntermountain Research Station, Shrub Sciences Laboratory, 735 North 500 East, Provo, Utah 84601. The use of trade, firm, or corporation names in this paper is for the information and convenience of the reader. Such use does not constitute an officia 1 endorsement or approval by the U.S. Department of Agriculture of any product or service to the exclusion of others that may be suitable. 281 282 GREAT BASIN NATURALIST Vol. 46, No. 2 TABLE 1. Locations of collection sites for 21 accessions of big sagebrush (Artemisia tridentata) used in this study to determine wintering mule deer preference. Subspecies Accession vaseyana Colton Sardine Canyon Benmore Petty Bishop’s Log Durkee Springs Salina Canyon Clear Creek Canyon Pinto Canyon Indian Peaks Hobble Creek tridentata Clear Creek Canyon Brush Creek Loa Dove Creek Evanston Wingate Mesa Dog Valley wyomingensis Evanston Kaibab Trough Springs Milford from 6.6 to 8.0. Average annual precipitation is 30.5-35.6 cm. The mean annual tempera- ture is 7.2-8.3 C, and the frost-free period is 100-120 days (Soil Conservation Service 1981). At each site the vegetation was killed by disking and kept weed free by hand and me- chanical means. Deer-proof fences were built around each site. We planted each site in spring 1978 with containerized stock of 21 accessions of big sagebrush (Table 1). Each accession was represented by 10 plants, and each plant was an experimental unit within the accessions. The resulting 210 plants in each set were placed at random on a3 x 3m grid for each site. Table 1 lists the locations of seed collection sites for the accessions. We measured the preference for the acces- sions by randomly selecting 15 annual leaders per plant through the entire crown and meas- uring them to the nearest centimeter in November 1980, 1981, and 1982. These meas- urements were used to calculate a mean an- nual leader length per plant before browsing. We remeasured plants using 15 randomly se- lected leaders during April 1981, 1982, and 1983 and calculated a mean annual leader length after browsing. Percentage used or preference was calculated by dividing the mean leader length after browsing by the County and state Elevation (m) Utah, Utah 2,260 Cache, Utah 1,800 Tooele, Utah 1,900 Sanpete, Utah 2,380 Sevier, Utah 2,270 Sevier, Utah 2,350 Sevier, Utah 2,130 Washington, Utah 1,850 Beaver, Utah 2,140 Utah, Utah 1,500 Sevier, Utah 1,720 | Uintah, Utah 1,830 ) Wayne, Utah 2,140 Dolores, Colorado 2,070 Uinta, Wyoming 2,020 San Juan, Utah 2,060 Juab, Utah 1,700 Uinta, Wyoming 2,130 Coconino, Arizona 2,340 Humboldt, Nevada 1,400 Beaver, Utah 1,540 mean leader length before browsing, multi- plying by 100, and subtracting from 100. Welch et al. (1981) reported a precision error — of 4% for this method. Data were expressed as percentage used (preference). Ultraviolet visable, fluorescent, water-sol- uble coumarin compounds (principally isosco- | poletin, scopoletin, and esculetin) were mea- © sured by placing leaf fragments from single | leaves (5 samples/plant) ofapproximately 1 mg | size in water in cuvettes (McArthur et al. | 1981). We used 1 mg leaf material/0.3 ml wa- ter up to 10 mg leaf material/sample. After 90 minutes equilibration time, spectrophoto- metric readings (percent transmittance) were | made at 364 nm in a Bausch and Lomb Spec- | tronic 700 spectrophotometer. Leaves for coumarin analysis were collected at the time ~ of the 1983 preference measurements. i The three subspecies of big sagebrush, A. f. _ ssp. tridentata, vaseyana, and wyomingensis, ‘ were represented in this study. Of the 21 accessions, 10 were vaseyana, 7 were triden- | tata, and 4 were wyomingensis . This gave the - capability of testing for subspecies effects on — preference. Analyses were performed using the SPSS* | statistical package (SPSS Inc., 1983). We used _ f analysis of variance techniques, Student- | ; Newman-Keuls multiple range mean com- | I ) ) April 1986 WELCH, MCARTHUR: MULE DEER 283 TABLE 2. Effects of subspecies, site, and year on preference of wintering mule deer for accessions of big sagebrush (Artemisia tridentata). Data expressed as percent of current year’s growth eaten (3 years, 630 plants, three gardens) Preference for subspecies A. t. ssp. tridentata A. t. ssp. wyomingensis A. t. ssp. vaseyana 32.6° Soin MATE Effects due to site Salt Creek Gordon Creek Springville 30.4" 35.0? 50.2° Effects due to years 1980 | 1981 1982 31.6* 41.7” 42.4? *Means sharing the same superscript are not significantly different at the 5% level. TABLE3. Preference of wintering mule deer for accessions of big sagebrush (Artemisia tridentata ) grown on different gardens. Data expressed as a percent of current year’s growth eaten. Data points for gardens represent a three-year mean. Accession Gordon Creek Evanston (t)** 2 e2 Trough Springs (w) 17.5 Dove Creek (t) 26.3 Clear Creek Canyon (t) 31.5 Loa (t) 25.0 Kaibab (w) 30.9 Dog Valley (t) 28.6 Brush Creek (t) 35.2 Wingate Mesa (t) 32.5 Milford (w) 34.9 Clear Creek Canyon (v) 34.4 Benmore (v) 40.1 Pinto Canyon (v) 36.8 Evanston (w) 31.1 Durkee Springs (v) 36.3 Salina Canyon (v) 40.5 Sardine Canyon (v) 45.8 Indian Peaks (v) 40.9 Petty Bishop's Log (v) 46.4 Colton (v) 43.1 Hobble Creek (v) 52.0 Garden Salt Creek Springville Mean 18.1 41.6 OStaas 30.9 41.9 Sales 31.3 Boal 30.9%" 19.7 42.7 oieeeee 18.5 51.3 S1aGme 27.2 42.4 Be) yee 23.6 49.1 Bougebede 24.6 47.2 Bo aod 33.9 42.8 Bou arbedel 29.0 47.4 Siraliecacke 25.6 55 38. 4bedete 39.3 38.4 39, grcdete 27.8 56.2 40. 3°3 43.8 46.3 40, 4% 27.7 58.1 40.73% 29.4 55.2 4).7°* 29.9 58.0 44.6% 40.9 55.0 45.6% 33.8 57.7 46.08 37.9 60.1 47.08 47.4 73.0 57.5" *Means sharing the same superscript are not significantly different at the 5 % level. **t =A. t. ssp. tridentata; y = A. t. ssp. vaseyana; w = A. t. ssp. wyomingensis. parison tests, and regression techniques. Data collected as percent were transformed by the arcsin percentage function for analysis and then returned to the percent values for final presentation (Snedecor and Cochran 1956). RESULTS AND DISCUSSION The overall mean percent of leader length used for all years, sites, and accessions was 38.6%. Two three-way analyses of variance detected significant effects due to all main effects of year, site, subspecies, and accession (Table 2). Wintering mule deer significantly pre- ferred the subspecies A. t. ssp. vaseyana over A. t. ssp. wyomingensis and A. t. ssp. triden- tata (Table 2). They significantly preferred A. t. ssp. wyomingensis over A. t. ssp. triden- tata. These results agree with reports of pre- vious studies (Stevens and McArthur 1974, Welch et al. 1981). Significantly more big sagebrush was con- sumed by wintering mule deer at the Springville site than at Salt Creek and Gordon Creek (Table 2). Big sagebrush consumption was significantly more at Gordon Creek than at Salt Creek. This differential use among the sites strengthens our ability to rank the pref- 284 GREAT BASIN NATURALIST Vol. 46, No. 2 TABLE 4. Analysis of variance of amount of coumarin by subspecies and effects due to site. Data are presented as percent transmittance (364 mm)*. Amount of coumarin by subspecies* A. t. ssp. tridentata A. t. ssp. wyomingensis A. t. ssp. vaseyana 61.9°" 54.7” Sela Effects due to site** Salt Creek Gordon Creek Springville 43.6" 48.6 57.9° *Percent transmittance is an inverse function of coumarin content. **Means sharing the same superscript are not significantly different at the 5% level. *** Springville site differences were compounded by disproportionate mortality of ssp. vaseyana prior to 1983 at that site. TABLE 5. Linear regression comparisons of mean preference and percent transmittance values. * Number of Comparison pairs r Significance All accessions 21 — 0.58 01 Subspecies 3 — 0.998 .05 A. t. ssp. tridentata 7 = (008) ca. .05 A. t. ssp. wyomingensis 4 + 0.82 NS A. t. ssp. vaseyana 11 + 0.01 NS *Values computed from Tables 2 and 4 using mean values only (Snedecor and Cochran 1956). Percent transmittance is an inverse function of coumarin content. erence of mule deer for the accessions of big sagebrush. Also, different mule deer herds occupied the three sites, thus eliminating any confounding resulting from previous expo- sure to a given type of big sagebrush. Significantly more big sagebrush was con- sumed during 1981 and 1982 than in 1980 (Table 2). We have no explanation for these differences. Wintering mule deer significantly pre- ferred the Hobble Creek accession over the other 20 accessions across all sites and years (Table 3). Overall use of the Hobble Creek accession was 57.5%. Use of Hobble Creek big sagebrush varied significantly between sites. At the Salt Creek site, the Hobble Creek accession received 47.4% use, 52.0% use at Gordon Creek, and 73.0% use at Springville. The consistency of ranking for preference of the Hobble Creek accessions was not present for the other accessions and further strengthens our contention that the Hobble Creek is the most preferred. This con- sistency was also true for years. These study results agree with our earlier study (Welch et al. 1981). We are currently collecting data needed to support the release of Hobble Creek through the Soil Conservation Service plant development program as a superior cul- tivar of big sagebrush for use on mule deer winter ranges. Different levels of coumarin compounds were found to be present in each subspecies (Tables 4 and 5). Earlier studies have shown that ssp. vaseyana had higher coumarin levels than ssp. tridentata and wyomingensis (Ste- vens and McArthur 1974, McArthur et al. 1981). Ina previous report, we suggested that these coumarin compounds would be useful for indirect selection for preference (Welch and McArthur 1979). This study shows that there is an environmental component to cou- marin content (Table 4). The effects due to site are significant. Nevertheless, genetic factors are of overriding importance. Subspecies were significantly different in coumarin con- tent (Tables 4 and 5), and individual acces- sions ranked rather consistently in coumarin content across sites (Table 6). Coumarin con- tent data from a set of plants over three years and all seasons showed that, whereas cou- | marin levels may vary temporally, individual _ plants maintain general positions in respect to other plants in the population (McArthur, © Welch, and Noller, data on file, Shrub Sci- | ences Laboratory). . | Coumarin compounds are useful indicators of A. tridentata subspecies—especially in dis- tinguishing ssp. vaseyana from ssp. tridentata and wyomingensis (Stevens and McArthur 1974, McArthur et al. 1981, Table 4). Their use as preference indicators for browsing April 1986 WELCH, MCARTHUR: MULE DEER 285 TABLE 6. Percent transmittance at 364 nm by accession and site. Mean preference* is also given. Percent transmittance Mean Gordon Salt Accession preference Creek Creek Springville Mean Dove Creek (t)** 30.9 63.0 62.9 72.9 aie" Clear Creek Canyon (t) SIRS 66.9 57.2 68.6 63.9" Evanston (t) 28.3 57.0 63.8 69.0 63.4°° Dog Valley (t) 33.8 67.7 58.4 63.7 63.3% Loa (t) 31.6 63.4 58.0 67.5 62.87 Evanston (w) 40.4 67.2 56.9 52.6 5OiSag Wingate Mesa (t) 36.4 58.2 53.9 64.5 58.97 Brush Creek (t) 35.7 64.3 49.3 49.0 54.6bc4 Milford (w) Salk 57.5 46.8 Sie 5Seiiacs Pinto Canyon (v) 40.3 49.4 49.4 63.2 Selec Trough Springs (w) 30.1 57.7 50.0 51.5 OOF Kaibab (w) 33.5 49.1 51.9 53.5 5Ol6rs Hobble Creek (v) 57.5 49.5 45.3 56.5 49.5% Colton (v) 47.0 47.3 40.8 48.2 45.5° Indian Peaks (v) 45.6 46.1 41.4 Soul 34.24 Durkee Springs (v) 40.7 44.2 Bal 57.4 45.0° Clear Creek Canyon (v) 38.4 29.1 26.0 53.0 Soule Sardine Canyon (v) 44.6 23.4 22.6 55.0 31.6° Salina Canyon (v) 41.7 aed 28.4 48.2 ROS Benmore (v) 39.3 26.0 16.8 43.5 Qs Petty Bishop's Log (v) 46.0 16.4 13.4 eae 14.8! *From Table 3. **t = A. t. ssp. tridentata; v = A. t. ssp. vaseyana; w = A. t. ssp. wyomingensis. ***Means sharing the same superscript are not significantly different at the 5% level. ****No surviving plants in 1983. mule deer is examined in Tables 5 and 6. We concluded that these compounds are good general indicators in the sense that the sub- species are browsed differentially by mule deer (Table 2), and those differences are re- flected by differences in coumarin content (Table 4). The relationship is ssp. vaseyana > ssp. wyomingensis > ssp. tridentata in brows- ing preference and in coumarin content. However, the relationship of coumarin con- tent to browsing preference is not strong nor consistent within subspecies (Tables 5 and 6). A regression analysis (SPSS*) interacting all (n=573) coumarin and preference data points gave an unimpressive but highly significant (p < .001) r° of 0.06. Linear regression values were strikingly different in magnitude and sign for the three subspecies (Table 5). Fur- thermore, an examination of the data for the most preferred accession, Hobble Creek (ssp. vaseyana), shows that it has a relatively low coumarin content for its subspecies (Table 6). Coumarin content is useful in identifying pre- ferred taxa but not in making distinctions within those taxa. The chromosome number level may impact ' coumarin content. Most ssp. tridentata and vaseyana accessions are diploid. The wyoming- ensis accessions are tetraploid (McArthur et al. 1981, McArthur and Welch 1982). Both tetraploid spp. tridentata accessions (Wingate Mesa and Brush Creek) are relatively high in coumarin content (Table 6). Two of the ssp. vaseyana accessions with relatively low cou- marin content (Colton and Pinto Canyon) are tetraploid. The additional genomes may be responsible for a moderating effect in the case of tetraploids for both ssp. tridentata and vaseyana. Tetraploid plants of both subspe- cies move toward intermediate values from the more extreme values of their respective diploids. ACKNOWLEDGMENTS The authors acknowledge the excellent technical assistance of Tracy Jacobson. They also acknowledge cooperative arrangements with the Utah Division of Wildlife Resources in obtaining plant materials and maintenance of the Springville and Gordon Creek sites (Central Regional Office and the Pittman- Robertson Wildlife Restoration Funds, Pro- ject W-82-R). LITERATURE CITED BRUNNER, J. R. 1972. Observations on Artemisia in Ne- vada. J. Range Manage. 25: 205-208. Hanks, D. L., E. D. MCARTHUR, R. STEVENS, AND A. P. PLUMMER. 1973. Chromatographic characteristics and phylogenetic relationships Artemisia section Tridentatae. USDA For. Serv. Intermt. For. and Range Expt. Stn. Res. Pap. INT-141. 24 pp. MCARTHUR, E. D., A. C. BLAUER, A. P. PLUMMER, AND R. STEVENS. 1979. Characteristics and hybridization of important intermountain shrubs. HI. Sunflower family. USDA Forest Serv. Res. Pap. INT-220. 82 pp. MCARTHUR, E. D., C. L. POPE, AND D. C. FREEMAN. 1981. Chromosomal studies of subgenus Tridentatae of Artemisia: evidence for autopolyploidy. Amer. J. Bot. 68: 589-605. MCARTHUR, E. D., AND B. L. WELCH. 1982. Growth rate differences among big sagebrush (Artemisia tri- dentata) accessions and subspecies. J. Range Manage. 35: 396-401. PLUMMER, A. P., D. R. CHRISTENSEN, AND S. B. MONSEN. 1968. Restoring big game range in Utah. Utah Div. Wildl. Resour. Publ. 68-3. 183 pp. SmiTH, A. D. 1950. Sagebrush as winter feed for mule deer. J. Wildlife Manage. 14: 285-289. SNEDECOR, G. W., AND W. C. Cocuran. 1956. Statistical methods applied to experiments in agriculture and biology (5th ed.). Iowa State College Press, Ames. 534 pp. SOIL CONSERVATION SERVICE. 1980. Unpublished data on file, Fairfield-Nephi Soil Survey Area, Utah. Sheet 61. Provo, UT: USDA, Soil Conservation Service. GREAT BASIN NATURALIST Vol. 46, No. 2 ____. 1981. Unpublished data on file, Gordon Creek Soil Survey Area, Utah. Price, UT: USDA, Soil Conservation Service. SPSS Inc. 1983. A complete guide to SPSS* language and operations. McGraw-Hill Book Co., New York. 806 pp. STEVENS, R., AND E. D. McArTHUR. 1974. A simple field technique for identification of some sagebrush taxa. J. Range Manage. 27: 325-326. SWENSON, J. L., Jk., W. M. ARCHER, K. M. DONALDSON, J. J. SHIOZAK, B. BRODERICK, AND L. WooDWaRD. 1972. Soil Survey of Utah County, Utah—central part. Provo, UT: USDA, Soil Conservation Service. WELCH, B. L., AND E. D. MCARTHUR. 1979. Feasibility of improving big sagebrush (Artemisia tridentata) for use on mule deer winter ranges. Pages 451— 473 in J. R. Goodin and D. K. Northington, eds., Arid land plant resources. Texas Tech University, Lubbock. 724 pp. WELCH, B. L., E. D. MCARTHUR, AND J. N. Davis. 1981. Differential preference of wintering mule deer for accessions of big sagebrush and for black sage- brush. J. Range Manage. 34: 409-411. WILLMS, W., A. MCLEAN, R. TUCKER, AND R. RITCEY. 1979. Interactions between mule deer and cattle on big sagebrush range in British Columbia. J. Range Manage. 32: 299-304. WINWARD, A. H. 1970. Taxonomic and ecological relation- ships of the big sagebrush complex in Idaho. Un- published dissertation, University of Idaho, ‘Moscow. 80 pp. WinwarbD, A. H., AND E. W. TISDALE. 1977. Taxonomy of the Artemisia tridentata complex in Idaho. Coll. Forest, Wildlife and Range Sci., University of Idaho, Moscow. Bull. 19. 15 pp. i COLEOPTERA OF THE IDAHO NATIONAL ENGINEERING LABORATORY: AN ANNOTATED CHECKLIST Michael P. Stafford’, William F. Barr” and James B. Johnson? ABSTRACT. —An insect survey was conducted on the Idaho National Engineering Laboratory during the summers of 1981-1983. This site is on the Snake River Plains in southeastern Idaho. Presented here is an annotated checklist of the Coleoptera collected. Successful collecting methods, dates of adult occurrence, and relative abundance are given for each species. Relevant biological information is also presented for some species. The Idaho National Engineering Labora- tory (INEL) was designated an Environmen- tal Research Park in 1975. Because of the increased number of ecological studies on the INEL since this time, there is a need for an entomofaunal list for this area. This informa- tion will be valuable to many biologists since insects interact with many other organisms, e.g., plants as pollinators and herbivores or vertebrates as food and parasites. Therefore, an insect survey was conducted on the INEL during the summers of 1981-1983. The objec- tives of this survey were to document the taxonomic composition of the insect fauna and the seasonal occurrence, relative abundance, and host information for the insect species present. Several arthropod surveys were conducted on the INEL, formerly known as the National Reactor Testing Station, under the auspices of Brigham Young University. Allred (1968) re- ported seven species of ticks from a variety of hosts. Forty-two species of spiders were col- lected in a pitfall trap survey (Allred 1969). Allred and Cole (1971) published a checklist of 22 species of ants, and Allred and Muma (1971) listed the solpugid species found on the INEL. Allred (1973) reported the occurrence of scorpions. Although these studies provide valuable information on the ant and arachnid faunae, information on other insect: taxa is lacking. STUDY AREA The INEL is on the Upper Snake River Plains at the base of the Lost River and Lemhi a on Fig. 1. Location and detail of the Idaho National Engi- neering Laboratory. Mountain Ranges in southeast Idaho (Fig. 1). The topography is flat to gently rolling; mean elevation is 1,490 m. Soils are generally aeo- lian sandy loams and loess and classified as Aridisols (McBride et al. 1978), with scattered lava outcroppings. The vegetation is domi- nated by big sagebrush (Artemisia tridentata Nutt.) Other conspicuous shrubs include green rabbitbrush (Chrysothamnus viscidi- floris [Hook.] Nutt.), gray rabbitbrush (C. nauseosus [Pall.] Britt.), gray horsebrush (Te- tradymia canescens D.C.), and winter fat ‘Published with the approval of the director of the Idaho Agriculture Station as Research Paper 85724. *Department of Plant, Soil and Entomological Sciences, University of Idaho, Moscow, Idaho 83843. 287 288 (Ceratoides lanata |Pursh] J. T. Howell). The dominant grasses are bluebunch wheatgrass (Agropyron spicatum [Pursh] S. & S.), Indian ricegrass (Oryzopsis hymenoides [R. & S.] Ricker), bottlebrush squirreltail (Sitanion hystrix [Nutt.] J. B. Smith), and needle-and- thread (Stipa comata Trin. & Rupr.). The more alkaline areas are dominated by shad- scale (Atriplex confertifola |Torr. & Frem. |] Wats.) and saltsage (A. nuttalii Wats.). In de- pressions prone to flooding are stands of Great Basin wildrye (Elymus cinereus Scribn. & Merrill). Extensive stands of Utah juniper (Juniperus osteosperma [Torr.] Little) occur on the foothills of the surrounding mountains. Harniss and West (1973) described the vege- tation patterns and presented a detailed map of the vegetation found on the INEL. Jeppson and Holte (1978) listed the plant species found in the area. RESULTS Presented here is an annotated checklist of the Coleoptera we collected in approximately 800 man-hours of collecting during this survey and previous collecting efforts on the INEL over the last 25 years by one of us (WFB). The list includes 214 species in 29 families, making it the most extensive faunal list yet published for the INEL. This information should be use- ful for those undertaking entomological and ecological studies in semiarid regions of the West, particularly in the Great Basin and Snake River Plains. The placement of family names follows A Catalog of the Coleoptera of America North of Mexico (Kingsolver 1979). Genus and species names are arranged in al- phabetical order. Determinations, except as noted in the acknowledgments, were made by the authors and Paul J. Johnson (also at the University of Idaho) using published keys, e,g. Arnett (1963) and Hatch (1953-1973), and the University of Idaho insect collection as reference material. Following each species are the collecting methods, dates of occur- rence, and abundance. Biological information is presented for some species. Voucher speci- mens are housed in the William F. Barr Ento- mological Museum at the University of Idaho. Collecting Methods BF = Berlese Funnel; GC = General Collect- ing; LT = Light Trap; MT = Malaise Trap; GREAT BASIN NATURALIST Vol. 46, No. 2 SW = Sweep Net; PE = Plant Examination; PF = Pitfall Trap; WP = Windowpane Trap. Relative Abundances U = Uncommon, 1—15 specimens; C = Com- mon, 16-50 specimens; A = Abundant, 51+ specimens. The assignment of the abundance rating is subjective. It is based upon both the number of specimens collected and on personal obser- vations. Because of population fluctuations we have observed over the years, these ratings may not always be indicative of the abundance of a species in that particular year. Also, the collecting methods employed may not always reflect the relative abundance of a species. COLEOPTERA CHECKLIST Carabidae Agonum balesi Gray: PF; May, Jun; U. Agonum placidum (Say): LT; Jul; U. Amara apricaria Paykull: LT; Jul; U. Amara impuncticollis Sav: PF; Jul; U. Amara laticollis LeConte: LT; Jul; U. Amara musculus Say: PF; Jun, Jul; U. Amara sp. (poss. quenseli Schonherr): PF; Jul; U Axinopalpus biplagiatus (Dejean): PF, WP; Jul; U. Bembidion immaculosum Hatch: LT, PF; Jul, Aug; U. Bembidion nebraskense LeConte: LT; Aug; WF Bembidion obscurellum Motschulsky: LT; Jul, Aug; A. Bembidion rupicola Kirby: LT; Aug; U. Bembidion timidum LeConte: LT; Aug; U. Calosoma luxatum Say: PF; May—Jul; U. Cicindela decemnotata Say: PF, GC; May— Aug; U. Cymindis planipennis LeConte: PF; Jun— Aug; U. Harpalus amputatus Say: PF; May; U. Harpalus basilaris Kirby: PF; Jun; U. Harpalus fraternus LeConte: LT, PF; Jun, Jul; U. Lebia vittata (Fabricius): WP; May—Jul, U. Microlestes nigrinus (Mannerheim): PF; Jun, Jul; U. Philophuga viridis Dejean: PF, PE; May—Jul; U. Larva observed feeding on larva of the sagebrush defoliator, Aroga websteri Clark, (Lepidoptera: Gelechidae), on A. tridentata. | { | April 1986 Piosoma setosa LeConte: PF, WP; Jun, Jul; U. New state record. Pseudomorpha behrensi Horn: PF; Jul, U. Dytiscidae Laccophilus decipiens LeConte: LT; Aug; U. Hydrophilidae Berosus fraternus LeConte: LT; Jul; U. Berosus styliferus Horn: LT; Jul, Aug; U. Cercyon quisquilius (Linnaeus): LT; Jul; U. Helophorus sp.: LT; Jul, Aug; U. Sphaeridium scarabaeoides (Linnaeus): PF: Jun—Aug; U. Histeridae Psiloscelis corrosa Casey: WP; Jun; U. New state record. Saprinus lugens Erichson: GC; May; U. Saprinus oregonensis LeConte: PF; Jun, Jul; We Xerosaprinus acilinea (Marseul): PF; Jun, Jul; WE Xerosaprinus lubricus (LeConte): PF; Jun— Aug; C. Scarabaeidae Aphodius denticulatus Haldeman: LT, PF; May-—Jul; U. Aphodius distinctus Miiller: PF; May; U. Aphodius fossor (Linnaeus): GC; Jul; U. Aphodius hirsutus Brown: PF; May; U. Aphodius granarius (Linnaeus): PF; Jun; U. Aphodius vittatus Say: PF; Jul; U. _ Aphodius sp. (poss. militaris LeConte): PF, WP; May; U. Boreocanthon simplex (LeConte): PF; May; (We Cremastocheilus crinitus bifoviatus Van Dyke: PF, GC; Jun, Jul; U. Dichelonyx truncata LeConte: GC; Jun, Jul; Ge Diplotaxis haydenii LeConte: GC, LT, WP; Jun, Jul; U. _ Diplotaxis obscura LeConte: PF; May—Aug; C Diplotaxis subangulata LeConte: LT, WP; | Jul; U. \| Glaresis canadensis Brown: PF: Jun—Aug; C. | Ochodaeus simplex LeConte: LT, WP; Jul, U. Paracotalpa granicollis (Haldeman): GC; Jul: U Serica anthracina LeConte: GC, PF; Jun; U. STAFFORD ET AL.: COLEOPTERA OF IDAHO 289 Serica barri Dawson: LT, PF: Jul; A. Trox sp.: LT; Jun; U. Leiodidae Hydnobius sp.: LT, PF; Jun, Jul; U. Ptomophagus californicus (LeConte): PF; Jul; lO: Silphidae Nicrophorus hecate Bland; PF, WP: May- Aug; C. Staphylinidae Anotylus sp.: LT; Aug; U. Bledius strenuus Casey: LT: Jul; U. Philonthus concinnus (Gravenhorst): PF; May; We Philonthus cruentatus (Gmelin): PF; May; U. Platystethus americanus (Erichson): LT; Aug: Wl, Tachinus angustatus Horn: WP; Jun; U. Pselaphidae Pilopius sp.: GC; May; U. Heteroceridae Lanternarius brunneus (Melsheimer): LT; |juras Buprestidae Acmaeodera immaculata Horn: GC, PE; Jul; C. Larvae bore roots of C. lanata, adults on flowers of Opuntia polyacantha Haw. Agrilus politus (Say): GC, PE; Jul-Aug; U. Larvae bore Salix spp. Agrilus pubifrons Fisher: GC, PE, SW; Jul, Aug; C. Larvae bore roots and adults feed in the flowers of C. viscidifloris. Agrilus walsinghami Crotch: GC, PE; Aug; U. Larvae bore roots of C. nauseosus. Anthaxia retifer LeConte: GC, PE; May; U. Adults on flowers of Balsamorhiza sagittata (Pursh) Nutt. Chrysobothris deleta LeConte: GC, PE; Aug; U. Larvae bore roots of A. tridentata. Chrysobothris horningi Barr: GC, PE; May; U. Larvae bore roots of Eriogonum spp. Chrysobothris idahoensis Barr: GC, PE; Jun— Aug; U. Larvae bore roots of Eriogonum spp. Chrysobothris texana WLeConte: GC, PE; Jun—Aug; C. Larvae bore branches of J. 0s- teosperma. Nannularia brunneata (Knull): GC, PE; Jul; U. Larvae bore roots of Eriogonum sp. Elateridae Ampedus ursinus (Van Dyke): PF; Jun; U. Anchastus cinereipennis (Ecshscholtz): PF; May-—Jul; C. Adults associated with E. cinereus. Cardiophorus sp. 1: BF, PF; Mar, May, Jun; We Cardiophorus sp. 2: PF; May, Jun; U. Cardiophorus sp. 3: GC, PF; May, Jun; U. Cardiophorus sp. 4: WP; Jun; U. Cardiophorus sp. 5: BF, PF, WP; Mar, May; WE Ctenicera noxia (Hyslop): PF; May, Jun; U. Ctenicera semivittata (Say): GC; May; U. Hypolithus bicolor (Eschscholtz): PF; Jul; U. Cantharidae Malthodes sp.: GC; May, U. Dermestidae Dermestes marmoratus Say: GC, LT; Jul, Aug; C. Cleridae Enoclerus acerbus Wolcott: GC, PF, WP; May-—Jul; C. Enoclerus barri Knull: GC; Jul, Aug; U. Phyllobaenus sp.: GC, PE, SW, WP; Jun-— Aug; C. Adults often found on A. tridentata preying on small Lepidoptera larvae. Trichodes ornatus Say: WP; Jul; U. Melyridae Amecoceris spp.: BF, PF, WP; May—Jul; A. Attalus glabrellus Fall: GC, WP; May—Jul; C. Attalus morulus smithi Hopping: GC, SW; May, Jun; U. Adults associated with E. cinereus. Attalus oregonensis Horn: GC, PE, PF; Jun, Jul; C. Adults feed on pollen in flowers of O. polyacantha. Collops bipunctatus (Say): PE, SW; Jul; U. Adults feed on pollen of E. cinereus. Collops bridgeri Tanner: GC, PF; May—Jul; U. Collops hirtellus LeConte: SW; Jun; U. Collops punctulatus LeConte: PF, WP; U. Dasytellus sp.: PF, WP; Jul; C. Hoppingiana nitida Hatch: WP; Jul; U. Trichochrous spp.: GC, PE, WP; C. Adults feed on pollen in flowers of O. polyacantha. GREAT BASIN NATURALIST Vol. 46, No. 2 Nitidulidae Brachypterolous pulicarius (Linnaeus): WP; Jul; U. Carpophilus pallipennis (Say): GC, PE, WP; Jun—Aug; C. Adults feed on pollen in flow- ers of O. polyacantha. Phalacridae Olibrus rufipes LeConte: GC, PE; Aug; C. Adults feed on pollen in flowers of C. visci- difloris. Phalacrus penicillatus Say: GC, PE, SW; U. Adults associated with E. cinereus. Coccinellidae Brachyacantha dentipes socialis Casey: GC; Aug; U. Brachyacantha ursina uteella Casey: GC, SW; Jun, Jul; U. Brumus septentrionus Weise: GC; Jun—Aug; We Coccinella difficilis Crotch: WP; May, Jun; U. Coccinella novemnotata degener Casey: SW; Aug; U. Coccinella prolongata Crotch: WP; Jun; U. Coccinella transversoguttata richardsoni Brown: GC, LT; May—Aug; U. Hippodamia apicalis Casey: GC, SW, WP; Jun—Aug; A. Hippodamia — convergens May—Aug; U. Hippodamia glacialis lecontei Mulsant: WP; Aug; WE Hippodamia quinquesignata Kirby: GC; May; \O, Hippodamia tredecimpunctata tibialis (Say): Lyle ZANE AN Hyperaspidius hercules Belicek: PF, WP; Jun—Aug; U. Hyperaspidius vittigera LeConte: WP; Aug; U Guerin: SW; Hyperaspis lateralis montanica Casey: MT, SW, WP; May—Aug; U. Nephus sordidus (Horn): PF; Jun; U. Scymnus caurinus Horn: SW; Jun; U. Scymnus marginicollis Mannerheim: PF, WP; Jun—Aug; U. ) Scymnus postpictus Casey: WP; Jul; U. Lathridiidae Melanopthalma americana Mannerheim: BF; May; U. Melanopthalma sp.: SW; Jul, U. April 1986 Tenebrionidae Alaudes singularis Horn: BF; Mar; U. Araeoschizus airmeti Tanner: GC, PF; May-— Aug; U. Inquiline in the nests of the harvester ant, Pogonomyrmex owhyheei Cole Blapstinus barri Boddy: PF, WP; May, Jun; U. Blapstinus substriatus Champion: PF, WP; May-Jul; C. Coelocnemis punctatus LeConte: PF; Aug; U. Coniontis obesa LeConte: PF; May, Jun; U. Coniontis setosa Casey: BF, PF; Mar, May, Jun; U. Eleodes cordata Eschscholtz: PF; May—Aug; U. Eleodes extricata cognata Haldemann: GC, PF; May—Aug; A. Eleodes hispilabris connexa LeConte: PF; May-Jul; U. Eleodes nigrina LeConte: GC; Jul, Aug; U. Eleodes obscura Say: PF; May—Aug; U. Eleodes pilosa Horn: GC, PF; May—Aug; A. Embaphion elongatum Horn: PF; May-—Jul; U. Helops californicus Mannerheim: PF; May, Jun; U. Helops convexulus LeConte: BF; Mar; U. Helops opacus LeConte: PF; May—Jul, U. Melanastus ater (LeConte): BF, PF, Mar, Jul; U. Oxygonodera hispidula (Horn): PF; May-— Aug; C. Sphaeriontis muricata (Casey): PF; Jun; U. Alleculidae Mycetochara procera Casey: GC, WP; May, Jun; U. Oedemeridae Oxacis bicolor (LeConte): WP; Jul; U. Mordellidae Mordellistena idahoensis Ray: PF; Jun; U. Mordellistena sericans Fall; MT, WP; Jun, Jul; U. Meloidae Epicauta immerita Walker: WP; Jun; U. Epicauta piceiventris Maydell: WP; Jul; U. Gnathium eremicola Macswain: GC, PE; Aug; A. Adults feed on nectar in flowers of C. viscidifloris. Lytta vulnerata cooperi LeConte: GC, PE; Aug; LOR Nemognatha lutea LeConte: GC; Jun; U. STAFFORD ET AL.: COLEOPTERA OF IDAHO Anthicidae Anthicus cervinus LaFerté: LT; Aug: U. Anthicus formicarius LaFerté: PF; Jul; U. Anthicus hastatus Casey: LT; Aug; U. Notoxus serratus LeConte: GC, PE, PF, WP: May—Jul; C. Adults feed on pollen and de- veloping seeds of E. cinereus. Cerambycidae Centrodera nevadica nevadica LeConte: LT, PF, WP: Jul, Aug; C. Cortodera barri Linsley & Chemsak: GC, MT, PE, PF, WP; Jun; U. Adults associated with E. cinereus. Crossidius ater LeConte: GC, PE; Aug, Sep: U. Larvae bore roots of A. tridentata. Crossidius coralinus LeConte: GC, PE; Aug; U. Larvae bore roots and adults feed on pollen of C. nauseosus. Crossidius hirtipes allgewahri LeConte: GC, PE; Aug; U. Larvae bore roots and adults feed on pollen of C. viscidifloris. Crossidius punctatus LeConte: GC, PE; Aug, Sep; U. Larvae bore roots and adults feed on pollen of C. nauseosus. Judolia gaurotoides Casey: GC, PF; May; U. Mecas bicallosa Martin: PE; Jun; U. Larvae bore roots of sagebrush. Megacheuma_ brevipennis (LeConte): GC, PE; Aug; C. Larvae bore roots of A. confer- tifolia, A. nuttali, and Sacrobatus vermicu- latus (Hook.) Torr. Megasemum asperum (LeConte): LT; Jul; U. Prionus californicus Motschulsky: GC; Aug; U. Bruchidae Acanthoscelides pauperculus (LeConte): GC, WP; Jun—Aug; U. Chrysomelidae Altica sp.: GC; Jun; U. Adults associated with E. cinereus. Anisostena californica Van Dyke: GC; Aug; U. Adults associated with E. cinereus. Brachycoryna montana (Horn): GC; Jul; U. Chaetocnema sp.: PF; Jun, Jul; U. Crepidodera nana (Say): GC; Jun; U. Cryptocephalus spurcus LeConte: GC, SW; Jul; U. Adults and larvae feed on foliage of C. viscidifloris. Disonycha latifrons Schaeffer: PF, SW; May— Aug; C. Larvae and adults feed on foliage of C. viscidifloris. 292 Exema conspersa (Mannerheim): SW; Jul, Aug; U. Glyptoscelis sp. (poss. artemisiae Blake): PF, SW, WP; May, Jun; U. Monoxia consputa LeConte: PF, SW, WP; May, Jun; U. Monoxia pallida Blake: WP; Jul; U. Monoxia puberula Blake: GC; Jul; U. Pachybrachys caelatus LeConte; PE, SW; Jun—Aug; C. Feeds on foliage of C. viscidi- floris. Pachybrachys jacobyi Bowditch: GC; Jun— Aug; U. Phyllotreta albionica LeConte: PF, SW; May—Aug; C. Adults commonly found on wild mustards, Descurania spp. and Schoenocrambe linifolia (Nutt.) Greene. Pyrrhalta luteola (Miller): GC; Jul; U. Pyrrhalta nymphaeae (Linnaeus): GC; Jun; U Saxinis saucia LeConte: GC; Jul; U. Scelolyperus nigrovirescens (Fall): GC; Jun; Stenopodius vanduzeei Blaisdell: GC; Jul; U. Systena blanda (Melsheimer): GC; Jul; U. Trirhabda nitidicollis LeConte: GC, PE, Jul; C. Adults and larvae feed on C. nauseosus. Anthribidae Trigonorhinus sp. (near annulatus Carr): WP; Jun, Jul; U. Curculionidae Acmaegenius granicollis Van Dyke: GC; Jun; U Anthonomus sp. 1: GC, PE, SW; Jun—Aug; U. Adults feed on foliage of C. viscidifloris. Anthonomus sp. 2: GC, PE, SW; Jun—Aug; C. Adults feed on foliage of C. viscidifloris . Apion sordidum Smith: PE, PF, SW, WP; May-—Jul; U. Inquiline in tephritid and ce- cidomyiid galls on A. tridentata. Brachyrhinus ovatus (Linnaeus): GC; Jun— Aug; WE Cleonus kirbyi Casey: PF; May, Jun; U. Cleonus quadrilineatus (Chevrolat): PF; May, Jun; U. Cosmobaris americana Casey: MT; Jun; U. Dinocleus denticollis Casey: PF; May, Jun; U. Dyslobus alternatus Horn: PF; May—Aug; C. Gyrotus sinuatus Hatch: PE; Jul; U. Larvae bore in Artemisia stems. Miloderoides maculatus VanDyke: PF; Jun, polls We GREAT BASIN NATURALIST Vol. 46, No. 2 Myrmex vittatus (Horn): GC; Jul, aug; U. Ophryastes latirostris LeConte: GC, PE; Jun; U. Larvae and adults feed on foliage of C. lanata. Sitona hispidula (Fabricius): LT; Aug; U. Smicronyx sp.: PF; SW; Jun—Aug; U. Sphenophorus gentilis LeConte: PE, PF, SW; May-Jul; A. Adults feed on developing seed heads of E. cinereus. Trachyphloeni sp. (undescribed genus): PF; Jul, Aug; U. Tychius tectus LeConte: PF; June; U. ACKNOWLEDGMENTS Appreciation is extended to the following persons for the identification of specimens: R. S. Anderson, University of Alberta (Cur- culionidae: Cleonus); A. F. Newton, Museum of Comparitive Zoology—-Harvard University (Staphylinidae); R. L. Penrose, California De- | partment of Agriculture (Cerambycidae: Cen- _ trodera); C. A. Triplehorn, Ohio State Uni- versity (Tenebrionidae: Coniontis); R. L. Wenzel, Field Museum of Natural History— — Chicago (Histeridae); and D. R. Whitehead, Systematics Entomology Laboratory, USDA | (Curculionidae: Trachyphloenini). We thank — O. D. Markham, U.S. Department of En- | ergy, and Frank Merickel, University of Idaho, for their help and support. This work — was conducted for the INEL Radioecology — and Ecology Programs sponsored by the Of- — fice of Health and Environmental Research, United States Department of Energy. LITERATURE CITED ALLRED, D. M. 1968. Ticks of the National Reactor Test- _ ing Station. Brigham Young Univ. Sci. Bull. Biol. — Ser., 10(1): 1-29. _____. 1969. Spiders of the National Reactor Testing Sta- — tion. Great Basin Nat. 29: 105-108. _____. 1973. Scorpions of the National Reactor Testing | Station. Great Basin Nat. 33: 251-254. ALLRED, D. M., AND A. C. COLE. 1971. Ants of the Na- tional Reactor Testing Station. Great Basin Nat. 31: 237-242. ALLRED, D. M., AND M. M. Muma: 1971. Solpugids of the National Reactor Testing Station, Idaho. Great | Basin Nat. 31: 164-168. ARNETT, R. H., JR. 1963. The beetles of the United States. | Catholic University Press, Washington, D.C. | 1,112 pp. Harniss, R. O., ANDN. E. West. 1973. Vegetative patterns _ of the National Reactor Testing Station, South- | eastern Idaho. Northw. Sci. 47: 30-43. April 1986 STAFFORD ET AL.: COLEOPTERA OF IDAHO Hatcu, M. H. 1953-1973. The beetles of the Pacific North- west, Parts I-V. University of Washington Press. Jeppson, R. J., AND K. E. Hote. 1978. Flora of the Idaho National Engineering Laboratory Sites. Pages 129-143 in O. D. Markham ed., Ecological studies on the Idaho National Engineering Laboratory Site 1978 Progress Report. IDO-12087. U.S. Department of Energy. National Technical Information Service, Springfield, Virginia, 371 pp. 293 KINGSOLVER, J. M., ed. 1979. A catalog of the Coleoptera of America north of Mexico. USDA Agric. Hand- book 529. MCBRIDE, R., N. R. FRENCH, A. H. DAHL, AND J. E. Det- MER. 1978. Vegetation types and surface soils of the Idaho National Engineering Laboratory Site. IDO-12084. National Technical Information Ser- vice, Spriagfield, Virginia. 29 pp. MICROHABITAT AFFINITIES OF GAMBEL OAK SEEDLINGS Ronald P. Neilson’? and L. H. Wullstein! ABSTRACT. —Previous work suggested that Gambel oak seedlings are rare in the northern parts of its range in Utah | where summer rainfall is relatively low but should be abundant in southern parts of the range where summer rainfall is usually high. Gambel oak grades from a relatively minor component of a ponderosa pine/mixed conifer assemblage in | the south to a virtually monotypic formation in the north, where it exists as long-lived clones. Quadrat analysis in Arizona and New Mexico, within the oak zone, revealed a seedling density ranging from 120 to 1320 per hectare. We found a significant tendency of seedlings to be located on the NE (cool, shady) side of sheltering objects in the environment. Mature ponderosa pine ranged in density from ca 40 to 500 stems per hectare, whereas mature Gambel oak ranged from ca 10 to 20 genets per hectare with ca | to 7 ramets per clone. These results support our previous conclusion that Gambel oak in northern Utah probably became established as a minor component of a mixed | pine/oak woodland at a time in mid-Holocene when summer rainfall was much higher than today. Gambel oak (Quercus gambelii Nutt.), a deciduous, white oak, is the dominant oak of the southern Rocky Mountain region. Its dis- tribution is primarily encompassed by the states of Utah, Colorado, Arizona, and New Mexico. We previously demonstrated that the northern limits of Gambel oak in Utah appear to be constrained by the combined effects of two distinct airmass gradients (Neilson and Wullstein 1983). Probabilities of late spring freeze as determined by the polar front gradi- ent and summer drought as determined by the “Arizona Monsoon’ gradient appear to co- vary during global warming and _ cooling trends and appear to have synergistically pro- duced a relatively sharp northern boundary (Neilson and Wullstein 1983). At present, seedling establishment of Gambel oak is rare in the northern part of its range (Neilson and Wullstein 1983). Our transplant studies (seeds and seedlings, Neilson and Wullstein 1983) and physiological studies (Neilson and Wullstein 1986) indicate that natural seedling establishment may be expected to occur only in the parts of the range where summer rains are sufficient for seedling survival. Gambel oak persists at its northern limits today by virtue of rhizomatous, asexual repro- duction (Neilson and Wullstein 1983). We be- lieve that these oaks became established at their northern limits through sexual repro- duction and seed dispersal at some time dur- ‘Department of Geography, University of Utah, Salt Lake City, Utah 84112. °Present address: University of Utah Research Institute, Salt Lake City, Utah 84108. ing the mid-Holocene thermal maximum _ when limiting stresses would have been re- | duced (Neilson and Wullstein 1983). At that | time the shrub and tree community composi- | tion in northern Utah, near the northern lim- | its of distribution for this species, might have | been similar to that where the species is capa- | ble of sexual reproduction and seedling estab- | lishment today. Near its northern limits to-_ day, where it reproduces asexually, Gambel | oak forms an essentially monotypic plant for- mation, or “mountain brush community — (Ream 1963). In the southern part of its range, | where it reproduces sexually, Gambel oak is a | relatively minor component of a mixed pine- oak woodland. The purpose of this study is to. document the density of Gambel oak seed- lings in regions of high summer rainfall, their | microhabitat affinities and the general canopy composition of their associated plant commu- , nities. METHODS In September 1979, 15 quadrats were es- tablished in Arizona and New Mexico to ascer- — tain the density of Gambel oak seedlings in various habitats. Excavation of root systems revealed that seedlings can usually be distin-, guished from suckers on the basis of cluster-. ing. Suckers tend to occur in tight clusters, while seedlings are widely dispersed. In many — 294 April 1986 cases the spent acorn was still attached to the seedling. It was also found that true seedlings possessed a readily extractable, vertically ori- ented taproot, which exhibited well-defined taper, whereas suckers were always attached to a horizontally oriented rhizome within a few cm of the surface, which exhibited little or no taper. Thereafter, seedlings were distin- guished from suckers by counting shoots that exhibited isolation from other shoots by more than two meters. In doubtful instances plants were examined for tap roots. All oaks less than 30 cm in crown height were arbitrarily classi- fied as seedlings unless otherwise determined to be suckers. Fourteen large seedlings were collected at random for estimations of age and taproot growth rates. Of the 15 quadrats, 13 were 0.1 ha and square. The remaining two quadrats were 0.04 and 0.05 ha, respectively, and roughly square. The quadrat sites were chosen to re- flect apparent extremes of habitat within an area. In areas exhibiting little variation in to- pography (e.g., two flat sites south of Flag- staff), the quadrat placement was essentially random. In areas of diverse topography, the quadrats were chosen to encompass previous seedling plantings (Neilson and Wullstein 1983), selected to reflect mesic and xeric ex- tremes within the region. Three quadrats were placed in each of two flat sites on the Mogollon Rim south of Flagstaff near the headwaters of Oak Creek Canyon. Three contiguous quadrats on Humphrey Peak (near Flagstaff, Arizona) sampled a gradient from swale to ridge on an east-facing slope. Three quadrats were estab- lished in the vicinity of Pinedale, Arizona, on the Mogollon Rim. Two of these were on flats separated by several kilometers. The third was on a south-facing slope. There were no previous plantings in this vicinity. Three quadrats were placed in the Sandia Mountains of New Mexico. Two of these were on east-fac- ing ridges of differing moisture regime located near Cold Spring Picnic Ground (see Table 1). The third was on a southfacing slope. Lati- tude, longitude, elevation, slope, aspect, gen- '. eral canopy composition and density were Lb - noted. These data are summarized in Table 1. As the natural seedlings were counted, they _ were marked with orange paint to eliminate double counting and to insure a complete NEILSON, WULLSTEIN: GAMBEL OAK 295 count. The location of each seedling was noted with regard to surface drainage patterns, whether it was sheltered or unsheltered from the sun (by trees, shrubs or rocks), and, if sheltered, its compass orientation (in 4 quad- rants) with respect to the sheltering object. Seedling distribution with regard to shelter- ing objects was analyzed with a Chi-square test. In 3 of the 15 quadrats seedlings were simply counted, since the canopy was essen- tially closed, i.e., all seedlings in these quad- rats were sheltered from the sun. RESULTS The general physical characteristics of the 15 quadrats are presented in Table 1. The communities ranged from dominant pon- derosa pine (Pinus ponderosa) to mixed conifer communities. Gambel oak was usually a relatively minor component. More com- plete descriptions of these communities may be found in Layser and Schubert (1979). The three contiguous quadrat sites on Humphrey Peak (Table 1) were of low-density ponderosa pine averaging 43 + 35 pines/ha with an average dbh of ca. 40 cm. Gambel oak density on each quadrat was ca 20 genets/ha with 1-2 stems per clone and an average dbh of 17 cm/stem. These quadrats represent a gradient from a swale to a neighboring ridge. This is most evident by acombined increase in Cowania sp. and Cercocarpus sp. from zero in quadrat | to 8 in quadrat 2 and more than 100 in quadrat 3. Gambel oak seedling density was relatively high (range 360-590) and without apparent pattern along this steep environ- mental gradient. Quadrats 4-9 were located on the flat Mogollon Rim south of Flagstaff in mature ponderosa pine forest averaging 350 + 112 stems/ha with an average dbh (in each quadrat) ranging from 31 to 46 cm. Mature oak density was low in these quadrats, averaging ca 10 genets/ha with 1-7 stems per clone and an average dbh of ca 16 cm per stem. Seedling density in these six quadrats ranged from 240 to 860 ha. Quadrat 9, which had the highest seedling density (860/ha) of this group, was flanked by five large oaks. Quadrats 10-12 represent three distinct habitats in the Pinedale area of the Mogollon Rim, a south-facing slope (No. 12), a relatively 296 GREAT BASIN NATURALIST Vol. 46, No. 2 TABLE 1. Site characteristics for the 15 quadrats (redundant data are not repeated within a geographic region). Quadrat Geographic Latitude Longitude Elevation Slope Aspect Soil WHC’ Canopy number region (decimal) (decimal) (meters) species” 1 Humphrey Peak 35.27 111.6 2226 Wa S 80° E 32-88/45-60 PIPO/QUGA 2 25° oN 70° E PIPO/QUGA 3 25° N 75° E PIPO/QUGA 4,5,6 Flagstaff 35.03 111.73 2165 0° 40-90/50-77 PIPO/QUGA 7,8,9 35.00 111.67 8° N 65° E 40-90/50-77 PIPO/QUGA 10 Pinedale 34.3 110.25 1982 4° N 45° W- unknown PIPO/QUGA 11 2012 0° PIPO/QUGA/ JUXX 12 1982 20° S 10° E PIPO/QUGA/ JUXX 13 Sandia Mountains 35.13 106.53 2226 15° S 80° E 46-122/58-65 PIPO/PIED/ JUXX/QUGA 14 10° S 60° E 40-85/42-46 PIPO/PIED/ JUXX/QUGA 15 35.15 106.53 2165 23° S 30° W 54-66/60-62 PIPO/PIED/ JUXX/QUGA 'WHC = Water Holding Capacity (%). A range of water holding capacities is reported for the soil surface layer and at a depth of 30 cm (Neilson and Wullstein 1983). 2PIPO = Pinus ponderosa, QUGA = Quercus gambelii, PIED = Pinus edulis, JUXX = Juniperus sp. flat lowland area (No. 10), and a flat plateau area. All three areas possessed a high density of seedlings, with the plateau exhibiting the greatest number (1,080/ha). All three areas had been artificially thinned, but still pos- sessed a fairly dense canopy. Quadrats 10 and 12 were primarily mature ponderosa pine with ca 500 stems/ha and an average dbh of 18 and 27 cm, respectively. Quadrat 11 was mixed conifer/oak woodland. Quadrats 13-15 in the Sandia Mountains, New Mexico, were characterized by mixed conifer forest with a Gambel oak component. Quadrat 15, on a south-facing slope, was simi- lar in seedling density to most other quadrats, with 513 seedlings/ha. Quadrats 13 and 14 were on east-facing ridges. These quadrats were of the same elevation and aspect and were within 1 km of each other. Quadrat 14 was located on a primary ridge separating ma- jor drainage basins. It was at a lower angle than No. 13 and contained more bare ground. Quadrat 13, by contrast, was located on a ridge within a major drainage basin, was at a steeper angle, and contained less bare ground than No. 14. By virtue of their close proxim- ity, the two quadrats would be expected to receive similar amounts of rainfall. Because of its position as a drainage divide, the primary input of moisture to Quadrat 14 was likely limited to incident rainfall. Quadrat 13, how- ever, received moisture from both rainfall and runoff from the surrounding basin within which it occurred. This was particularly evi- dent from extensive rill development, which was absent on Quadrat 14. Quadrat 13 pos- sessed 1,320 seedlings/ha in contrast to 120 seedlings/ha in Quadrat 14. Table 2 indicates the number of seedlings in sheltered and unsheltered positions for each of the quadrats. Quadrats 11, 13, and 15 were too dense to allow judgements of sheltering. Sixty percent (341/571) of the seedlings in 12 quadrats (i.e., excluding quadrats 11, 13, and 15) were scored for their orientation with re- gard to a sheltering object (Table 2, Fig. 1). The distribution is significantly skewed (X* = 33.74, P < .01) to the NE quadrant. Age estimates of 14 randomly collected seedlings (<30 cm crown height) ranged from 6 to 17 years. Regressions of root length against root diameter indicated root taper ranging from —.06 to —.19 mm/cm (r° ranged from .81 to .99). Since these measurements were obtained from broken taproots, total root length was extrapolated from the regres- sion equations and found to range from 40 to 114 cm. Root growth rates were estimated to — range from 2.7 to 11.4 cm/yr. DISCUSSION Several points are apparent from these data. First, Gambel oak seedlings are abun- dant and widely distributed in the southern part of the range of this species and are rare in April 1986 TABLE 2. Total seedlings (sheltered and unsheltered) per quadrat. Seedlings/ Quadratno. Sheltered Unsheltered _ hectare a 40 19 590. 2 16 20 360 3 32 16 480 4 24 20 440 5 29 6 350 6 14 10 240 lh 19 20 390 8 12 13 250 9 58 28 860 10 50 37 870 ll unknown unknown 1080 12 35 4] 760 13 unknown unknown 1320 14 12 0 120 15 unknown unknown 513 the northern part of the range. Second, ma- ture Gambel oak is a relatively minor compo- nent of the ponderosa pine—mixed conifer forest. Third, although Gambel oak does clone in Arizona and New Mexico, the appar- ent number of ramets per clone is relatively few (1-7 in the quadrats sampled, although larger numbers can be found) compared to the hundreds to thousands of ramets per clone that are common in the northern parts of the range (Brown 1958, Ream 1963). A corollary to this point is that mature ramets are much larger in the south than in the north, where dbh is typically in the range of 5-8 cm (Brown 1958, Ream 1963) compared to 16-17 cm in the south. Fourth, although seedlings are widespread across a range of microhabitats, their distribution and abundance does appear to reflect some dependence on soil moisture. This last point is most interesting. We pre- viously reported (Neilson and Wullstein 1983) _ that seedling survival in the southern parts of _ the range was largely independent of micro- habitat, being high in all situations from shel- tered to unsheltered. The three contiguous quadrats on Humphrey Peak are in support of this contention. Whereas the shrub distribu- ‘tion indicated considerable edaphic and/or | hydrologic differences between the swale and i ridge habitats, the seedling density across this ‘gradient indicated no trend. Nevertheless, ' 60% of the seedlings in these and the other quadrats did exhibit a strong pattern of shel- tering, apparently favoring mesic microhabi- NEILSON, WULLSTEIN: GAMBEL OAK 297 N 41+14% W E 22+8% n= 341 P<0.01 S Fig. 1. Compass orientation of seedlings (% per quadrant) associated with a sheltering object in all quadrats, excluding quadrats 11, 13, and 15 (see text for explanation). tats. This is further supported by the two ridge quadrats in the Sandia Mountains. One ridge was apparently swept frequently by sur- face flow, as indicated by the extensive rill development and the location of the ridge within a major drainage basin. Intense sum- mer thunder showers are common in this re- gion (Bryson and Lowry 1955). This ridge con- tained the highest density of seedlings observed in any of the quadrats, notwith- standing the apparently high surface wash. A neighboring ridge, a major drainage divide, apparently received little to no runoff from the surrounding landscape, as indicated by the absence of rills. This ridge contained the lowest density of oak seedlings observed (all of which were sheltered) in any of the quadrats. In almost every respect but hydrology, the two ridges appear to be similar. This suggests that even in a region where summer rain is relatively high, a consistently mesic micro- habitat may be required to provide adequate soil moisture for seedling survival. The six quadrats on the relatively flat Mogollon Rim south of Flagstaff present some evidence that the density of seedlings is not entirely independent of the density of adults. Five of these quadrats possessed seedling densities ranging from 250 to 440 per hectare (quadrats 4-8), whereas one (quadrat 9) pos- sessed a seedling density of 860 per hectare (Table 2). All these quadrats contained a simi- lar density of mature oaks. However, quadrat 9 with the highest seedling density in this 298 region, was flanked by five large oaks, sug- gesting some dependency of seedling density on the local density of seed—bearing trees. Since seedling density was found to be rela- tively high throughout the oak zone in the southwest, why are mature oaks relatively rare in these forests? It may be that conifer seedlings, which are also abundant in these forests, grow much more rapidly than the oaks and gain dominance through competition for light and moisture. Several oak seedlings (of less than 30 cm stature) were aged on the basis of taproot rings and were judged to be from 6 to 17 years old. By contrast, ponderosa pine raised in several provenance gardens in the Great Plains grew between 0.7 and 3.9 m in 10 years (Read 1983). Thus, overtopping of oak seedlings by ponderosa pine seedlings may competitively inhibit the growth of oaks. Once past the establishment phase, the po- sition of oaks in the landscape is apparently independent of minor differences between microhabitats. The density of mature oaks was relatively uniform between all the quadrats, which varied considerably in topography. The ultimate fate of seedlings, once established (i.e., taproot developed), is likely determined by biotic factors, including competition with pines and herbivory from insects and verte- brates (primarily rabbits and cows) (Neilson and Wullstein 1983). In conclusion, where Gambel oak is least abundant in its geographic range, it is capable of sexual reproduction, seed dispersal, and seedling establishment. Conversely, where Gambel oak is most abundant in its geo- graphic range, near the northern limits, its mode of reproduction is primarily asexual. The establishment of Gambel oak seedlings is dependent on consistent soil moisture, even in areas where summer rainfall is high. These GREAT BASIN NATURALIST Vol. 46, No. 2 results support the hypothesis that Gambel oak in northern Utah, existing today by asex- ual reproduction, was likely established as a relatively minor component of a mixed pine/ oak woodland under a mid-Holocene climate with more favorable summer moisture condi- tions in northern Utah than presently occur (Cottam, Tucker, and Drobnick 1959, Neilson and Wullstein 1983). ACKNOWLEDGMENT We thank R. Seiler for valuable assistance in data collection. This work was supported by the Department of Biology, University of Utah. LITERATURE CITED Brown, H. E. 1958. Gambel oak in west-central Colo- rado. Ecology 39: 317-327. BrySON, R. A., AND W. P. Lowry. 1955. Synoptic climatol- ogy of the Arizona summer precipitation singular- ity. Bull. Amer. Meteorol. Soc. 36: 329-339. CotraM, W. P., J. M. TUCKER, AND R. DROBNICK. 1959. .Some clues to Great Basin post-pluvial climates provided by oak distributions. Ecology 40: 361-377. LaysER, E. F., AND G. H. SCHUBERT. 1979. Preliminary classification for the coniferous forest and wood- land series of Arizona and New Mexico. USDA For. Serv. Res. Pap. RM-208. 27 pp. NEILSON, R. P., AND L. H. WULLSTEIN. 1983. Biogeogra- phy of two southwest American oaks in relation to atmospheric dynamics. J. Biogeog. 10: 275-297. . 1986. Comparative drought physiology and bio- geography of Quercus gambelii and Quercus tur- binella. Amer. Mid]. Nat. 114: 275-297. READ, R. A. 1983. Ten-year performance of ponderosa pine provenances in the Great Plains of North America. USDA For. Serv. Res. Pap. RM-250. 17 pp- REAM, R. R. 1963. The vegetation of the Wasatch Moun- | tains, Utah and Idaho. Unpublished dissertation, University of Wisconsin, Madison. 178 pp. ; | q i} | FEEDING HABITS OF METAMORPHOSED AMBYSTOMA TIGRINUM MELANOSTICTUM IN PONDS OF HIGH pH (© 9) Brian T. Miller’ and John H. Larsen, ita ABSTRACT.—During the spring breeding season throughout the channeled scablands of eastern Washington, meta- morphosed male and female blotched tiger salamanders (Ambystoma tigrinum melanostictum) utilize oropharyngeal suction to capture large quantities of small aquatic invertebrates. Stomach content analysis on salamanders from three populations of this subspecies revealed that they consume the following taxa: Copepoda, Cladocera, Culicidae, Anostraca, and Chironomidae. Although the amount of energy obtained by adults via in-water feeding was not calculated, the large volume of aquatic invertebrate material flushed from salamander stomachs suggests that this feeding strategy should add significantly to their total annual nutrient consumption. Investigations on the feeding habits of Am- bystoma tigrinum have centered primarily on the diets of branchiate individuals. This onto- genetic stage, depending on individual size, season, and locality, feeds daily on large quan- tities of a variety of prey items (Olenick and Gee 1981, Brophy 1980, Dodson and Dodson 1971). As a result of their secretive terrestrial habits (Nussbaum et al. 1983, Smith 1961, Sever and Dineen 1978) similar information on diet and feeding for transformed individu- als has been difficult to obtain. Reports on the feeding habits of this developmental stage are largely restricted to general accounts that list a variety of terrestrial invertebrates (most of- ten those accepted by captive salamanders) as potential prey (Conant 1975, Nussbaum et al. 1983, Johnson 1977, Smith 1961). Since some metamorphosed A. t. melanostictum in cen- tral and eastern Washington remain in the ponds for considerable period of time as adults during the breeding season or as subadults immediately following transformation, such individuals could utilize the abundant aquatic invertebrate communities as an _ energy source. In numerous locations larval A. t. mavor- tium metamorphose during late summer or early fall and remain in the ponds throughout the winter (Webb and Roueche 1971). The extent to which these transformed individuals feed during this period is unknown, but Webb and Roueche (1971) found aquatic prey in the stomachs of 13 subadults collected in the stomachs of 13 subadults collected in Dona Ana County, New Mexico, in February, and Burger (1950) reported that the principal item in the stomachs of adult A. t. nebulosum was the pond snail, Lymnaea stagnalis. Sexually mature, metamorphosed tiger salamanders return to the ponds to breed. Egg deposition typically occurs during late winter or early spring (Semlitsch 1983, Hassinger et al. 1970, Bishop 1943, Sever and Dineen 1978), whereas migrations to the ponds occur during spring and/or autumn (Semlitsch 1983, Stine 1984, Smith 1961). Even though transformed A. tigrinum may spend several months in the water, they reportedly feed very little if at all (Rose and Armentrout 1976, Stine 1984). However, we found the stomachs of trans- formed male and female A. t. melanostictum, collected from ponds in Washington State during the spring breeding season, often filled with aquatic invertebrates. The following re- ports on in-pond feeding by breeding congre- gations of transformed A. t. melanostictum and its possible significance. MATERIALS AND METHODS Transformed tiger salamanders were seined from three fish-free ponds in the chan- neled scablands of central and eastern Wash- ington (14 April to 6 May 1984). This region is a network of scoured canyons and deep valleys carved out of basalt bedrock and loess by the Spokane flood of 20,000 years ago (Bretz Department of Zoology, Washington State University, Pullman, Washington 99164-4220. Electron Microscopy Center, Washington State University, Pullman, Washington 99164-4210. 299 300 1959). Thin soils characterize the scablands, and the dominant vegetation is a sagebrush (Artemesia )-wheatgrass (Agropyron) associa- tion (Daubenmire 1970). Nineteen salaman- ders were collected from a saline (300 mOs,,), alkaline (10.1 pH) pond in southern Grant County, 11 km from an alkaline (10.1 pH) pond of low salinity (<5 mOs,,) in southern Lincoln County, and 9 km from an ephemeral pond in northernWhitman County. Follow- ing capture the specimens were immediately immersed in an ice solution and returned to the laboratory, where they were either anes- thetized in 1:1000 MS222 or frozen to —80 C. Stomach contents were obtained from either the anesthetized individuals using a flushing method modified after that of Legler and Sul- livan (1979) or from the excised stomachs of the frozen animals. All stomach contents were stored in 70% ETOH until identified. RESULTS AND DISCUSSION Thirty-six (92%) of the 39 transformed A. t. melanostictum examined contained aquatic invertebrates in their stomachs; material in the remaining three individuals could not be identified. No apparent differences in the per- centage of stomachs containing food were de- tected between males and females from any of the three populations. The structure of the aquatic invertebrate communities varies from pond to pond, and therefore the differences in the diets of these salamander populations re- flect this factor. The kinds of prey utilized by this subspe- cies in eastern Washington ponds are similar to those reported for various larval con- specifics (Brophy 1980, Dodson and Dodson 1981, Dineen 1955) and related larval con- generics (Branch and Altig, 1981). Although the total number of food items per stomach varied from 7 to 431 (x = 118), the most com- mon prey eaten by salamanders inhabiting the ponds included: Copepoda and Cladocera (Lincoln County); Copepoda, Cladocera, and Anostroca (Whitman County); and Chirono- midae and Culicidae (Grant County) (Table 1). Although not quantified, these inverte- brates also composed the largest stomach con- tent volume in the majority of water-collected salamanders. Therefore, small aquatic inver- tebrate taxa are responsible for the bulk of the GREAT BASIN NATURALIST Vol. 46, No. 2 TABLE 1. Summary of stomach content analyses for metamorphosed A. t. melanostictum from eastern and central Washington ponds. Percentage of stomachs containing taxon/ percentage of total items consumed / average number per stomach containing taxon Lincoln Co. Whitman Co. Grant Co. N=11 N=9 N = 16 Copepoda 91/55/78 100/75/102 6/10/164 Cladocera 73/26/47 = — Anostraca — 89/21/32 — Ostracoda 73/04/06 67/02/03 — Culicidae 9/ */O1 — 100/45/044 Chironomidae 9/ */01 = 94/41/043 Trichoptera 73/06/11 33/01/04 — Notonectidae 27/ */01 22/ */01 31/01/002 Corixidae — 1l/ */01 38/ */001 Odonata 9/ */O1 — 13/ */001 Dytiscidae 82/03/05 78/02/04 19/02/009 Curculionidae 36/ */01 = — Gastropoda 45/05/14 — — Acarina — — 6/ */001 *< 0.5% diet of transformed adult A. t. melanostictum from the ponds investigated. It is clear from our observations on meta- morphosed A. t. melanostictum maintained in aquaria and supplied quantities of aquatic in- vertebrates that prey capture is accomplished by oropharyngeal suction, a nearly universal method of those vertebrates feeding in water (Bramble and Wake 1985). The large number of prey items present in the stomachs of both | male and female transformed salamanders in- | dicates that they feed extensively while in the ponds during the breeding season. This ob- — servation differs from that of Rose and Armen- trout (1976); they rarely found food in the stomachs of metamorphosed A. t. mavortium collected from ponds. Smallwood (1928) de- termined that adult A. maculatum stop eating when they become aquatic during the breed- ing season, and Anderson (1968) reported that transformed male, but not female A. macro- dactylum feed in the ponds. The amount of time that A. t. melanos- tictum spends in water during the year is un- known. Therefore, the importance of energy obtained from in-water prey capture has not been determined. However, such feeding — probably adds significantly to the total yearly | energy intake, particularly within the chan- | neled scablands of Washington where terres- trial conditions are harsh and land-living prey may be less readily available. April 1986 Metamorphosed A. t. melanostictum appar- ently spends long periods of time each year in these high pH ponds that readily support ex- ploitable, abundant aquatic invertebrate com- munities and contain few predatory fish (Tillett 1984). This strategy is probably adaptive, since the terrain adjacent to these scabland ponds is, at best, marginal salamander habitat. ACKNOWLEDGMENTS The authors are grateful to R. E. Herrington and E. Johnson for their assistance in the field. LITERATURE CITED ANDERSON, J. D. 1968. A comparison of the food habits of Ambystoma macrodactylum _ sigillatum, Am- bystoma macrodactylum croceum, and Am- bystoma tigrinum californiense. Herpetologica 24: 273-284. BisHop, S. C. 1943. Handbook of salamanders, Comstock Publ. Co., Ithaca, New York. 555 pp. BRAMBLE, D. M., AND D. B. WAKE. 1985. Feeding mecha- nisms of lower tetrapods. Pages 230-261 in M. Hildebrand, D. M. Bramble, K. F. Liem, and D. B. Wake, eds., Functional vertebrate morphol- ogy. Belknap Press of Harvard University Press, Cambridge, Massachusetts. BRANCH, L. C., AND R. ALTIG. 1981. Nocturnal stratifica- tion of three species of Ambystoma larvae. Copeia 1981: 870-873. BrETz, J. H. 1959. Washington’s channeled scablands. Washington State Dept. Cons., Div. Mines and Geol. Bull. 45. 57 pp. Bropny, T. E. 1980. Food habits of sympatric larval Am- bystoma_ tigrinum and Notophthalmus _ viri- descens. J. Herpetol. 14: 1-6. BurRGER, W. L. 1950. Novel aspects of the life history of two Ambystomas. J. Tenn. Acad. Sci. 25: 252-257. ConanT, R. 1975. A field guide to reptiles and amphibians of eastern and central North America. Houghton- Mifflin Co., Boston. 429 pp. DAUBENMIRE, R. 1970. Steppe vegetation of Washington. Washington Agr. Exp. Sta. Tech. Bull. 62. 131 pp. DINEEN, C. F. 1955. Food habits of the larval tiger sala- mander, Ambystoma tigrinum. Proc. Indiana Acad. Sci. 65: 231-233. MILLER, LARSEN: SALAMANDER FEEDING HABITS 301 Dopson, S. I., AND V. E. Dopson. 1971. The diet of Am- bystoma tigrinum larvae from western Colorado. Copeia 1971: 614-624. HASSINGER, D. D.,J. D. ANDERSON, AND G. H. DALRYMPLE. 1970. The early life history and ecology of Am- bystoma tigrinum and Ambystoma opacum in New Jersey. Amer. Mid]. Nat. 84: 474-495. JOHNSGN, T. R. 1977. The amphibians of Missouri. Univ. Kansas Mus. Nat. Hist. Publ. Ed. Ser. 6; 1-134. LEGLER, J. M., AND L. J. SULLIVAN. 1979. The application of stomach-flushing to lizards and anurans. Her- petologica 35: 107-110. NussBauM, R. A., E. D. BRODIE, JR., AND R. M. StoRM. 1983. Amphibians and reptiles of the Pacific Northwest. University Press of Idaho, Moscow. 332 pp. OLENICK, R. J., AND J. H. GEE. 1981. Tiger salamanders (Ambystoma tigrinum) and stocked rainbow trout (Salmo gairdneri): potential competitors for food in Manitoba prairie pothole lakes. Canadian Field- Naturalist 95: 129-132. Rose, F. L., AND D. D. ARMENTROUT. 1976. Adaptive strategies of Ambystoma tigrinum Green inhabit- ing the Llano Estacado of west Texas. J. Anim. Ecol. 45: 713-729. SEMLITSCH, R. D. 1983. Structure and dynamics of two breeding populations of the eastern tiger salaman- der, Ambystoma tigrinum. Copeia 1983: 608-616. SEVER, D. M., AND C. F. DINEEN. 1978. Reproductive ecology of the tiger salamander, Ambystoma ti- grinum, in northern Indiana. Proc. Indiana Acad. Sci. 87: 189-203. SMALLWOOD, W. M. 1928. Notes on the food of some Onodaga Urodela. Copeia 1928: 89-98. SMITH, P. W. 1961. The amphibians and reptiles of IIli- nois. Bull. Illinois Nat. Hist. Surv. 28: 1-298. STINE, C. J. 1984. The life history and status of the eastern tiger salamander, Ambystoma tigrinum tigrinum (Green) in Maryland. Bull. Maryland Herp. Soc. 20: 65-108. TILLETT, W. E. 1984. Aquatic studies in the Columbia Basin project. Pages 149-208 in J. H. Foster, W. E. Tillett, W. L. Meyers, J. C. Hoag, eds., Co- lumbia Basin Wildlife/irrigation development study. Report No. REC-ERC-83-6 U.S. Dept. Interior, Bureau Reclamation. Wess, R. G., AND W. L. ROUECHE. 1971. Life history as- pects of the tiger salamander (Ambystoma ti- grinum mavortium) in the Chihuahuan Desert. Great Basin Nat. 4: 193-212. NOTES ON THE SWAINSON’S HAWK IN CENTRAL UTAH: INSECTIVORY, PREMIGRATORY AGGREGATIONS, AND KLEPTOPARASITISM Neil D. Woffinden! ABSTRACT. —A premigratory flock of Swainson’s Hawks numbering at least 213 individuals was observed during July and August of 1984. Aerial feeding on grasshoppers was noted and kleptoparasitism was recorded between the Swainson’s Hawk and the American Kestrel. Swainson s Hawk (Buteo swainsoni) inhab- its open spaces such as plains, prairies, and deserts. Prior to the turn of the century this species was an abundant resident of several western states (Bent 1937). A specimen was collected in Utah’s Wasatch Mountains as early as 1868, and the species was reported as a common nesting resident of Summit County, Utah, in 1877 (Hayward et al. 1976). Yet, Swainson’s Hawk could not be consid- ered abundant in central Utah today. For in- stance, during a 12-month Great Basin raptor survey 1,275 individual raptors representing 12 species were sighted, yielding an average of one sighting per every 10 km traveled. By comparison, nearly 400 km were traversed between each Swainson’s Hawk sighting (Woffinden and Murphy 1977a). Further- more, only one nesting pair has been reported for Cedar Valley, Utah County, Utah (40°00'N, 111°55’—112°35’W) during the past decade (Woffinden and Mosher 1979). Cedar Valley, situated near the eastern limit of the Great Basin, is characterized by typical cold desert vegetative associations (Shelford 1963). Two small villages, Cedar Fort and Fairfield, are located in the exten- sively cultivated northern portion of the val- ley. Except for numerous black willows (Salix nigra) associated with these communities, the valley is devoid of large trees. The area sup- ports a sizable raptor community and has been the site of anumber of recent studies (Murphy et al. 1969, Smith and Murphy 1973, Woffinden and Murphy 1977b). On 23 July 1984, 36 Swainson’s Hawks were observed immediately west of Fairfield. Sub- sequently, 213 individuals were counted dur- ing a visit to the area on 6 August, and 59 were observed on 16 August 1984. All the birds were dark or rufous phase individuals (Brown and Amadon 1968:585), and many had lost flight and/or tail feathers. This unusual accu- mulation represents the largest flock of Swain- son's Hawks ever reported for this portion of the state. Although the aggregation was apparently a premigratory gathering, it was impossible to determine accurate arrival and departure dates. - However, a local resident stated that large numbers of hawks first appeared near the middle of July. Large flights of Swainson’s Hawks have been reported for several areas throughout the United States (Bent 1937, Cruickshank 1937, Fox 1956, Martin and McEneaney 1984), but premigratory accumu- | lations of the magnitude reported here have | rarely been recorded. Swainson ’s Hawk is noted for its insectivo- rous diet (Bent 1937, Taylor 1946, Hayward et al. 1976). For instance, White (1966) recov- ered 230 crickets from a female collected on a Kansas farm. However, the species consumes a variety of other organisms, including mam- mals, birds, reptiles, amphibians, fish, cray- fish, and beetles (Bent 1937, White 1966, Sex- ton and Marion 1974, Dunkle 1977, Bechard 1983). In spite of the early reports of insec- tivory, recent studies suggest that mammals make up the major portion of the hawk’s diet (Dunkle 1977, Fitzner 1977, Bechard 1980, 1983). soaring near Fairfield, Utah, on the afternoon of 23 July 1984. An additional 16 individuals ‘Division of Natural Sciences, University of Pittsburgh at Johnstown, Johnstown, Pennsylvania 15904. 302 Twenty Swainson’s Hawks were observed © ~ : a a April 1986 were perched in a group of nearby trees. Swarms of grasshoppers (Melanoplus fermur- rubrum) were observed in the area flying from the ground or vegetation and ascending to considerable heights. Soaring hawks were observed repeatedly extending one or both feet to quickly snatch ascending grasshoppers. All but four of the perched birds joined the soaring flock during the 15 minutes they were observed actively feeding. By 1745 h the hawks had drifted out of sight to the west, and only a few airborne grasshoppers were still visible. This unique feeding behavior corresponds exactly with an account in Bent (1937:229). Similar Swain- sons Hawk feeding behavior was noted dur- ing one other visit to the area. Although the hawks were taking advantage of an abundant food source, it is not known if the grasshopper outbreak was responsible for the unusual aggregation noted. Grasshoppers were also abundant during the summer of 1972, but large groups of Swainson’s Hawks were not observed in the area at that time. The bulk of the hawks were confined to the Fairfield locale; only four were sighted during searches throughout much of the remainder of the valley. Kleptoparasitism (interspecific food steal- ing or piracy) has been reported for several avian families (see Brockman and Barnard 1979 for a review). Heredia and Clark (1984) summarized the occurrence of this behavior among the raptors, including four buteonine species as participants. Swainson ’s Hawk was excluded. Piracy involving this species and the American Kestrel (Falco sparverius) is reported here. Approximately 34 Swainson’s Hawks and 3 Kestrels were observed in the immediate vicinity of Fairfield on 16 August 1984. Sev- eral of the buteos were soaring; the remainder were perched in trees, on fence posts, and on utility poles. A Kestrel holding an unknown species of small mammal was perched on a utility line some distance from Swainson’s Hawks. It flushed as the vehicle approached and flew a few meters before alighting again on one of the conductors. The procedure was repeated several times. Each flight brought the Kestrel nearer to the perched hawks. Finally, two Swainson’s Hawks left their perch in a nearby tree and flew directly at the WOFFINDEN: SWAINSON’S HAWK 303 Kestrel. After flushing the Kestrel from the line, they actively pursued it for several sec- onds. In response to the harassment, the Kestrel dropped the prey it was holding. One of the pursuing hawks caught the falling prey before it reached the ground, only to be vigor- ously chased by the second. The prey was dropped once more and fell to the ground. Both hawks made a few cursory passes over the site and then flew to nearby fence posts where they remained perched during the du- ration of my visit. The Kestrel returned to the utility line. Brockman and Barnard (1979) suggest that continual raptor interactions at feeding areas may promote piracy. The deliberate actions of the Swainson’s Hawk I observed suggest pre- vious piratical experience. Since the site is prime Kestrel habitat and the Swainson’s Hawks had been residents of the area for sev- eral weeks, the possibility of previous inter- specific interactions was likely. Heredia and Clark (1984) suggest that similar niche overlap may have played a role in the piracy they observed between the Black-shouldered Kite (Elanus caeruleus leucurus ) and White-tailed Hawk (Buteo albicaudatus ). Although klepto- parasitism is a frequently observed avian be- havior, it appears to occur opportunistically and is probably not an important feeding strat- egy (Fuchs 1977, Maxson and Bernstein 1982). I thank Gary B. Jolley for field assistance. D. M. McNair and M. Bechard commented on an earlier draft. Financial support was provided by the University of Pittsburgh at Johnstown. LITERATURE CITED BECHARD, M. J. 1980. Factors affecting nest productivity of Swainson’s Hawks (Buteo swainsoni) in south- eastern Washington. Unpublished dissertation, Washington State University, Pullman, Washing- ton. _ 1983. Food supply and the occurrence of brood reduction in Swainson’s Hawk. Wilson Bull. 95: 233-242. BENT, A. C. 1937. Life histories of North American birds of prey. (Pt.1.) U.S. Natl. Mus. Bull. 167. BROCKMAN, R. J., AND C. J. BARNARD. 1979. Kleptopara- sitism in birds. Anim. Behav. 27: 487-514. Brown, L.. AND D. AMADON. 1968. Eagles, hawks and falcons of the world. McGraw-Hill Book Co., New York. 304 CRUICKSHANK, A. D. 1937. A Swainson’s Hawk migration. Auk 54: 385. DUNKLE, S. W. 1977. Swainson’s Hawks on the Laramie Plains, Wyoming. Auk 94: 65-71. Fitzner, R. E. 1977. Behavioral ecology of Swainson’s Hawk (Buteo swainsoni) in southeastern Washing- ton. Unpublished dissertation, Washington State University, Pullman, Washington. Fox, R. P. 1956. Large Swainson’s Hawk flight in south Texas. Auk 73: 281-282. Fucus, E. 1977. Kleptoparasitism of Sandwich Terns (Sterna sandvicensis) by Black-headed Gulls (Larus ridibundus). Ibis 19: 183-190. HaywakbD, C. L., C. Cotram, A. M. Woopsury, AND H. H. Frost. 1976. Birds of Utah. Great Basin Nat. Mem. No. 1. HEREDIA, B., AND W. S. CLARK. 1984. Kleptoparasitism by White-tailed Hawk (Buteo albicaudatus) on Black-shouldered Kite (Elanus caeruleus leucu- rus) in southern Texas. Raptor Res. 18: 30-31. MARTIN, S. J., AND T. P. MCENEANEY. 1984. Observations of migrating Swainson’s Hawks in Wyoming. Prairie Nat. 16: 62. MAXxsON, S. J., AND N. P. BERNSTEIN. 1982. Kleptopara- sitism by South Polar Skuas on Blue-eyed Shags in Antarctica. Wilson Bull. 94: 269-281. GREAT BASIN NATURALIST Vol. 46, No. 2 Murphy, J. R., F. J. CAMENZIND, D. G. SMITH, AND J. B. WESTON. 1969. Nesting ecology of raptorial birds in central Utah. Brigham Young Univ. Sci. Bull., Biol. Ser. 10(4): 1-36. SHELFORD, V. E. 1963. The ecology of North America. University of Illinois Press, Urbana, Illinois. SEXTON, O. J., AND K. R. Marion. 1974. Probable preda- tion by Swainson’s Hawks on swimming spadefoot toads. Wilson Bull. 86:167—168. SMITH, D. G., AND J. R. Murpuy. 1973. Breeding ecology of raptors in the eastern Great Basin of Utah. Brigham Young Univ. Sci. Bull., Biol. Ser. 28(3): 1-76. TayLor, W. P. 1946. Swainson’s Hawks working on grasshoppers again. Condor. 48: 95. WHITE, C. M. 1966. Notes on the food of the Swainson’s Hawk. Kansas Ornith. Soc. 17:10. WOFFINDEN, N. D., AND J. A. MOSHER. 1979. Ground nest- ing and aggressive behavior by the Swainson’s Hawk. Great Basin Nat. 39: 253-254. WOFFINDEN, N. D., AND J. R. Murpuy. 1977a. A roadside raptor census in the eastern Great Basin— 1973-1974. Raptor Res. 11: 62-65. —__. 1977b. Population dynamics of the Ferruginous Hawk during a prey decline. Great Basin Nat. 37:411-425. _ Refuge, TURKEY VULTURES DECLINE AT A TRADITIONAL ROOSTING SITE Daniel M. Taylor’ ABSTRACT. —A population decline of about 50% from 1974 to 1981 for Turkey Vultures (Cathartes aura) was found at a traditional roosting site at Malheur National Wildlife Refuge in Oregon. This decline may have been due to a region-wide reduction in this species, to the possible but improbable formation of a new but undiscovered roost. or toa reduction in feeding opportunities caused by the decreased use of the refuge by cattle. Concern has been expressed in some re- gions of the United States over population declines of Turkey Vultures (Cathartes aura), resulting in the placement of this species on the National Audubon Society's Blue List twice since 1972 (Tate 1981). Vulture popula- tions appear to be increasing in parts of the east and decreasing in some areas of the west (Tate and Tate 1982), but more quantitative data are needed. Brown (1976) analyzed winter population trends for Turkey and Black Vultures (Cor- agyps atratus) from Christmas Bird Count data from 1950 to 1973 for 16 eastern states, Arizona, Texas, California, and the District of Columbia. He found that Turkey Vultures had declined steadily over this 24-year period in all of these states except California. He attributed the exception of California to large vulture populations from areas censused only in recent years. Since at least 1935, Turkey Vultures have traditionally roosted at the P Ranch Station at the south end of Malheur National Wildlife Harney County, Oregon (Davis 1974). The vultures roost in a row of tall cot- tonwoods (Populus sp.) but use a metal obser- vation tower for pre- and postroosting. They can be accurately censused in both places. In 1973 Davis (1974) censused the vultures at _ P Ranch from late spring to late summer and found a mean of 104 birds per night (range / 68-151: N = 28). There was some seasonal varia- tion, with more birds in the spring and late sum- mer. Her census periods from 21 May to 20 June _ and 21 June to 20 July had high counts of 106 and 110, respectively. There was a mean of 90 birds | from seven counts during these periods. At P ranch I made 16 censuses of Turkey Vultures in 1981 from 23 May to 2 July and 22 counts in 1982 from 25 May to7 July. In 1981 I counted only the peak number of birds on the tower, but the next year I also counted addi- tional birds in the cottonwoods. The mean in 1981 was 46.6 birds (range 35-54), and the following year it was 54.0 (range 33-74) alto- gether, with 49.6 birds (range 33-69) on just the tower. The means of these two years is little more than half that of 1973, and there is almost no overlap between the ranges of the 1973 counts and those of 1981 and 1982. The only count between 1973 and 1981 that I am aware of was one of 92 vultures on 1 July 1976, indicating populations were still com- paratively high that year. There are at least three possible reasons for this population decrease. One is that there has been an overall decrease of vultures in the region. This would support Brown’s earlier findings (1976). Another possibility is the es- tablishment of new roost sites, but no new large roost sites are known in the area, which receives considerable attention from amateur bird watchers. A third explanation for this population decrease is that it is a local decline caused by the cutting back in the number of cattle on the refuge, with a shift to greater winter use. In 1972-73 Animal Unit Months (AUMs) on the refuge were 126,593 (Malheur National Wildlife Refuge, unpublished re- port). This has decreased to 45,000 AUMs in 1981 and 46,000 AUMs in 1982. Since vul- tures are known to scavenge dead cattle (Bent 1937, Burleigh 1972), the reduction of this potential food source might be responsible for the vulture decline. ) ‘Department of Biology, Idaho State University, Pocatello, Idaho 83209. Present Address: 2903 Greenvale Place, Nampa, Idaho 83651. 305 306 Whatever the reason, this summer popula- tion of vultures has evidently suffered a large decrease since 1973. Because this species may be in trouble in general, and many roosts can easily be censused, efforts should be made to do so over the entire range of the vulture to determine population trends. I thank the personnel of Malheur National Wildlife Refuge for providing housing and support while this study was conducted. The study was funded in part by the nongame program of the Oregon Department of Fish and Wildlife. GREAT BASIN NATURALIST Vol. 46, No. 2 LITERATURE CITED BENT, A. C. 1937. Life histories of North American birds of prey (Part I). U.S. Nat Mus. Bull. 167: 1-407. Brown, W. H. 1976. Winter population trends in Black and Turkey vultures. Amer. Birds 30: 909-912. BURLEIGH, T. D. 1972. Birds of Idaho. Caxton Printers, Caldwell, Idaho. xiii + 467 pp. Davis, D. 1974. Roosting behavior of the Turkey Vulture (Cathartes aura). Unpublished thesis, Idaho State University, Pocatello, Idaho. 93 pp. TATE, J., JR. 1981. The blue list: the first ten years. Amer. Birds 35: 3-10. TATE, J., JR., AND D. J. TATE. 1982. The blue list for 1982. Amer. Birds 36: 126-135. BARN OWL DIET INCLUDES MAMMAL SPECIES NEW TO THE ISLAND FAUNA OF THE GREAT SALT LAKE Carl D. Marti! ABSTRACT. —An investigation of the diet of the Common Barn-owl (Tyto alba) on Antelope Island, Great Salt Lake, Utah, yielded four mammal species not previously known to occur on any island in the Great Salt Lake (Microtus pennsylvanicus, M. montanus, Ondatra zibethicus, and a Sorex sp.). Two other species, known from other islands, were added to the list of fauna of Antelope Island (Perognathus parvus and Reithrodontomys megalotis ). The barn owl diet on Antelope Island was remarkably like that of barn owls feeding in farmlands adjacent to the Great Salt Lake despite major vegetational differences. Relatively little collecting for small mam- mals has occurred on islands in the Great Salt Lake, Utah; the remoteness of several islands and private ownership of others have discour- aged a thorough examination of their mam- malian fauna. Bowers (1983) compiled a list of nonvolant, native mammal species known to occur on these islands from previously pub- lished studies. In this paper I document the occurrence of four mammal species previously unreported on any island in the Great Salt Lake and add two additional species to the list for Antelope Island. These mammals were identified among prey remains in pellets of the Common Barn-owl (Tyto alba) nesting on Antelope Is- land. Barn owls have taken certain small mammals in several other localities before mammalogists were aware of the existence of _ the mammals there (e.g., Stickel and Stickel _ 1948, Twente and Baker 1951). also compare the island diet with prey eaten by barn owls _ feeding in farmlands adjacent to the Great Salt | Lake. METHODS Regurgitated pellets were collected from a _ barn ow] nest site at the Dooley Ranch on the east-central shore of Antelope Island, Davis County, Utah. The nest was in an abandoned agricultural silo. Pellets were gathered once ’ or twice a year in spring or summer from 1980 {through 1984. I documented nesting at the ‘collection site from 1981 through 1984 and ———— ‘Department of Zoology, Weber State College, Ogden, Utah 84408. believe that nesting probably occurred in 1980 as well. Thus, the material consisted of prey of pairs of owls and their young. I treated pellets with a sodium hydroxide solution to dissolve hair and feathers. Bones were identified and quantified by standard procedures (Marti 1974). RESULTS AND DISCUSSION The barn owl diet on Antelope Island was typical of the foods of this species elsewhere (Wallace 1948, Glue 1967, Marti 1974), being heavily dominated by mammalian prey (98.4%, Table 1). Three rodents in the diet, meadow vole (Microtus pennsylvanicus), montane vole (M. montanus), and muskrat (Ondatra zibethicus), had not previously been known to occur on any island in the Great Salt Lake, nor had shrews (Sorex sp.). The shrews were most likely S. vagrans, which occurs on the adjacent lake shore (Dur- rant 1952). The Great Basin pocket mouse (Perognathus parvus) and the western har- vest mouse (Reithrodontomys megalotis), which also occurred in the owl diet, had not been reported previously for Antelope Island but were known from other islands in the Great Salt Lake (Bowers 1983). Even though little is known about small mammal populations on these islands, several apparent anomalies existed between the owl diet and probable prey abundance and distri- bution. The heavy domination by voles was the most surprising aspect of the barn owl 307 308 GREAT BASIN NATURALIST Vol. 46, No. 2 TABLE 1. Prey of Common Barn-Owls on Antelope Island, Great Salt Lake, Utah. Percent of prey numbers Totals Prey 1980 1981 1982 1983 1984 No. Percent Sorex sp. 2.0 2.3 3.6 2.8 3.6 73 2.7 Sylvilagus nuttallii (juv.) 0.1 — — — — 1 tr. Perognathus parvus 0.5 — — — — 6 0.2 Dipodomys ordii 0.2 — — — 0.3 5 0.2 Reithrodontomys megalotis 9.5 19.2 11.9 3.3 5.9 246 9.0 Peromyscus maniculatus 5.0 4.3 Ll 2.5 6.6 124 4.5 Neotoma lepida 0.1 — — — — 1 tr. Microtus pennsylvanicus 55.6 49.8 62.8 70.7 61.7 1617 59.2 Microtus montanus 25.3 20.8 16.6 16.8 19.7 590 21.6 Ondatra zibethicus (juv.) 0.2 — — — — 2 0.1 Mus musculus 0.5 Dall 1.4 0.2 0.5 21 0.8 Birds Porzana carolina -= 0.4 — — — 1 tr. Sturnus vulgaris — _ — 0.5 0.3 4 0.1 Unidentified bird 0.8 0.4 2.5 3.0 0.7 34 1.2 Insects Unidentified coleopteran 0.2 — — — 0.5 5 0.2 Totals 1231 255 NOT 393 574 2730 100.0 *tr = <0.01% diet. Habitat suitable for voles was very lim- ited on Antelope Island; dense vegetation usually associated with them occurred only in a narrow band at a few places along the lake shore, around a few springs, and in an irri- gated hay field. The owls must have concen- trated their foraging efforts in those limited areas. The deer mouse (Peromyscus manicu- latus) is the most abundant small mammal on the islands in the Great Salt Lake (Neil Jensen, personal communication), but they ranked only a distant fourth in species abun- dance in the owl diet. The valley pocket go- pher (Thomomys bottae) was the only small nocturnal mammal known to exist on the is- land not taken by the owls. My continuing, long-term study of barn owl ecology in Box Elder, Weber, and Davis counties, Utah (Marti unpublished data) over- lapped with the years reported here for the Antelope Island material. Data from the long- term study, being done in agricultural land, provides some interesting comparisons for the Antelope Island data. Antelope Island and the main study area are 27 km apart at the closest point. Major differences between the two sites are in vegetation and topography. The mainland study area is essentially flat, and irrigated crops and pastures replace native vegetation. Antelope Island has much more topographic relief, rising from about 1,260 m to 1,999 m, and is covered almost entirely with vegetation typical of Great Basin deserts. Despite these differences, the same five prey genera (Microtus, Peromyscus, Reithrodonto- mys, Mus, and Sorex) contributed 97% of the diet in both areas. Only minor differences in the order of species abundance occurred be- tween diets in the two areas. Meadow voles and montane voles were first and second in both, whereas deer mice and harvest mice exchanged places (third and fourth), as did house mice (Mus musculus) and shrews (fifth and sixth) between island and mainland. Year- by-year comparisons between island and mainland diets showed that shrews and house mice were taken less frequently in every year on the island than on the mainland. Deer mice occurred less frequently in the island diet for four of five years, but the reverse was true for harvest mice. The two vole species combined were more frequent prey on the island in three of five years and overall (80.8% versus 77.5%); meadow voles were taken at a higher frequency in every year of the island than on the mainland, but the opposite was true for montane voles. The desert woodrat (Neotoma lepida) and Ord’s kangaroo rat (Dipodomys ordii) were in the island but not the mainland _ diet. Habitat for these has been eliminated by irrigation agriculture in the mainland study area. Two other species, the Norway rat (Rat- tus norvegicus) and pocket gophers, were eaten by owls on the mainland but not on the April 1986 island. Norway rats probably do not occur on the island, which has had very little human habitation, but pocket gophers are found there. I cannot offer a satisfactory explanation of why island barn owls did not appear to capture gophers. It is presumed that the Antelope Island barn owls foraged entirely on the island and, thus, that their diet reflected prey species found there. I base this on investigations of barn owl foraging ranges done elsewhere and on the tenets of central-place foraging theory. Orians and Pearson (1979) contended that ani- mals should reduce costs of obtaining food as much as possible. One way to do this is to forage close to the nest site (central place) and reduce traveling time. For the Antelope Is- land barn owls this means foraging on the island and not crossing over to the mainland to hunt. Minimum distance from the island col- lection site (nest/roost) to the mainland was more than 10 km during 1980-1984. Hegdal and Blaskiewicz (1984) found a maximum dis- tance from roost to hunting areas of 5.6 km in radio-tagged barn owls. Note that these comparisons are between the diet from one collection site each year on the island and 26-31 sites on the mainland. The sample size from the mainland was much larger (n = 41,453) and represented year- round prey data whereas the island sample was mainly from late winter through summer. Although these differences could affect com- _ parisons between diets from the two sites, it seems unlikely that they would cause major _ misconceptions. In conclusion, barn owls selected very simi- lar prey on Antelope Island and in agricultural lands adjacent to the Great Salt Lake despite the very different vegetation in the two MARTI: BARN OWL DIET 309 places. Dietary evidence indicated that the owls concentrated their foraging efforts in habitat suitable to voles in both localities. This type of habitat is abundant and widespread in the mainland study area but limited and con- centrated on Antelope Island. ACKNOWLEDGMENTS I thank the following employees of the Utah Division of Wildlife Resources for help in gaining access to Antelope Island and in col- lecting pellets: Phillip Wagner, Don Paul, and John Kimball. LITERATURE CITED Bowers, M. A. 1983. Insular biogeography of mammals in the Great Salt Lake. Great Basin Nat. 42: 589-596. Durrant, S. D. 1952. Mammals of Utah: taxonomy and distribution. Natural History Museum, Univ. Kansas Publ., Lawrence, Kansas. GLUE, D. E. 1967. Prey taken by the barn owl in England and Wales. Bird Study 14: 169-183. HEGDAL, P. L., AND R. W. BLASKIEWICZ. 1984. Evaluation of the potential hazard to barn owls of talon (Brodi- facoum bait) used to control rats and house mice. Environ. Toxicol. Chem. 3: 167-179. Martl, C. D. 1974. Feeding ecology of four sympatric owls. Condor 76: 45-61. ORIANS, G. H., ANDN. E. PEARSON. 1979. On the theory of central place foraging. Pages 155-177 in D. J. Horn, G. R. Stairs, and R. D. Mitchell, eds., Analysis of ecological systems. Ohio State Univer- sity Press, Columbus, Ohio. STICKEL, W. H., AND L. F. STICKEL. 1948. Mammals of northwestern Texas found in barn owl pellets. J. Mammal. 29: 291-293. TWENTE, J. W., AND R. H. Baker. 1951. New records of mammals from barn owl pellets. J. Mammal. 32: 120-121. WALLACE, G. J. 1948. The barn owl in Michigan. Mich. Agric. Expt. Sta. Tech. Bull. 208. DISTRIBUTIONAL STUDY OF THE ZION SNAIL, PHYSA ZIONIS , ZION NATIONAL PARK, UTAH F 1,2 David Ng’ and James R. Barnes! ABSTRACT. —The major hanging gardens and associated water seeps in Zion National Park were surveyed for the presence of the Zion Snail (Physa zionis ). Environmental parameters, including water depth, water velocity, substrate slope, and algal cover, were measured to determine their effect on the local distribution of the snail. Large populations (densities 125 to 250/m*) were found in the Virgin River Narrows area of the park and at a hanging garden and seep located 1.0 km north of Scout Lookout. Densities in other localities were low in comparison. Snails were not found in all hanging gardens or seeps. The major factor controlling within seep distribution was determined to be water velocity. Experiments were conducted to test the ability of the snail to remain attached during differing water flows. The snail showed an ability to remain attached during high flows, but few snails were found in areas of high flow. The Zion Snail, Physa (Petrophsa) zionis Pilsbry (1926), an endemic to Zion National Park, Washington County, Utah, is found in water seeps and associated hanging gardens on the canyon walls along the Virgin River Narrows region of the park. The limpetlike characteristics of P. zionis, a reduced shell spire, enlarged last whorl, and large foot, were reported as being adaptations for attach- ment on vertical surfaces of seeps (Pilsbry 1926, Chamberlain and Jones 1929). There is no literature on the ecology or natural history of the Zion Snail other than the mention of a snail-algae association and the hypothesis that P. zionis evolved from a common Physa type (Woodbury 1933, Talmadge 1970). The purposes of this study were: (1) to sur- vey and determine the distribution and popu- lation density of the Zion Snail within Zion National Park, (2) to determine the impor- tance of specific habitat variables which may limit snail distribution within seeps, and (3) to test the hypothesis that the limpetlike mor- phological adaptations provide the snail with greater substrate attachment abilities in the seep environment. STUDY AREA Zion National Park, in southwestern Utah, is in an area of sandstone formations cut by the Virgin River and its tributaries. Seeps and hanging gardens that offer suitable habitat for ‘Department of Zoology, Brigham Young University, Provo, Utah 84602. “Department of Zoology, University of Texas, Austin, Texas 78712. the Zion Snail are found at the junction of the Navajo and Kayenta sandstone layers. Seeps are formed when water percolating through the porous Navajo layer contacts the impervi- ous underlying Kayenta formation and flows laterally to the canyon walls (Welsh and Toft 1981). The Navajo-Kayenta junction is at dif- ferent elevations along the canyon wall in rela- tion to the Virgin River. In the Main Canyon area of the park, the river has cut the deepest and seeps are farthest removed from the river. In the Narrows the river is near the Navajo- Kayenta junction elevation, and many of the seeps are close to the river. Malanson (1978, and 1980) describes the plant communities for some hanging gardens in the Narrows region. Seeps in Zion National Park are located in three regions. They are the Narrows, the Gateway to the Narrows, and Main Canyon. The Narrows, including Orderville Canyon, is a narrow box canyon where vertical canyon walls form the channel of the Virgin River. The canyon walls are less than 10 m apart in some areas. Seeps in the Narrows are mainly simple dripline and window-blind types (Welsh and Toft 1981), though a few low al- coves have been carved out by the river. All seeps have varying amounts of calcium car- bonate precipitate and algae. A film of water usually less than 1 mm in depth flows over the vertical and horizontal surfaces. Clinging herbaceous plants and mosses are common, indicating the presence of relatively stable or | 310 April 1986 NG, BARNES: SNAIL ECOLOGY Silat TABLE 1. Classes of ecological parameters used to analyze local distributional patterns within seeps. Parameters 1 2 Depth (mm) 0-1 2-3 Flow (mm) 1 Slope (degrees) 0-10 11-45 Algae (% cover) 0-12.5 12.5—25 Classes of parameters 3 4 5 4-10 >10 >45 25-50 50-75 >75 old seeps (Malanson 1980). New flows without vegetation are also present. The Gateway to the Narrows region, imme- diately downstream from the Narrows, is a more open area that receives heavy visitor use. The Gateway is approximately 1.5 km long, with a paved trail along the east canyon wall. Seeps in this section are large and of the window-blind and alcove types (Welsh and Toft 1981). Plant communities are well devel- oped, with many herbaceous and woody plants present. Calcium carbonate precipita- tion is extensive, with deposits several cen- timeters in depth common. Seeps in this sec- tion are isolated from the Virgin River. The Main Canyon, which extends down- stream from the Gateway, widens consider- ably. Seeps and gardens are large alcoves, alcove-plunge basins, or terraces (Welsh and Toft 1981). Water flow in these seeps is gener- ally high, with depths of several millimeters measured. Woody plants and pockets of soil are present in most seeps. Most seeps are connected to the Virgin River by small (< 2.0 +m wide) streams with rock-gravel-sand sub- _ Strates. A terraced seep-spring area approximately 1 km north of Scout Lookout in the Main Canyon was used to study the ecological vari- ables that may limit local distribution and at- ‘tachment abilities of the snail. This seep is 10 +m wide, with two small springs at the south end. The terraces receive surface flow from areas high on the canyon wall, whereas the "springs feed into a small stream. The stream flows over a 5 m waterfall, is joined by flow | | from the terraces, and then flows for approxi- mately 30 m into the Virgin River. Water {temperature in the springs is relatively con- «stant (19-21 C), but the terrace water temper- sature fluctuates with the ambient air tempera- ture. Moss, algae, and herbaceous plants are present. The substrate varies from thick cal- cium carbonate deposits on the sandstone wall | and terraces to the cobble-gravel-sand stream bottom. Water depth and velocity in the stream range from 0 to > 200 mm and 0 to > 1.0 m/sec, respectively. METHODS The major seeps in Main Canyon and Gate- way to the Narrows were surveyed during the summers of 1981 and 1982. The Narrows was surveyed only in July 1982 because of the limited time it was accessible. The terraced seep-spring north of Scout Lookout in Main Canyon was sampled 24 October 1981 and 2 April, 24 April, 8 June, and 16 July 1982. Detailed quantitative work in the Gateway region was impractical because of heavy visi- tor use. A quadrat (10 x 15 cm) was used to estimate population density. Four variables, including water depth, water velocity, substrate slope, and algal cover, were measured in each quadrat. Quadrats were placed every 30 cm apart along random transects across the water course. The number of transects and quadrats required to sample a seep or spring area varied with changes in the amount of flowing water. Water velocity (force of flow) was de- termined by measuring in millimeters the amount of water forced vertically into a piece of 2 mm diameter plastic tubing. Substrate slope was measured to the nearest 5 degrees. Snail size was measured to the nearest 0.5 mm in situ. Sampling was limited to seeps accessi- ble without climbing equipment. Statistical analysis of the measured vari- ables was accomplished by using the Chi- square analysis program in MINITAB (Ryan etal. 1981) and the ANOVA program in RUM- MAGE (Scott et al. 1982). For analysis, the measurements were divided into classes (Table 1). To test the attachment ability of the snail, snails were subjected to various flows in a 3) 1 40 A 20 > 40 B {S) P4 5 G 20 uJ [aa re x 40 Cc 20 10 20) so Ga Be) G0). “KO SIZE CLASSES (MM) Fig. 1. Size distribution of the Zion Snail found in the Narrows area of Zion National Park. A and B are from seeps in the Main Narrows; C is from Orderville Canyon. plastic channel and on natural substrate. In the plastic-channel test, snails were placed at the end of a 1 m long plexiglass channel in which water from the stream was channeled through in increasing amounts. In the natural- substrate wash experiment, two volumes of water (5 and 10 liters) were poured down a 15 cm wide plexiglass channel at 5 and 10 second duration onto snails in situ. The channel was held at 30 degrees. For comparison, the more common stream snail, Physa gyrina, was tested in the laboratory. RESULTS Population Density and Distribution Our surveys show the Zion Snail occurring from a seep-spring stream area 1 km north of Scout Lookout into Orderville Canyon. The Narrows area above Orderville Canyon was not surveyed in this study. Zion Snails were not found in seeps on the west side of the Virgin River nor south of the seep-spring stream area above Scout Lookout in the Main Canyon area. NARROWS.—Six seeps examined in this re- gion contained snails. All snails were found in dripline or window-blind type seeps where vegetation was abundant. Many snails were observed less than 10 cm above the Virgin GREAT BASIN NATURALIST Vol. 46, No. 2 River. Snails were not found on the northwest canyon wall. Of the six seeps that contained snails, three were sampled quantitatively, two in the main part of the Narrows and one in Orderville Canyon. The first seep was 4 m wide, located approximately 0.3 km into the Narrows, and had a density of 30 snails/m’. The second was 6 m wide, located 0.5 km into the Narrows, and had approximately 100 snails/m*. The seep in Orderville Canyon is a long, continuous dripline with approximately 130 snails/m’. The other three seeps that con- tained snails were in the main part of the Narrows. These seeps were all less than 3 m wide and had estimated densities of < 20 snails/m*. The largest snails, 6.5 to 7.0 mm in length, were found in the main Narrows. Ay- erage length ranged from 3.0 to 6.0 mm (Fig. 1). Based on the above densities, 3,000 snails is a conservative estimate of the minimum population in the Narrows region. GATEWAY.—This area was only qualita- tively sampled because the large number of park visitors using the trail made quantitative sampling impractical. Very few seeps con- tained snails, and even in large or continuous seeps the snails were found in discrete patches. Densities of 5 to 10 snails/m* were common, with 20 to 30 snails/m? maximum. Snail sizes throughout this area were 3 to 4 mm at the time of sampling. MAIN CANYON.—The only snails found in the Main Canyon area were in a terraced seep-spring 1.0 km north of Scout Lookout. Three in-seep snail distributional patterns were evident: (1) on the terraces of the seep; (2) at the head of the northern-most spring; and (3) in the main stream below the waterfall | down to the confluence with the Virgin River. Snails were not found in the southern spring. Densities ranged from 40/m’ in the stream to | more than 250/m° in the spring. However, because of the small surface area of the spring, the total number of snails there is less than | 1,000. The highest densities were found in spring season with the appearance of young snails. In the terrace region the density was around 175/m*. The size ranged from < 1.0 | mm (immatures) to 4.0 mm. Snails > 5.0 mm | were rare in this area. There was a marked | decrease of snails > 3.5 mm after June. The | total population in this area apparently ranged - from 2,000 to 6,000 snails. April 1986 NG, BARNES: SNAIL ECOLOGY 313 TABLE 2. Results of natural substrate wash experiment using in situ Physa zionis. Duration Volume Rate of Number of snails (Seconds) (Liters) flow Replicates Start Finish Percent remaining 5 5 1Vsec 1 16 13 81.25 2 11 8 De, 3 12 11 91.66 4 13 11 84.62 10 2 Vsec 1 10 10 100.00 2 10 vil 70.00 3 10 U 70.00 4 12 9 75.00 10 5) 0.5 l/sec 1 12 12 100.00 2 12 12 100.00 3 13 od 53.85 4 13 12 92.31 10 1 V/sec 1 12 10 83.33 9 11 9 81.81 3 11 9 81.81 4 14 12 85.71 Wash Experiments In the preliminary plastic channel test, eight snails were used. At the initial flow of water through the channel, two snails were lost; the others remained attached up to flows of 280 mm in the measuring tube, the maxi- mum achievable using water available in the field. Size did not appear to be a factor in attachment ability, since the largest (5 mm) and the smallest (1.5 mm) were among those remaining. In the natural-substrate wash experiment, 78% of the snails remained attached after the highest flow (10 liters for 5 sec = 2 I/sec), and 86% remained after the lowest flow (5 liters for 10 sec = 0.5 I/sec). With flows of 1 I/sec, 83% remained after 5 and 10 sec. durations (Table 2). Snails became detached when the carbon- ate or sandstone substrate they were on _ washed away. Physa gyrina, a coiled shell gastropod that - was common in the area, was tested in the laboratory for comparison with the attach- ment ability of the Zion Snail. In the prelimi- nary plastic channel test, only two snails re- mained after the initial flow of water. At < 75 - mm of flow in the measuring tube, the shells of the remaining snails started to lift from the | foot and soon pulled the foot off the channel. | | | | | | No snails remained in the plastic channel after flows of > 75 mm. In the controlled wash - experiment, only 2 of 16 and 1 of 12 snails _ remained at 5 liters and 10 seconds (0.5 I/sec), the lowest flow possible. A third replicate with 12 snails left no snails remaining, and the experiment was stopped. Within Seep Distribution The seep-spring area was chosen for the analysis of factors that influence local distribu- tions. Snails were found in 126 of 262 quadrats (48.1%). Using a Chi-square test of indepen- dence, snail distribution was found to be inde- pendent of slope and algae cover while depen- dent on depth and flow (Table 3). Flow class 1 (<1 mm) accounted for 125 of the 126 quadrats (99.2%) that contained snails. The proportion of quadrats that contained snails also decreased with increasing water depth. The single quadrat with a snail in high flow (> 1 mm) is not significantly different from 0 (Table 4). In an attempt to determine the overall ef- fects of the variables and their interactions on snail distribution, the ANOVA model Y (snails) = D (depth) + V (velocity) + S (slope) + DV + DS + VS +E was used in weighted least squares analysis in RUMMAGE. Algal cover was not used because it was not significant and the five cover classes were contributing too many missing cells. The data used were from a single sampling date and from a specific area in the seep-spring to mini- mize effects of unaccounted factors. The three factors plus interactions accounted for 19% of the variation (R? = 0.193, R’ adj. for df = 0.101). The terms water depth and velocity were significant at p < 0.05 (Table 5). 314 TABLE 3. Chi-square analysis of independence to de- termine the significance of water depth, water flow (ve- locity), substrate slope, and algal cover in determining the distribution of the Zion Snail in the seep-spring area. Variable x df Significance Depth* 33.78 2 as Flow 19.33 1 are Slope 0.50 2 N.S. Algae 2.10 4 N.S. *Depth class 4 was added to 3 since less than five samples were taken at depths greater than 10 mm. **p < 0.05 DISCUSSION The highest population densities were found in Orderville Canyon and the seep- spring. Both areas had densities exceeding 100/m*. Early accounts of the snail did not include density estimates; consequently, his- toric comparisons cannot be made. However, Gregg (1940) described “Fairy Land” 3/4 mi south of Sinawava as a terraced region with numerous snails on horizontal surfaces. From his description and the location, Fairy Land is possibly the seep-spring area we studied. Pils- bry (1926) collected the type specimens at the Narrows mouth and found no Zion Snails above three miles into the Narrows. Because of accessibility, Zion Snails were often col- lected at the mouth of the Narrows. Wood- bury (cited in Chamberlain and Jones 1929: 167) reported that cliffs (seeps) near the type locality and beyond had been “stripped of snails by collectors on previous occasions, but that in a few days migration from above had soon renewed the supply.” The migration re- ported is most likely of snails from higher, inaccessible parts of the seep to the lower regions visited by collectors. Presently, the lower, accessible seeps in the Gateway and the mouth of the Narrows contribute only a small percentage to the total number of Zion Snails in the park. The distribution of the Zion Snail is patchy. They occur at differing distances in reference to the Virgin River, with various distances between subpopulations. The only snails found in deep water were those in the stream below the waterfall at the seep-spring area. However, density in the stream was very low and most likely was the result of snails being washed down from the seep above. GREAT BASIN NATURALIST Vol. 46, No. 2 TABLE 4. Number of quadrats and occurrence of snails within various water depth and flow categories. Parameter® # with Snails and category N (% + 95% C. 1.) Depthiaeeel 142 90 (63.4 + 7.9) 2 74 287167287211) 3 46 8 (17.4 + 11.0) Flow ] 239 125 (52.3 + 6.3) 2 23 L( ABs Sosy “See Table | for description. Depth class 4 was added to 3 since less than five samples were taken at depths greater than 10 mm. “Not significantly different from 0 at p < 0.05. The silt-laden Virgin River, despite being the physical factor that joins most of the seeps together, may be a barrier to snail movement and contributes to the patchy distribution. The absence of snails in the river s main chan- nel may indicate an avoidance of the silt or an inability to survive there. However, the river may still be a major avenue of dispersal. Some snails in the Narrows occur very close to the river and may be carried downstream during highwater or flooding. These snails are cer- tainly. covered during periods of high flow. If the snail can survive in the river for a short period of time, the Narrows may be a source of its origin for downstream areas. The seeps along the Gateway that are presently isolated from the river may have been colonized dur- ing extreme flooding or by other vectors. The extreme patchiness may be a reflection of rare colonization events and local extinction. Within seeps, the snail distribution is patchy. Water depth and velocity are viewed as limiting factors on a gross scale. Snails are found in decreasing frequencies with increas- ing water depth and are not found at all in areas of high flow (Table 4). However, as shown by the ANOVA model, depth and wa- ter velocity only accounted for 10% of the variation when adjusted for df. Even in areas of favorable depth and velocity, much of the distributional variation is unexplained by the measured factors. The localized patchiness may be linked with food sources, variations in water flow, and the movement of snails that were not measured. As shown by the flow experiments, the Zion Snail is suited for living in water flows much higher than those normally encountered in the seeps. The attachment abilities are also much greater than Physa gyrina, which has a April 1986 NG, BARNES: SNAIL ECOLOGY 315 TABLE 5. Results of RUMMAGE ANOVA testing occurrence of snails with environmental variables Depth (D), Flow (F) and Slope (S). (N = 262). Source df ss D 3 31.70 F 1 24.52 S 2 0.72 DF oF 8.80 DS 6 15.25 FS 2, 8.62 Error 141 345.05 R? = 0.193 R? adjusted for df = 0.101 *dflost due to missing cell. higher, coiled shell. The early literature (Pilsbry 1926, Woodbury 1929, Chamberlain and Jones 1929) states that the purpose of the limpetlike shell and large foot is to allow the snail to live on the vertical surfaces in the trickling water. How- ever, most snails are not found in visibly moving water and are not restricted to vertical surfaces. Slope was not a significant factor (Table 5), and snails are found on horizontal surfaces. The hypothesis (Woodbury 1933) that the large foot and reduced spire evolved from a more typical physoid type snail as a result of high flows or floods in the seep environment is probably correct. We feel the selective value of attach- ment during high flows is quite significant. Seeps near the river are occasionally flooded (Malanson 1978), and the ability to remain at- tached during floods would be an advantage. Based on the experiments, the higher, coiled shell of a typical snail would have caused it to be washed away even in a minor flood, whereas a majority of the Zion Snails would have remained attached to the seep. The adaptive value to the Zion Snail of a large foot and limpetlike shell is, in our opinion, to remain attached during peri- ods of fast flows and flooding, allowing the snail to remain in and exploit seep habitats. SUMMARY This research, along with past studies on the Zion Snail, provides a basic understanding of its general distribution and habitat use. Future studies should be done to determine the distri- bution further up the Narrows and in Orderville Canyon; the role of temperature in controlling reproduction, food habits, and mortality factors; the role of drying seeps on distribution; and, fiinally, the effect of floods on the Narrows and Orderville Canyon populations. Some of these proposed studies would necessitate collecting MS F Significance 10.58 4.32 0.006 24.52 10.02 0.002 0.36 0.15 0.863 4.4 1.80 0.169 2.54 1.04 0.403 4.31 1.76 0.175 2.45 adult snails and raising them in the laboratory, but others could only be accomplished after long-term field observations. Lastly, it must be recognized that we know nothing of populations in gardens high up on the canyon wall or if the snail even exists in those gardens. Although the snail is endemic to Zion National Park, there are no large populations, but the populations we studied contained sufficient numbers for reproduction to take place. The Zion Snail has probably never existed in large numbers and, in comparison to other snails, it may be considered rare. LITERATURE CITED CHAMBERLAIN, R. V., AND D. T. JONEs. 1929. A descriptive catalog of the Mollusca of Utah. Bulletin of the University of Utah 19: 1-203. Grecc, W. O. 1940. Mollusca of Zion National Park, Utah. Nautilus 54: 30-32. MALANSON, G. P. 1978. Distribution of plant species in the hanging gardens of the Narrows, Zion National Park, Utah. Unpublished thesis, University of Utah, Salt Lake City. Pitspry, H. A. 1926. A fresh-water snail, Physa zionis, living under unusual conditions. Proceedings of the Academy of Natural Sciences of Philadelphia 77: 325-328. T. A, B. L. JOINER, AND B. F. Ryan. 1981. MINITAB, Minitab reference manual. MINITAB Project, Statistics Department, Pennsylvania State University, University Park, Pennsylvania. D. T., M. W. CarTER, AND G. R. Bryce. 1982. RUMMAGE. Statistics Department, Brigham Young University, Provo, Utah. TaLmapceE, R. R. 1970. How far ecology? Of Sea and Shore 1: 131. WELSH, S. L., AND C. A. Tort. 1981. Biotic communities of hanging gardens in southeastern Utah. National Geographic Society Research Reports 13: 663— 681. Wooppsury, A. M. 1929. The snails of Zion National Park. Nautilus 43: 54-61. ____. 1933. Biotic relationships of Zion Canyon, Utah, with special reference to succession. Ecological Monographs 3: 147-245. RYAN, SCOTT, BLOCKAGE AND RECOVERY OF NITRIFICATION IN SOILS EXPOSED TO ACETYLENE! Steven J. Burns”? and Richard E. Terry” ABSTRACT.—Acetylene gas is very useful in laboratory and in situ assay procedures for nitrogen fixation and denitrification. There is concern, however, that measurements of denitrification may be underestimated because nitrification, a major source of nitrate, is inhibited by C,H). The objective of this study was to examine the effects of C,H, partial pressure and length of exposure time on nitrification in soils. Acetylene partial pressures of 0.1, 1.0, and 10.0 kPa were found to effectively inhibit nitrification in soil samples incubated in the laboratory. Both the partial pressure of C,H, and the length of exposure time were found to affect the recovery time of nitrification in soil samples. Nitrification recovered within seven days in samples exposed to 0.1 and 1.0 kPa C,H, for only 24 hours. The recovery of nitrification in samples exposed to 10.0 kPa CH, for 24 hours or to 0. 1 and 1.0 kPa C,H, for 216 hours was delayed for an additional seven days, however. Acetylene gas has been effectively used in laboratory and in situ techniques for the mea- surement of both nitrogen fixation and deni- trification. The use of acetylene (C,H,) is at- tractive due to the low cost and _ high sensitivity of the procedures. For N, fixation studies, C,H, is used to saturate enzymes re- sponsible for fixation; in the process, C,H, is reduced to ethylene (C,H,). Ethylene can be readily detected by gas chromatography and N, fixation rates estimated (Hardy et al. 1968, Bergerson 1980). Acetylene has an inhibitory effect on the bacterial enzymes that reduce N,O to N,; therefore, C,H, has been used in denitrification studies (Balderson et al. 1976, Yoshinari et al. 1977). In the presence of C,H, partial pressures greater than 0.1 kPa, N,O is the sole gaseous product of denitrification and is detectable by gas chromatography. Concerns have been expressed about the use of C,H, in denitrification studies, because C,H, has been found to be an effective nitrifi- cation inhibitor at a partial pressure of 0.01 kPa (Walter et al. 1979, Berg et al. 1982). In some situations denitrification measurements may be affected by C,H,, because nitrifica- tion, amajor source of NO, , is inhibited. This is not a problem with short-term laboratory incubations or experiments that involve the addition of nitrate to the soil, but it may be inappropriate to use acetylene in experiments where NH,’ is used as the starting point in deni- trification studies or in long-term experiments where mineralization and subsequent nitrifica- tion could be expected to produce significant nitrate. Researchers have studied nitrification in the presence of C,H, partial pressures ranging from 1,000 to 0.01 Pa (Walter et al. 1979, Berg et al. 1982). The minimum effective partial pressure that inhibited nitrification was 10 Pa. However, partial inhibition was found at 0.1 Pa (Berg et al. 1982). These researchers reported that the in- hibitory effects of the low C,H, levels (1,000 to 10 Pa) ceased within 7 to 10 days of removal of GH Acetylene partial pressures of 10 kPa are rou- tinely used in nitrogen fixation studies (Hardy et al. 1968, Bergerson 1980) and in laboratory deni- trification studies (Yoshinari et al. 1977, Terry and Tate 1980a, 1980b, Terry et al. 1981). Field measurement of denitrification by the C,H, inhi- bition technique involves introduction of C,H, to the soil atmosphere through perforated tub- ing or C,H, treated irrigation water (Ryden, I., Laboratory evaluation, 1979; Ryden, II., Devel- opment and application, 1979, Rolston et al. 1982, Hallmark and Terry 1985). It is likely, with these procedures, that C,H, partial pressures in excess of 10 kPa will exist in portions of the soil atmosphere that will maintain minimum effec- tive levels of 0.1 kPa throughout the soil atmo- sphere. 'This publication is based on work supported in part by funds provided by the Bureau of Land Management. “Department of Agronomy and Horticulture, Brigham Young University, Provo, Utah 84602. 5Present address: Department of Agronomy, University of Missouri, Columbia, Missouri 65201. 316 | ) : ) | April 1986 BURNS, TERRY: NITRIFICATION IN SOILS TABLE 1. The properties of soils used in this investigation. Soil series pH Sand Timpanogos 7.9 348 Woodrow 8.1 239 The effects of C,H, partial pressures greater than 1 kPa on nitrification and the resumption of nitrification following exposure have not been studied. The objective of this research was to determine the effects of various C,H, partial pressures and length of C,H, exposure time on nitrification in soils. MATERIALS AND METHODS Soils used in this investigation were Tim- panogos clay loam (a fine, loamy, mixed mesic Calcic Argixeroll) collected at the Brigham Young University Agriculture Station near Spanish Fork, Utah, and Woodrow clay loam (a fine silty, mixed mesic Xeric Torrifluvent), collected at Camp Floyd State Park, Fairfield, Utah. The properties of the soils used in this study are presented in Table 1. Air-dried soil samples were analyzed for total N by the mi- cro-Kjeldahl method of Bremner (1965) and for organic C by the method of Allison (1965). The soil pH was measured by glass electrode on a 1:1 soil:water ratio. Duplicate moist soil samples, the equiva- lent of 10.0 g dry weight, were placed into 120 ml serum bottles and sealed with septum stoppers. The samples were brought to ap- proximately —0.333 MPa matric potential by addition of 1.25 ml of a solution containing 3.96 g (NH,), SO, L ’ to equal an application rate of 100 mg N kg * soil. The soils were preincubated for 24 hours before treatment with C,H,. To compare the inhibitory effects of various levels of C,H, with those of the commercial nitrification inhibitor, nitrapyrin, soil samples were continuously exposed to C,H, partial pressures of 0.1, 1.0 or 10.0 kPa. Impurities were removed from C,H, by passage of the gas through concentrated H,SO, and water traps (Hardy et al. 1968). Nitrapyrin was dissolved in the ammonium solution and added to the soil samples at the rate of 2 mg kg '. The continuously exposed samples were unsealed, aerated for 10 minutes once a week, then Sy Silt Clay Organic-C Total-N gkg 309 343 1229 1.27 390 375 8.8 1.40 reexposed to acetylene, thus eliminating anae- robic conditions. Nitrapyrin samples were also aerated for 10 minutes once a week. To examine the effects of various partial pressures of acetylene on nitrification and the recovery of nitrification following exposure, acetylene was added at rates of 0, 0.1, 1.0, and 10.0 kPa for a period of 24 hours and then removed by flushing the incubation vessels with air. The samples were then incubated at 21 C and continuously aerated with laboratory air at 100% relative humidity using a manifold delivery system and an aquarium air pump. The air flow rate for each sample was 0.5 mL Sie To determine the effects of length of acetylene exposure time on nitrification, soil samples were exposed to 0.1, 1.0, or 10.0 kPa C,H, for 24 or 216 hours. Following exposure, acetylene was removed by flushing with air and the samples were then continuously aer- ated. Sufficient samples were prepared to allow for duplicate analyses at 0, 7, 14, 21, and 28 days. At the end of incubation inorganic N was extracted from the samples with 50 mL of 2M KCl and NH,’, NO, , and NO, were deter- mined by the steam distillation procedure of Bremner and Keeney (1965). RESULTS AND DISCUSSION The effectiveness of C,H, and the commer- cial nitrification inhibitor, nitrapyrin, on inhi- bition of nitrification in Timpanogos clay loam was tested. Nitrapyrin, and C,H, at partial pressures ranging from 0.1 to 10.0 kPa, effec- tively inhibited nitrification in this soil throughout the 28-day incubation (Table 2). The soil samples were preincubated for 1 day to allow the (NH,),SO, or (NH4), SO, + ni- trapyrin solutions to equilibrate with the soil prior to C,H, addition and incubation. The difference in the NH,*—N levels between the nitrapyrin and C,H, treatments at day 0 indi- cated that approximately 20% of the added 318 GREAT BASIN NATURALIST Vol. 46, No. 2 TABLE 2. The effects of various acetylene partial pressures and nitrapyrin on nitrification in Timpanogos clay loam. C3H. Form Partial pressure # of N 0 kPa 0.1 NH," 79 NO; +NO,° 60 1.0 NH,” 80 NO, +NO, 60 10.0 NH, 80 NO, +NO,_ 59 NITRAPYRIN NH,” 98 NO, +NO,_ 5D *n.d. = not determined #Except Nitrapyrin TABLE 3. The effects of a 24-hour exposure to C,H, partial pressures ranging from 0 to 10 kPa on nitrification in Timpanogos cl. C,H, Form Partial pressure of N 0 kPa 0 NH,* 52 NO ie NOs se 0.1 NH,* 78 NO w-tNOse Gl 1.0 NH,* 70 NOs NOK = (8 G8 10.0 NH,* 68 NON ENO shines NH,’ —N was nitrified during preincubation. The finding that C,H, partial pressures rang- ing from 0.1 to 10.0 kPa inhibited nitrification concurred with earlier work of Walter et al. (1979) and Berg et al. (1982), who showed that C,H, partial pressures of 1.0 and 0.1 kPa effec- tively inhibited nitrification in soils. The effects of 24 hours of exposure to C,H, partial pressures ranging from 0 to 10.0 kPa on subsequent changes in NH,’ —Nand(NO, + NO, )—N concentrations in the Timpanogos soil are shown in Table 3. The zero time of incubation followed the 24-hour preincuba- tion and the 24 hours of exposure of C,Hg. Nitrification continued in the control samples (0 kPa) during the 24-hour period that the remaining treatments were exposed to C,H. For this reason more (NO, + NO, )—N had accumulated in the control samples during preincubation. During the first week if incu- bation following removal of C,H,, nitrification of added NH,’ —N was complete in all sam- ples except those treated with 10.0 kPa. Nitri- fication in the samples exposed to 10.0 kPa C,H, for 24 hours was slowed for approxi- Days of incubation 0 14 21 28 Inorganic N mg kg ' 79 82 83 81 62 60 61 60 81 83 84 84 61 62 61 62 80 81 84 87 61 61 59 61 95 97 97 n.d.* ol 56 53 n.d.* Days of incubation i 14 21 28 Inorganic N mg kg ' 0 0 0 0 140 147 148 150 0 0 0 0 146 146 es 5 148 0 0 0 0 144 144 152 156 45 2 0 0 93 140 150 155 mately two weeks due to the lingering effects of acetylene exposure. There were no differences in the NH, —N and (NO, +NO, )—N concen- trations during incubation of the 0.1 and 1.0 kPa treatments. During the first week following re- moval of C,H, from the samples, the nitrification rate in samples exposed to 0.1 and 1.0 kPa C,H, was 12 mg N kg ' day | compared to 4.3 mg N kg ‘day ' in those treated with 10 kPa. Samples of the Woodrow clay loam were incubated aerobically following a 24-hour ex- posure to, and subsequent removal of, C,H, partial pressures ranging from 0 to 10.0 kPa. The effects of this brief exposure on subse- quent changes in NH,’—N and (NO, + NO, )—N concentrations are shown in Table 4. During the first week following C,H, re- moval, the nitrification rates for 0, 0.1, 1.0, and 10.0 kPa C,H, treatments were 5.2, 5.7, 2.1, and 3.6 mg N kg ' day ‘, respectively. Accumulation of (NO, +NO, )—N in the samples exposed to 1.0 and 10.0 kPa was slower during the first week of incubation. Nitrification proceeded in the Woodrow cl, an unfertilized rangeland soil, at a slower pace April 1986 BURNS, TERRY: NITRIFICATION IN SOILS 319 TABLE 4. The effects of a 24-hour exposure of C,H, partial pressures ranging from 0 to 10 kPa on nitrification in Woodrow cl. C,H, Form Days of incubation Partial pressure of N 0 7 14 ik 28 kPa Inorganic N mg kg ! 0 NH," 79 31 0 0 0 NO, +NO,_ 19 56 105 106 114 0.1 INJaL” 87 54 5 0 0 NO, +NO,~ 12 52 98 104 lll 1.0 NH,” 83 70 19 0 0 NO, +NO, 12 27 82 101 lll 10.0 NH," 83 62 18 D 0 NO, +NO,— 13 38 83 93 110 TABLE 5. The effects of C,H, partial pressure and length of exposure time on nitrification in Timpanogos cl. C,H, Exposure Form Days of incubation Partial pressure time of N 0 7 14 21 28 kPa Hours Inorganic) Nima} pay aaaeennennnnn 10 24 NH,* 68 45 2 0 0 NO, +NO, 63 93 140 151 155 216 NH,” 8] 48 1 0 0 NO, +NO,~ 59 89 149 149 148 1.0 24 NH,” 70 0 0 0 0 NO, +NO, 63 144 145 153 156 216 NH," 81 47 1 0 0 NO, +NO,- 60 95 139 148 158 0.1 24 NH,” 78 0 0 0 0 NO, +NO,- 61 146 146 153 148 216 NH,* 80 47 I 0 0 NO, +NO, 61 94 148 152 158 than in the Timpanogos cl, a fertilized cropped soil. The results reported above indi- cate that brief exposure (24 hours) to C,H, partial pressures ranging from 0.1 to 1.0 kPa have little effect on nitrification in these soils. Brief exposure of soil to 10.0 kPa C,H, slowed subsequent nitrification for as long as two weeks, however. The effects of length of C,H, exposure time on recovery of nitrification in Timpanogos cl are shown in Table 5. Nitrification rates in soil sam- ples treated with 10.0 kPa acetylene were equiv- alent during the first week of incubation follow- ing C,H, removal whether exposed to the gas for 24 or 216 hours. Nitrification was nearly com- plete within one week in samples exposed to 0.1 or 1.0 kPa for only 24 hours. Nitrification rates in samples exposed to 0.1 and 1.0 kPa C,H, for 216 hours were much slower, however. The effects of lengthy exposure (216 hours) to C,H, partial pressures of 0.1 and 1.0 kPa on subsequent nitri- fication were similar to the effects of brief expo- sure (24 hours) to 10.0 kPa. Walter et al. (1979) reported that nitrifica- tion rates in soil samples exposed to C,H, partial pressures ranging from 0.1 to 1.0 kPa for 24 hours returned to those of control sam- ples after an 8 to 10 day lag period. Similar results were reported by Berg et al. (1982), who exposed soil samples to 0.01 kPa C,H, for seven days. They reported that rates of nitrate production were similar to the rates in control samples seven days after C,H, removal. The findings of this experiment indicate that both the partial pressure of C,H, and the length of exposure time affect the recovery time of nitrification in soil samples. Exposure of Woodrow cl to 0.1 and 1.0 kPa C,H, for 24 hours slowed nitrification for approximately seven days. Exposure of Timpanogos cl to 0.1 and 1.0 kPa C,H, for 24 hours had little effect on nitrification once the inhibitor was re- moved. Exposure of this soil to 0.1 and 1.0 kPa acetylene for 216 hours slowed nitrification for approximately seven days, however. These levels of C,H, exposure are commonly used in 320 laboratory and in situ denitrification studies (Yoshinari et al. 1977, Ryden, II., Develop- ment and application, 1979; Rolson et al. 1982, Ryden and Dawson 1982). The recovery of nitrification in Timpanogos soil exposed to 10.0 kPa C,H, was delayed for at least seven days whether the samples were exposed to the gas for 24 or 216 hours. Acetylene partial pres- sures of 10.0 kPa have been used in denitrifi- cation studies and in studies of concurrent denitrification and nitrogen fixation (Terry and Tate 1980a, 1980b, Yoshinari et al. 1977). Problems with the use of C,H, in denitrifi- cation studies would likely be encountered in soils where the nitrate supply is limited by nitrification. Ryden (1982) reported that deni- trification was underestimated in soil samples incubated in the laboratory in the presence of C,H, for 168 hours. Nitrate became exhausted in the samples during incubation. Problems with nitrification inhibition in denitrification studies may be avoided in laboratory studies by adding supplemental nitrate and/or adopt- ing short incubation periods (<24 hours) (Terry and Tate, 1980b). In the design of field studies on denitrification, it would be wise to rotate study sites every 7 to 14 days to allow nitrification to proceed in the soil. The use of sites previously exposed to C,H, should be avoided for 14 to 21 days because of the slow recovery of nitrification following prolonged exposure to C,H),. LITERATURE CITED ALLISON, F. E. 1965. Organic carbon. In C. A. Black, ed., Methods soil analysis. Part 2—Chemical and mi- crobiological methods. Amer. Soc. of Agron., Madison, Wisconsin. Agronomy 9: 1367-1378. BALDERSON, W. L., B. SHERR, AND W. J. PAYNE. 1976. Blockage by acetylene of nitrous oxide reduction Pseudomonas Perfectomarinus. Appl. Environ. Microbiol. 31: 504—508. BERG, P., L. KLEMEDTSSON, AND T. ROssWALL. 1982. In- hibitory effect of low partial pressure of acetylene on nitrification. Soil Biol. Biochem. 14: 301-303. GREAT BASIN NATURALIST Vol. 46, No. 2 BERGERSON, F. J. 1980. Methods of evaluating biological nitrogen fixation. John Wiley and Sons Inc., New York. BREMNER, J. M. 1965. Total nitrogen. In C. A. Black, ed., Methods of soil analysis. Part 2—Chemical and microbiological methods. Amer. Soc. of Agron.., Madison, Wisconsin. Agronomy 9: 1149-1178. BREMNER, J. M., AND D. R. KEENEY. 1965. Steam distilla- tion methods for determination of ammonium, nitrate, and nitrite. Anal. Chim. Acta. 32: 485-495. HALLMARK, S. L., AND R. E. TERRY. 1985. Field measure- ment of denitrification in irrigated soils. Soil Sci. 140: 35-44. Harpy, R. W. F., R. D. HOLSTEN, E. K. JACKSON, AND R. C. Burns. 1968. The acetylene-ethylene assay for Np fixation: Laboratory and field evaluation. Plant Physiol. 42: 1185-1207. ROLSTON, D. E., A. N. SHARPLEY, D. W. Toy, AND F. E. BROADBENT. 1982. Field measurement of denitri- fication: III. Rates during irrigation cycles. Soil Sci. Soc. of Amer. J. 46: 289-296. RYDEN, J.C. 1982. Effects of acetylene on nitrification and denitrification in two soils during incubation with ammonium nitrate. J. Soil Sci. 33: 263-270. RYDEN, J. C., AND K. P. Dawson. 1982. Evaluation of the acetylene-inhibition technique for the measure- ment of denitrification in grassland soils. J. Sci. Food Agric. 33: 1197-1206. RYDEN, J. C., L. J. LUND, AND D. D. Focut. 1979. Direct measurement of denitrification loss from soils: I. Laboratory evaluation of acetylene inhibition of nitrous oxide reduction. Soil Sci. Soc. Amer. J. 43: 110-140. RYDEN, J. C., L. J. LUND, J. LETEy, AND D. D. Focur. 1979. Direct measurement of denitrification loss from soils: II. Development and application of field methods. Soil Sci. Soc. of Amer. J. 43: 110-118. TERRY, R. E., AND R. L. TaTE. 1980a. Denitrification as a pathway for nitrate removal from organic soils. Soil Sci. 129: 162-166. . 1980b. The effect of nitrate on nitrous oxide reduc- tion in organic soils and sediments. Soil Sci. Soc. Amer. J. 44: 744-746. Terry, R. E., R. L. TATE, AND J. M. Duxsury. 1981. The effect of flooding on nitrous oxide emissions from an organic soil. Soil Sci. 132: 228-232. WatteR, H. M., D. R. KEENEY, AND I. R. FILLERY. 1979. Inhibition of nitrification by acetylene. Soil Sci. Soc. Amer. J. 43: 195-196. YOSHINARI, T., R. HyNES, AND R. KNOWLES. 1977. Acetylene inhibition of nitrous oxide reduction and measurement of denitrification and nitrogen fixation in soil. Soil Biol. Biochem. 9: 177-183. NEW THAGRIINE LEAFHOPPERS FROM THE ORIENTAL REGION, WITH A KEY TO 30 SPECIES (HOMOPTERA: CICADELLIDAE: COELIDIINAE) M. W. Nielson ABSTRACT.—Five new species of Thagria from the Oriental region are described and illustrated. These include melichari from Thailand, unidentata from Indonesia, marissae from southern China, bifida from Nepal, and insolentis from an undetermined locality in the Oriental region. There are presently 166 species in this large and unique genus. A key to males of 30 species is included. The genus Thagria Melichar is the largest group of coelidiine leafhoppers. Although they occur primarily in the Oriental region, many species are found in the Australian re- gion (not known in Australia proper) and sev- eral are in the southern Palearctic region (southern China, southern Korea, and south- ern Japan). Prior to 1977 only 36 species were known. Since then 125 species have been de- scribed (Kwon and Lee 1979, Nielson 1977, 1980a, 1980b, 1980c, 1980d, 1982). The five new species described herein bring the present total to 166 species. The genus is uniquely characterized by the males possessing a distinctive and highly di- verse ventral paraphysis on which a tubular aeadeagal shaft is attached basally to and freely articulates dorsally with the paraphysis. The many configurations of the ventral para- physis in combination with highly modified structures of the 10th segment and caudodor- sal processes of the pygofer differentiate the numerous species. A key to males of 30 species including those described in previous papers (except Kwon and Lee 1979) after my 1977 revision and those treated herein is presented. A regional key for all known species will be presented later. Host plants and biology of species in the group are very poorly known. Key to Males of Thagria 1. Clypellus broad, swollen basally or nearly so, basal width equal to or greater than basal width of clypeus, lateral margins usually nar- rowedimnedial yaar ne em hee eles See 2 Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah 84602. 9(8). 321 Clypellus narrow, never swollen basally, basal width narrower than basal width of clypeus, lateral margins usually parallel, sometimes expanded distally ............. 16 Ventral paraphysis curved ventrally at distal 1/2 to 1/3 in lateral view, apex decurved ..... 3 Ventral paraphysis not as above, in lateral view straight or recurved ................. 5 Style with apex bifurcate or divided into 2 slenderramilprene eats ener 4 Style not as above (Fig. 16, Nielson 1980a). . siaGegl Snes Mace tee rae erceceng eA blockeri Nielson Aedeagus long, extending beyond midlength of ventral paraphysis; 10th segment process with dentate process on middle of dorsal mar- ain (Ie, BF) seccccsoccccecueons bifida, n. sp. Aedeagus shorter, reaching to about mid- length of ventral paraphysis; 10th segment process with longer process on ventral margin (Fig. 25, Nielson 1980b) . thailandensis Nielson Ventral paraphysis symmetrical............ 6 Ventral paraphysis asymmetrical.......... 10 Tenth segment with paired processes ....... 7 Tenth segment without paired processes (Fig. 13, Nielson 1980b) ....... ampla Nielson Ventral paraphysis without basal paired pro- cesses on dorsal margin................+-. 8 Ventral paraphysis with basal paired pro- cesses on dorsal margin (Fig, 22, Nielson LOSOb) eres ence serrastyla Nielson Style very long, exceeding midlength of ven- tral paraphysis; ventral paraphysis without Spines|istal lyanmerck essen arc cena 9 Style very short, not reaching midlength of ventral paraphysis; ventral paraphysis with lateral spines distally (Fig. 28, Nielson 1982) barbata Nielson Style attenuated distally (Fig. 2, Nielson OSD eee ees, Poses ea Someenaets fossiata Nielson 14(10). 15(14). GREAT BASIN NATURALIST Style forked distally (Fig. 11, Nielson 1980c) RRO ies sees Heroes RCN furculata Nielson Ventral paraphysis with basal processes on Glorasall WHA ooo donc asccgo0000000000006 11 Ventral paraphysis without basal processes but with medial or subapical processes on dorsalimangin'=seaen ae reese tere 14 . Ventral paraphysis with paired basal pro- CESSESPr tes ao eters Rute oath ihn ate etn SEER 12 Ventral paraphysis with single basal process. 13 . Basal processes on paraphysis symmetrical (atie43,. UL) 6 coccococe sod a0h melichari, n. sp. Basal processes on paraphysis asymmetrical (Fig. 40, Nielson 1982) ..... hollowayi Nielson . Style broad throughout in dorsal view, with- out dentate subapical processes (Fig. -39, Nielson 1980b)............. boulardi Nielson Style narrowed at distal 1/4 in lateral view, with dentate subapical process (Fig. 35, Niel- Song! 98 0b) sameeren paraornata Nielson Tenth segment and caudodorsal margin of pygofer with processes of equal length in lat- eral view; ventral paraphysis with short lat- eral process distad of middle.............. 15 Tenth segment and caudodorsal margin of pygofer with processes of unequal length in lateral view; ventral paraphysis with short lateral process on middle (Fig. 4, Nielson NOSOD Saye etary undulata Nielson Tenth segment processes very narrow and sinuate, nearly needlelike at distal 2/3 in dor- sal view (Fig. 8, Nielson 1980b) . capilla Nielson Tenth segment processes broader and nearly straight, not needlelike in dorsal view (Fig. 20, Nielson 1980a)....... paradigitata Nielson Ventral paraphysis symmetrical........... 17 Ventral paraphysis asymmetrical.......... 25 . Ventral paraphysis keeled ventrally........ 18 Ventral paraphysis not as above........... 20 . Ventral paraphysis with subbasal ventral keel Ventral paraphysis with subapical ventral keel (Fig. 10, Nielson 1980d) . paraloae Nielson . Style very long, extending beyond apex of ventral paraphysis (Fig. 22, Nielson 1980d) oc ee MOCO eae trim aa eee samuelsoni Nielson Style very short, extending only to base of ventral paraphysis (Fig. 3, Nielson 1980c) ventrocarina Nielson . Ventral paraphysis with paired basal process onidorsalimanc nee eee rece 21 Ventral paraphysis not as above........... 22 . Paired basal processes of paraphysis very long, nearly reaching to apex of paraphysis (Fig. 3, Nielson 1980d) ..... bilateralis Nielson Paired basal processes of paraphysis shorter, not reaching midlength of paraphysis (Figs. (Geel LY AR Be nematic nite: eta insolentis, n. sp. Vol. 46, No. 2 22(20). Caudoventral lobe of pygofer without spines = Caudoventral lobe of pygofer with 2 short spines apically (Fig. 7, Nielson 1982)....... bidentata Nielson . Ventral paraphysis without lateral processe — Ventral paraphysis with lateral processes sub- apically (Fig. 3, Nielson 1980a)............ srilankensis Nielson 3). Style with subapical bifurcation (Fig. 21, Nielson1982) anne eee bifurcata Nielson _- Style without subapical bifurcation (Fig. 11, Nielson 1980a).............. brincki Nielson . Style with distal half straight or nearly so... 26 — Style with distal half hooked (Fig. 17, Nielson 19800) se ecinn ookt Oe eee paraexilis Nielson . Ventral paraphysis without ventral keel .... 27 — Ventral paraphysis with ventral keel sub- basally (Fig. 17, Nielson 1982)............ re een ME RRA A 5 6b c mutabilis Nielson . Ventral paraphysis with 1—2 lateral processes On ORNear apex... 0 eee eee 28 — Ventral paraphysis without such processes.. 29 . Ventral paraphysis with a pair of unequal dis- tal processes (Fig. 23) ........ marrisae, n. sp. — Ventral paraphysis with a single, large, retrorse lateral process subapically (Fig. 35, Nielson 1982).............. retrorsa Nielson . Caudoventral lobe of pygofer with a single long spine (Fig. 26) ........ unidentata, n. sp. Caudoventral lobe of pygofer without such spine (Fig. 13, Nielson 1980c) kaloostiani Nielson Thagria bifida, n. sp. Figs. 1-6 LENGTH: Male 6.90 mm. Moderate-sized, slender species. General color black with tannish translucent costa, face black. Head small, subconical, much narrower than pronotum; crown broad, width about equal to width of eyes, produced beyond ante- rior margin of eyes, elevated above level of eyes, lateral margins convergent basally; eyes moderately large, semiglobular; pronotum with length about equal to length of crown; scutellum large; forewings long and narrow, venation typical of genus; clypeus long and broad, lateral margins excised near middle; clypellus short and broad, base broad and swollen, lateral margins below converging to truncate apex. April 1986 NIELSON: NEW ORIENTAL LEAFHOPPERS 323 Figs. 1-6. Thagria bifida: 1, Male pygofer and 10th segment, lateral view. 2, Tenth segment, and pygofer processes, dorsal view. 3, Connective, aedeagus, ventral paraphysis and right style, dorsal view. 4, Aedeagus and ventral paraphysis, lateral view. 5, Right style, lateral view. 6, Plate, ventral view. MALE: Pygofer in lateral view with rather long, broad, caudoventral lobe, apex narrowly rounded; caudodorsal margin with long, nar- row, slightly sinuate process, nearly reaching apex of caudoventral lobe (Fig. 1); 10th seg- ment with pair of long slender acuminate pro- cesses nearly reaching to apex of anal tube, processes with 2 small dentate projections, one subapical and one near middle on dorsal margin (Figs. 1, 2); aedeagus symmetrical, long, extending beyond midlength of ventral paraphysis (Fig. 3); ventral paraphysis short, very broad at basal half in dorsal and lateral views, narrowed at distal halfand decurved in lateral view (Figs. 3, 4); style very long, ex- tending beyond apex of ventral paraphysis, bifurcate subapically, inner bifurcation shorter than outer one (Fig. 3); plate long and narrow with few lateral macrosetae and few short microsetae apically (Fig. 6). HOLOTYPE (male): NEPAL: Ktmd. [Kat- mandu], Pulchauki, 8000’, 27. VII. 1967. Can. Nepal Exp. (CNC) REMARKS: This species is similar to obrienae Nielson but can be distinguished by the diag- nostic bifurcate style. Thagria melichari, n. sp. Figs. 7-13 LENGTH: Male 6.60 mm. Moderately robust species. General color ochraceous with narrow transverse ivory markings on forewings, veins embrowned with small irregular ochraceous spots, on veins. Head much narrower than pronotum (Fig. 7); crown narrow, produced distally beyond anterior margin of eyes, length twice basal width, anterior margin angulate, lateral mar- GREAT BASIN NATURALIST Vol. 46, No. 2 Figs. 7-13. Thagria melichari: 7, Head and pronotum, dorsal view. 8, Face, ventral view. 9, Male pygofer and 10th segment, lateral view. 10, Tenth segment and pygofer processes, dorsal view. 11, Aedeagus and ventral paraphysis, dorsal view. 12, Aedeagus and ventral paraphysis, lateral view. 13, Right style, lateral view. gins convergent basally; eyes large, elongate- ovoid; pronotum large with median longitudi- nal carina; forewing with venation typical of genus; clypeus long and rather broad; clypel- lus slightly swollen basally, basal width nearly equal to basal width of clypeus (Fig. 8). MALE: Pygofer with long, narrow, cau- doventral lobe, caudodorsal margin with pair of broad processes (Fig. 9); 10th segment with pair of long ventral processes, processes broad basally, abruptly tapered distally with small projection laterally near middle of process (Figs. 9, 10); aedeagus symmetrical, moder- ately long, about half as long as ventral para- physis (Fig. 11); ventral paraphysis slightly asymmetrical, very broad basally in dorsal view, asymmetrically clefted distally, with pair of long basal processes (Figs. 11, 12); style very long, slender, pointed distally and curved laterally in lateral view (Fig. 13); plate long and narrow, typical of genus. HOoLotyPE (male), THAILAND: Muok- Lek, 1,000 ft, _._—=«. «XI. ___., H.. Fruhstorfer. Additional labels with following information: “H. Fruhstorfer, vend. 25. V. 1924,” “Arya hyalinopunctata n. sp., manuscript name, L. Melichar det.” (MM). Allotype (female), THAILAND: Pakchong, 100 m N of Bangkok, Dec. 2, 1957, J. L. Gressitt (BPBM). Paratypes: VIET NAM: 33 km NE Ban Me Thuot, 500 m, 1 female, 16-18. V. 1960, L. W. Quate (author's collection). REMARKS: This species is similar in male genitalia characters to sarawakensis Nielson but can be separated by the configuration of the 10th segment processes and caudodorsal processes of the pygofer, by the asymmetri- cally clefted apex of the ventral paraphysis, and by the current geographical range. This species is named for Dr. Leopold Melichar in recognition of his outstanding contributions to leafhopper systematics. April 1986 NIELSON: NEW ORIENTAL LEAFHOPPERS 325 19 16 Figs. 14-19. Thagria insolentis: 14, Male pygofer and 10th segment, lateral view. 15, Tenth segment and pygofer processes, dorsal view. 16, Connective, aedeagus, ventral paraphysis, and right style, dorsal view. 17, Aedeagus and ventral paraphysis, lateral view. 18, Right style, lateral view. 19, Plate, ventral view. Thagria insolentis, n. sp. Figs. 14-19 LENGTH: Male 5.90 mm. Small, slightly robust species. General color light brown with numerous irregular tannish markings on forewings, bullae on dark pronotum ochraceus, crown light tan basally with blackish markings anteriorly, face red- dish brown. Head large, subconical, narrower than pronotum; crown somewhat narrow, width less than transocular width, elevated above level of eyes, produced beyond anterior mar- gin of eyes; eyes large, semiglobular; prono- tum short, median length about equal to me- dian length of crown, with short median longitudinal carina originating on anterior margin; scutellum large; forewing moderately long, venation as in description of genus; clypeus long, narrow, excised near antennal sockets; clypellus long and narrow, lateral margins nearly parallel. MALE: Pygofer in lateral view with elongate triangular caudoventral lobe (Fig. 14); caudodor- sal margin with ornate process, process broad basally, abruptly decurved medially with nar- row, asymmetrical bifid apex, ventral margin with narrow, hooked secondary process on mid- dle, dorsal margin with short secondary process (Figs. 14, 15); 10th segment with pair of narrow long processes nearly reaching to apex of cau- doventral lobe (Fig. 14); aedeagus symmetrical, very long and tubular, curved dorsally at distal half and extending to about apex of ventral para- physis in lateral view (Figs. 16, 17); ventral para- physis symmetrical, broad basally with pair of long narrow processes basally on dorsal margin, lateral margins of paraphysis convergent distally to narrow convex apex with short dentate sub- apical projections laterally (Figs. 16, 17); style very long, attennuated, and sharply pointed api- GREAT BASIN NATURALIST Vol. 46, No. 2 Figs. 20-25. Thagria marissae: 20, Male pygofer, lateral view. 21, Pygofer processes, dorsal view. 22, Connective, aedeagus, ventral paraphysis, and right style, dorsal view. 23, Aedeagus and ventral paraphysis, lateral view. 24, Right style, lateral view. 25, Plate, ventral view. cally, exceeding apex of paraphysis (Fig. 16); plate long and narrow, with many long microse- tae apically (Fig. 19). HoLotyPE (male): [ORIENTAL REGION]: Friese, Teoor (or Tevor), no date, no collector (NM). REMARKS: The species is near luteifascia (Walker). It can be easily distinguished from that species by the ornate caudodorsal processes of the pygofer. The locality of this species is not known but is presumed to be in the Oriental region. In a recent communication from Dr. A. Kaltenbach, Naturhistorishces Museum, Vi- enna, he stated that the specimen may have come from the Friese collection (H. Freise, 1860-1948) but did not know if Friese collected in the Oriental region. Thagria marissae, 0. sp. Figs. 20-25 LENGTH: Male 5.75 mm. Small, slender species. General color light golden brown, suffused with brown markings near apex of forewings and near middle of costa. Head large, narrower than pronotum, sub- conical; crown broad, about as wide as eyes, produced distally beyond anterior margin of eyes, lateral margins convergent basally, ele- vated above level of eyes; eyes large, semi- globular; pronotum and scutellum short, me- dian length of each nearly equal; forewing long and narrow, venation typical of genus; clypeus broad anteriorly, clypellus short, lat- eral margins nearly parallel. MALE: Pygofer in lateral. view with short broad caudoventral lobe, tapered toward apex, apex rounded, caudodorsal margin with short narrow lobelike process extending dis- tally and not reaching apex of caudoventral lobe (Figs. 20, 21); 10th segment short, sim- ple, without ventral processes (Fig. 20); aedeagus symmetrical, very long and narrow, ae ———— April 1986 NIELSON: NEW ORIENTAL LEAFHOPPERS 327 31 Ag | El 28 Figs. 26-31. Thagria unidentata: 26, Male pygofer and 10th segment, lateral view. 27, Tenth segment and pygofer processes, dorsal view. 28, Connective, aedeagus, ventral paraphysis, and right style, dorsal view. 29, Aedeagus and ventral paraphysis, lateral view. 30, Right style, lateral view. 31, Plate, ventral view. nearly reaching to apex of ventral paraphysis (Figs. 22, 23); ventral paraphysis asymmetri- cal, broad basally with gradual constriction along middle and slightly expanded distally in dorsal view with pair of short unequal, sharply pointed, lateral processes apically (Figs. 22, 23); style very short, extending just beyond base of aedeagus in dorsal view, narrowly tri- angular in dorsal view (Fig. 22); plate long and very narrow throughout with tuft of long mi- crosetae apically (Fig. 25). HOLOTYPE (male): CHINA: Iwa Bi, Hainan Isl., 25. VII. 1935, L. Gressitt (NCSU). REMARKS: Thagria marissae is similar to T. lurida (Melichar). It can be separated from lurida by the narrower caudoventral lobe of the pygofer, by the longer aedeagus that reaches to the apex of the ventral paraphysis, by the asymmetrical ventral paraphysis, and by its known geographical range. I name this species for my granddaughter, Marissa Jean Hammer. Thagria unidentata, n. sp. Figs. 26-31 LENGTH: Male, 7.25—7.75 mm. Moderately long, slender species. General color tannish brown; eyes tan to brown; crown and pronotum tan, posterior margin of prono- tum blackish; scutellum tan to brown; forewing translucent, veins blackish; face tan. Head much narrower than pronotum, sub- conical; crown narrower than width of eyes, produced beyond anterior margin of eyes, narrowly rounded distally, lateral margins convergent basally, slightly carinate laterally; pronotum and scutellum equal in length, each 328 equal in length of crown; forewing long and narrow, venation typical of genus; clypeus long and narrow, lateral margins constricted near antennal sockets; clypellus short, lateral margins wider distally than proximally. MALE: Pygofer in lateral view with short caudoventral lobe, lobe with long spine on caudoventral margin, spine as long as lobe, caudodorsal margin of pygofer with single long process, process sharply pointed api- cally, curved posterioventrally, and reaching to apex of caudoventral lobe of pygofer (Fig. 26); 10th segment with pair of long processes, processes decurved ventrally at distal 1/3 (Fig. 27); aedeagus short, tubular, reaching to about midlength of ventral paraphysis (Fig. 28); ventral paraphysis slightly asymmetrical in dorsal view, broad basally with lateral mar- gins gradually convergent distally, distal 1/3 slightly undulated with apex slightly curved laterally (Figs. 28, 29); style short, not reach- ing midlength of ventral paraphysis, distal half narrowly attenuated (Figs. 28, 30); plate long and narrow throughout, with long microsetae on lateral margins and at apex (Fig. 31). HOLOTYPE (male): INDONESIA: Siberat Isl., West Sumatra, .LX. 1924, B. K. and N. Raffles, Singapore Museum (BMNH). Paratypes. 1 male, same data as holotype (au- thor’s collection). REMARKS: Thagria unidentata is similar to T. fryeri (Distant) but lacks the distinctive lateral processes on the dorsal margin of the ventral paraphysis and has a much longer spine on the caudoventral margin of the cau- doventral lobe of the pygofer. GREAT BASIN NATURALIST Vol. 46, No. 2 ACKNOWLEDGMENTS The loan of specimens from the following indi- viduals and their institutions is much appreci- ated: the late Dr. J. Linsley Gressitt, Bernice P. Bishop Museum, Honolulu (BPBM), Dr. K. G. A. Hamilton, Canada National Collection, Ott- awa (CNC), Dr. A. Kaltenbach, Naturhis- torische Museum, Vienna (NM), Dr. W. J. Knight, British Museum (Natural History), Lon- don (BMNH), Dr. Pavel Lauterer, Moravian Museum, Brno (MM), and Dr. David A. Young (retired), North Carolina State University, Raleigh (NCSU). I thank Jean Stanger for the fine illustrative work and Dr. James P. Kramer, U.S. National Museum, Washington, D.C., and Dr. Paul W. Oman, Oregon State University, Corvallis, for reviewing the paper. LITERATURE CITED Kwon, Y. J., AND C. E. LEE. 1979. On some new and little known Palearctic species of leafhoppers. Nature and Life (Kyungpook J. Biol. Sci.) 9(2): 69-97. NIELSON, M. W. 1977. A revision of the subfamily Coelidiinae (Homoptera: Cicadellidae) I. Tribe Thagriini. Pacific Insects Monog. 34. 218 pp., 808 figs. . 1980a. New Oriental species of leafhoppers in the Genus Thagria (Homoptera: Cicadellidae: Thagriini). J. Kansas Entomol. Soc. 53: 123-131. . 1980b. Seven new species of Thagriine leafhoppers from Southeast Asia (Homoptera: Cicadellidae: Thagriini). J. Kansas Entomol. Soc. 53: 305-319. . 1980c. New leafhopper species of Thagria from Malaysia (Homoptera: Cicadellidae: Thagriini). J. Kansas Entomol. Soc. 53: 343-349. . 1980d. Four new leafhoppers species of Thagria from the Australian region with notes on Thagria sum- bawensis (Jacobi) (Homoptera: Cicadellidae: Thagri- ini). J. Kansas Entomol. Soc. 53: 607-616. . 1982. Some additional new species of Thagriine leafhoppers from Malaysia and Indonesia (Cicadelli- dae: Coelidiinae: Thagriini). J. Kansas Entomol. Soc. 55: 461-473. =a ea GENUS PARALIDIA WITH DESCRIPTIONS OF NEW SPECIES (HOMOPTERA: CICADELLIDAE: COELIDIINAE) M. W. Nielson! ABSTRACT.—The generic concept of Paralidia Nielson, type-species, Coelidia plaumanni Linnavuori, is reeluci- dated after five additional new species were found in South America. These are described and illustrated. Four species, all from Brazil, include spinata, retrorsa, denticulata, and bispinosa. One species, singularis, is from Ecuador. A key to males of the six known species is also presented. The monobasic genus Paralidia was estab- lished for Coelidia plaumanni Linnavuori in the tribe Coelidiini by Nielson (1982). Since its erection five additional species have been found. These not only broaden the concept of the genus but also solidify its taxonomic and zoological base. Generic characterization, in addition to that previously described, includes the fol- lowing features: Small to moderately large, slender species; head slightly to distinctly nar- rower than pronotum; crown always narrow and produced slightly to nearly 1/3 of its me- dian length beyond anterior margin of crown, lateral margins parallel to convergent basally and varying from slightly to prominently cari- nate; pronotum and scutellum usually short and nearly always equal to or shorter than crown; clypeus nearly always narrower anteri- orly than basally; male pygofer with cau- doventral process always present, slender and long, but varying in length, degree of fusion basally with caudal margin of pygofer and con- figuration; aedeagus nearly always asymmet- rical, usually narrow and somewhat tubular, always with dorsally curved apex that usually has 1-3 spines, sometimes with serrations or spines on middle of shaft; style varies from moderately long to very long (reaching apex of aedeagus) and very slender at distal 2/3, some- times enlarged preapically; plate long with middle third of outer margin enlarged and convex, always profusely setose with long mi- crosetae. The known species occur in the eastern states of Brazil, with one species from Ecuador. The host plants, biology, and eco- nomic importance of the group are not known. Females are rarely collected. Key to Males of Paralidia 1. Aedeagus with 1-3 spines on or near apex, spines along middle of shaft absent .......... 2 _ Aedeagus without apical spines; spines along middle of shaft present (Fig. 4). . . .spinata, n. sp. Aedeagus with 2—3 spines on or near apex .... 3 — Aedeagus with a single terminal spine (Fig. 13) SEI Ra tee hart tan ne singularis n. sp. Aedeagus with 2 apical or subapical spines.... 4 -- Aedeagus with | apical and 2 subapical spines (EST eal) ee ooo eI raat retrorsa, N. Sp. Aedeagus with 2 subapical, needlelike spines, spines asymmetrical and not originating on the same sagittal plane..............-..--0005- 5 — Aedeagus with 2 apical, basally broad spines, spines symmetrical and originating on the same sagittal plane (Fig. 799°, Nielson 1982) Bihar Ain aio area hicks Seow plaumanni (Linnavuori) Aedeagus with dentate, lateral flange on one side of middle of shaft; pygofer with a partial- ly fused, membranous caudoventral process; style needlelike at distal 2/3 (Fig. 28) series create aia Me Sree Nes even denticulata, n. sp. — Aedeagus without such flange; pygofer with 4 well-developed caudoventral process; style with a preapical expansion (Fig. 35)......... paar ete st § 0000000000 00000 04 I DIMORE 5 > So) Paralidia spinata n. sp. Figs. 1-8 LENGTH: Male 6.15-6.80 mm, female 7.70 mm. Moderate-sized, slender species. General color pale ochre with broad (narrow in fe- Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah 84602. ?Separation of apical spines not apparent in Figure 799. 329 GREAT BASIN NATURALIST Figs. 1-8. Paralidia spinata: 1, Head, pronotum and scutellum, dorsal view; 2, Face, ventral view; 3, Male pygofer, lateral view; 4, Aedeagus, lateral view; 5, Connective, aedeagus and right style, dorsal view; 6, Style, lateral view; 7, Plate, ventral view; 8, Female abdominal venter, ventral view. male), black, medial longitudinal band from crown to near forewing apex, eyes light red- dish brown, face ochre. Head rather large, narrower than prono- tum (Fig. 1); crown very narrow, produced slightly beyond anterior margin of eyes, rounded distally, lateral margins carinate and parallel; eyes very large, about twice as wide as coronal width, semiglobular; pronotum and scutellum short, median length of each slightly less than median length of crown; forewing elongate, venation typical of genus; clypeus long and broad, lateral margins broadly convex; clypellus short, lateral mar- gins broader distally (Fig. 2). MALE: Pygofer in lateral view with a long, narrow slightly sinuate, acuminate caudoven- tral process, process broad at basal 1/3, cau- dodorsal margin with a shorter, narrow sinu- ate process, apex curved slightly downward and bluntly pointed (Fig. 3); aedeagus sym- metrical, long, nearly tubular throughout, slightly compressed laterally with numerous conspicuous dentate microspines laterally on distal half of shaft before apex, spines not as readily apparent in some specimens, apex curved dorsally in lateral view, rounded api- cally and without terminal spines (Figs. 4, 5); gonopore terminal; connective as in singu- laris ; style long, reaching to about midlength of aedeagal shaft, needlelike at distal half (Figs. 5, 6); plate long, similar in shape to singularis except apex more pointed, pro- fusely setose (microsetae) (Fig. 7). FEMALE: Seventh sternum nearly twice as long as preceding segment, caudal margin sin- uate (Fig. 8). HOLOTYPE (male): BRAZIL: Minaes Gerais, Santa Barabara, Caraca, ___.I. 1970, F. M. Oliveira, B. M. 1971-165 (BMNH). Allotype April 1986 NIELSON: NEW NEOTROPICAL LEAFHOPPERS 331 Figs. 9-15. Paralidia singularis: 9, Head, pronotum and scutellum, dorsal view; 10, Face, ventral view; 11, Male pygofer, lateral view; 12, Connective, aedeagus and right style, dorsal view; 13, Aedeagus, lateral view; 14, Style, lateral view; 15, Plate, ventral view. (female), Minaes Gerais, Delfim Moreira, 1,100 m,___. 1.1972, F. M. Oliveira, B. M. 1972-541 (BMNH). Paratypes: 4 males, same data as holotype, 4 males, same data as allo- type (BMNH), 2 males, same data as holotype (USNM, author's collection). REMARKS: Paralidia spinata is similar to denticulata but can be separated from it and all other species in the genus by the minute spines on the aedeagal shaft, which also lacks terminal spine(s). Paralidia singularis, n. sp. Figs. 9-15 LENGTH: Male 5.50 mm. Small, slender species. General color dark brown with large, triangular, pale translucent spot on middle of forewing and with numerous pale spots on veins; eyes deep reddish brown, pronotum with small pale spots; clypeus and clypellus black with small pale spots. Head large, narrower than pronotum (Fig. 9); crown very narrow, produced slightly be- yond anterior margin of eyes, rounded dis- tally, lateral margins carinate; eyes very large, more than twice as wide as basal width of crown, semiglobular; pronotum and scutel- lum short; elytra long and moderately broad; clypeus long and narrow, narrower anteriorly than basally; clypellus short, narrow, lateral margins broad distally (Fig. 10). MALE: Pygofer in lateral view with very long, slender, caudoventral process reaching nearly to apex of caudodorsal process, cau- dodorsal process moderately long, lobelike, apex curved dorsally (Fig. 11); aedeagus sym- metrical or nearly so, long, nearly tubular throughout with single prominent, sharply pointed spine apically (Fig. 12), apex of shaft curved dorsally with spine projecting cau- dodorsally (Fig. 13); gonopore apical near base of spine; connective stout, Y-shaped, 332 Figs. 16-23. Paralidia retrorsa: 16, Head, pronotum and scutellum, dorsal view; 17, Face, ventral view; 18, Male pygofer, lateral view; 19, Aedeagus, lateral view; 20, Terminus of aedeagal shaft, caudal view; 21, Connective, aedeagus and right style, dorsal view; 22, Style, lateral view; 23, Plate, ventral view. with short stem; style very long and slender, nearly reaching apex of aedeagus, distal third expanded except for abruptly pointed apex (Fig. 12, 14); plate long, narrowed subbasally, broad subapically and abruptly narrowed to a bluntly rounded apex, with numerous long microsetae (Fig. 15). HOLOTYPE (male): ECUADOR: Quito (Equateur), .X.1930, R. Benoist (MNHN). REMARKS: Paralidia singularis is nearest to retrorsa n. sp. and can be distinguished by the single long spine on the apex of the aedea- gus and by the large pale triangular spot on the middle of the forewing. Paralidia retrorsa, n. sp. Figs. 16-23 LENGTH: Male 6.90 mm, female 7.25—8.00 mm. GREAT BASIN NATURALIST Vol. 46, No.2 | color dark brown to black except for pale face, crown and costa of forewing deep tan, veins of forewing sometimes mottled. Head large, narrower than pronotum (Fig. 16); crown very narrow, about 1/2 width of eyes, much produced distally beyond anterior margin of eyes, distal length nearly 1/3 entire median length, anterior margin acutely rounded, disk | foveate medially, lateral margins nearly parallel, slightly carinate; eyes very large, elongate- _ ovoid; pronotum and crown short, each shorter than median length of crown; forewing elongate, | Moderate-sized, slender species. General venation typical of genus; clypeus long and nar- row, narrower anteriorly than posteriorly with | short broad ridge medially and originating on | anterior margin then dividing into several | rugose lines that terminate about middle; clypel- _ lus long and narrow, lateral margins expanding _ distally (Fig. 17). | } April 1986 NIELSON: NEW NEOTROPICAL LEAFHOPPERS 333 Figs. 24-31. Paralidia denticulata: 24, Head, pronotum and scutellum, dorsal view; 25, Face, ventral view; 26, Male pygofer, lateral view; 27, Connective, right style and aedeagus, dorsal view; 28, Aedeagus, lateral view; 29, Terminus of aedeagal shaft, caudal view; 30, Style, lateral view; 31, Plate, ventral view. MALE: Pygofer in lateral view with weakly sclerotized caudoventral process, process similar to denticulata but fused to pygofer at more than half its base, caudodorsal margin with short, broad lobe, lobe rounded apically with short, bluntly pointed process subapi- cally on dorsal margin, process directed mesally (Fig. 18); aedeagus asymmetrical, very long and narrow, apex abruptly curved dorsally in lateral view and terminating with single curved, sharply pointed, retrorse spine (Fig. 19), two additional membranous spines on caudal margin of curved apex, one above the other, not on same sagittal line and both pointed dorsally (Fig. 20), dorsolateral mar- gins of shaft basad of apex not parallel at distal 1/3 in lateral view (Fig. 19); gonopore apical near base of terminal spine; connective as in genus; style moderately long, distal half very narrow and membranous (Figs. 21, 22): plate long and narrow, similar in configuration to P. plaummani (Linnavuori) with numerous long microsetae (Fig. 23). FEMALE: Seventh sternum about twice as long as preceding segment, caudal margin sinuate as in spinata. HOLOTYPE (male): BRAZIL: no locality, no date, M. Alvarenga, B.M. 1971-165 (BMNH). Allotype (female), same data as holotype (BMNH). Paratypes: 3 females, same data as holotype (BMNH), 1 female, Sao Paulo, S. Bar- reiro, S. Bocaina, 1,650 m, ___. XII. 1968, M. Alvarenga, B.M. 1970-484 (author's collection). REMARKS: This species is similar in. general habitus and male genital characters to spinata n. sp. but can be distinguished by the short, ante- rior median clypeal ridge, by the prominent asymmetrical aedeagus, and by the retrorse ter- minal spine with 2 subterminal attendant spines on the aedeagus. The well-produced crown will also separate this species from all others in the genus. 334 GREAT BASIN NATURALIST Vol. 46, No. 2 SE FA SSS RK NK ne SSS 39 37 Figs. 32-39. Paralidia bispinosa: 32, Head, pronotum and scutellum, dorsal view; 33, Face, ventral view; 34, Male pygofer, lateral view; 35, Aedeagus, lateral view; 36, Terminus of aedeagal shaft, caudal view; 37, Connective, right style and aedeagus, dorsal view; 38, Style, lateral view; 39, Plate, ventral view. Paralidia denticulata, n. sp. Figs. 24-31 LENGTH: Male 6.90-7.10 mm. Medium-sized, slender species. General color light reddish brown with suffused deep brown medially on distal half of forewings, face tannish brown. Head moderately large, subconical, nar- rower than pronotum (Fig. 24); crown very narrow, produced distally beyond anterior margin of eyes, narrowly rounded distally, lateral margins broadly convex, carinate; eyes very large, elongate-ovoid; pronotum long, median length slightly greater than median length of crown; scutellum small; forewing elongate, venation typical of genus; clypeus long and narrow, narrower anteriorly than basally; clypellus long and narrow (Fig. 25). MALE: Pygofer in lateral view with long, very narrow, acuminate caudoventral pro- cess, process completely hidden by caudal margin of pygofer, caudodorsal margin with short truncate lobe (Fig. 26); aedeagus asym- metrical, long, subtubular throughout with lateral finely dentate asymmetrical flange along middle of dorsal margin in dorsal view (Fig. 27) and rows of fine teeth below on mid- dle 2/3 of shaft, apex of shaft abruptly curved dorsally in lateral view with 2 very short, sharp spines apically, one below the other but not originating on the same sagittal line (Figs. 28, 29): gonopore apical between spines; con- nective typical of genus; style long, extending beyond midlength of aedeagal shaft, needle- like at apical 2/3 (Fig. 30); plate typical of | genus, profusely setose (Fig. 31). HOLOTYPE (male): BRAZIL: Rio de Janeiro, Guanabara, .X.1970, M. Alvarenga, B.M. 1971-165 (BMNH). Paratypes: BRAZIL: Bahia, Itapetinga, ID IRIE). = —= — April 1986 F. M. Oliveira, B. M. 1971—165 (author's col- lection). REMARKS: Paralidia denticulata is similar to spinata n. sp. but can be separated from it by the lateral tooth flange that occupies the greater middle of the aedeagal shaft and by the presence of apical spines. Paralidia bispinosa, n. sp. Figs. 32-39 LENGTH: Male 8.10 mm. Large, slender species. General color tan- nish throughout, pale spots on dark veins of forewings. Head large, distinctly narrower than prono- tum (Fig. 32); crown very narrow, about 1/2 as wide as transocular width, produced beyond anterior margin of eyes, anterior margin an- gled, lateral margins slightly convergent basally, carinate; eyes very large, semiglobu- lar; pronotum and scutellum short, each with median length about equal to length of crown; forewings elongate, venation typical of genus; clypeus long and narrow, narrower anteriorly than posteriorly; clypellus long and narrow, lateral margins broad distally (Fig. 33). MALE: Pygofer in lateral view with long, narrow, heavily sclerotized, broadly curved caudoventral process, process extending be- yond caudodorsal lobe and subtriangulate dis- tally, caudodorsal margin with broad, moder- ately long lobe, apex dentate dorsally (Fig. 34); aedeagus asymmetrical, long, tubular, with apex curved dorsally in lateral view (Fig. 35), 2 membranous spines on caudal margin of curved apex, spines unequal in length, one NIELSON: NEW NEOTROPICAL LEAFHOPPERS 335 below the other and not originating on the same median sagittal line (Fig. 36); gonopore terminal; style very long, nearly reaching apex of aedeagus, narrow except for base and expansion subapically (Figs. 37, 38); plate long, configuration very similar to other spe- cies in the genus, profusely setose (microse- tae) (Fig. 39). HOLOTYPE (male): BRAZIL. Sao Paulo, Salesopolis county, Estaceo Biologica de Bo- racela. Cloud Forest, 850 m, 23.11.1969, W. J. Knight. B.M. 1970-484 (BMNH). REMARKS: P. bispinosa is the largest known species. It is similar in aedeagal characters to denticulata, but it is larger, lacks the serrate flange, and has a well-developed caudoventral process on the pygofer. ACKNOWLEDGMENTS The material described herein was kindly fur- nished by the generosity of Dr. W. J. Knight, British Museum (Natural History) (BMNH), and Dr. Michel Boulard, Museum National d'Histoire Naturelle, Paris. I thank Jean Stanger for her fine illustrations and Dr. James P. Kramer, U.S. National Museum, Washington, and Dr. Paul W. Oman, Oregon State University, Corvallis, for reviewing the manuscript. LITERATURE CITED NIELSON, M.W. 1982. A revision of the subfamily Coelidi- inae (Homoptera: Cicadellidae). IV. Tribe Coelidiini. Pacific Insects Monogr. 38. 318 pp., 1,104 figs. MONTANE INSULAR BUTTERFLY BIOGEOGRAPHY: FAUNA OF BALL MOUNTAIN, SISKIYOU COUNTY, CALIFORNIA Arthur M. Shapiro! ABSTRACT. —Ball Mountain is an isolated, mostly heavily forested peak reaching the subalpine zone (2,330 m) in eastern Siskiyou County north of Mt. Shasta, California. It supports a rich fauna of at least 68 butterfly species showing affinities to the faunas of the Trinity Alps and Eddies, the Warners, and the Cascades. Rare or endemic entities include Speyeria mormonia, Lycaena heteronea gravenotata, and melanic forms of Speyeria atlantis and Agriades “glandon.” Several zones of intergradation or hybridization impinge on the fauna as well. Both the physical and biotic geography of northern California are very complex. The jumbled terrain of the Kiamath-Trinity- Siskiyou upland and the volcanic southern Cascades provides rapid climatic gradients that are reflected in the plant communities of the region. Most of these communities are still inadequately characterized, and several important areas are poorly known even froma floristic standpoint. The butterfly faunas of northern California were extremely poorly documented prior to the 1970s, with only one major paper (Williams 1909) and scattered specimens in museums, often inadequately labeled. From 1976 through 1980 a major ef- fort was mounted to document the butterfly fauna of the Trinity Alps and the Eddies (Shapiro, Palm, and Wcislo 1981) in the hope of using these faunas to test some historical scenarios advanced by botanists to account for the origins of the subalpine and alpine biota of the Sierra Nevada range. Although no defini- tive tests of those scenarios emerged, this study uncovered so much unanticipated com- plexity (along with anticipated Klamath en- demism) in the Trinity-Eddy faunas that it has been continued at several sites which, by virtue of unique location, topography, or veg- etation seemed most likely to provide impor- tant information on the biogeographic history of the northern California butterfly faunas. An especially rewarding site is Ball Mountain, Siskiyou County. Ball Mountain is one of only three peaks between Mount Shasta and the Oregon bor- der, east of U.S. Highway 5, to reach above Department of Zoology, University of California, Davis, California 95616. 2,270 m. The highest of them, Goosenest, at 2,812 m, is a recent Cascade volcano with a poorly vegetated lava cone, though it does have some rare alpine plants (e.g., Hulsea nana Gray, Compositae, which it shares with two other recent volcanoes, Mts. Lassen and Shasta, and with the nonvolcanic Mt. Eddy). The other two, Willow Creek Mountain (2,676 m) and Ball Mountain, (2,330 m), are only 8 km apart and share-a common base and access by road. Both are older, Pliocene volca- noes, mostly basaltic (Ball) or andesitic (Wil- low Creek). Ball Mountain has two old vents marked by pyroclastic jumbles; the higher of these bears a fire watchtower. There is no evidence of recent (Holocene) volcanic impact on the vegetation of the mountain, which has presumably evolved to its present state through the Pleistocene and thereafter. Ball Mountain and Willow Creek Mountain rise fairly gradually from a rolling volcanic upland, the Little Shasta country, to the west (largely Tertiary flows, with altitudes from 750—1,200 m); Ball Mountain, the more easterly, drops off abruptly to the Quaternary lake-bed allu- vial plain near Macdoel, ca 1,275 m (Fig. 1). Although Willow Creek Mountain is the higher of the two, it is more continuously forested and has a smaller variety of habitats and less access by road than Ball Mountain. Access to both is provided by USFS road 47NO3, the extension of York Road, which near Lodgepole Station turns northeastward, whereas road 46N11 goes due south to Willow Creek Mountain and 46N10 goes along the flank of Ball Mountain, looping around Little 336 April 1986 SHAPIRO: BUTTERFLY BIOGEOGRAPHY 337 Fig. 1. Detail of Ball Mountain area from USGS 15’ topographic series, Macdoel quadrangle, 1954. Contour interval 40 feet (12.1 m). Shasta Meadow and Martin Dairy Camp- ground as 46N09; another spur leads to the lookout. Access for hikers and heavy-duty ve- hicles is also possible from Ball Mountain Road (unnumbered), which is paved beyond Table Rock (1,130 m) but becomes nearly im- passable to most vehicles thereafter. This road joins the USFS roads south of Martin Dairy. The entire area is mapped on the USFS map of the Klamath National Forest, Goosenest Ranger District. The best detail is on the 1968 version. The Ball Mountain lookout is R6.E, T45.N. The entire area reported on in this paper can be collected comfortably by two people in one day, or by one person in two days, with a vehicle. VEGETATION The vegetation of Ball Mountain is appar- ently undescribed in the botanical or forestry literature. Like many mountains in northern California, it provides a “telescoped” se- quence of Merriam’s “Life Zones,” such that one may drive from “Upper Sonoran’ to “Subalpine’” in less than 20 km (this very short linear distance facilitates altitudinal migration of valley and foothill butterflies to the high montane meadows). The general zonation of forest vegetation on Ball Mountain and the platform on which it sits more or less corresponds to the outline provided by Rundel, Parsons, and Gordon (1977) for the California portion of the Cas- cades. The lowest elevations, representing an extension of the Little Shasta country, have an open woodland dominated by shrubby forms of Oregon Oak (Quercus garryana Dougl.), with juniper (Juniperus occidentalis ssp. occi- dentalis Hook., near its southwestern limit) as an associate. This is quickly replaced by a mixed conifer association beginning at just over 900 m, dominated by yellow pine (Pinus ponderosa Laws.), with incense cedar (Calo- 338 GREAT BASIN NATURALIST Vol. 46, No. 2 Fig. 2. Rare butterflies from Ball Mountain, dorsal surfaces: a, Speyeria atlantis (melanic form), male, viii. 10.83; b, same (normal form), male, vii.3.85; c, same (melanic form), female, viii.10.83; d, same (melanic form), female aberration, ix.5.85; e, Speyeria mormonia, 2 females, ix.5.85; f, Euphydryas editha, male, vi.12.85; g, Euphydryas chalcedona, male, vi.12.85; h, Agriades “glandon,” male, vii.3.85; i, same, female, vii.3.85; j, Hesperia harpalus, male, ix.5.85; k, same, light female, ix.5.85; 1, same, dark female, ix.5.85. cedrus decurrens Torr.) and white fir (Abies concolor Lindl.) locally abundant. At about 1,450 m lodgepole pine (Pinus contorta ssp. murrayana Grev. & Balf.) first appears in cold and poor sites and quickly becomes dominant; above 1,650 m it is joined by Red Fir (Abies magnifica var. shastensis Lemm.) and west- ern white pine (Pinus monticola Dougl.), and this association continues nearly to the sum- mit. Within it are extensive sedgy and grassy meadows ranging from dry to wet and rimmed by quaking aspen (Populus tremuloides Michx.), many of which are uncharacteristi- cally large for the species. These are among the most diverse sites for both herbs and but- terflies. The area surrounding both summits supports a virtually pure stand of whitebark pine (Pinus albicaulis Engelm.). This sub- alpine forest is characteristically open, with many herbs and shrubs in the understory; the shrubs, which include sagebrush (Artemisia tridentata Nutt.) and Haplopappus bloomeri Gray, predominate among the pyroclasts, whereas herbaceous perennials (Monardella odoratissima ssp. pallida Epl., Eriogonum umbellatum Torr., etc.) cover the light, sandy soils elsewhere. Understory vegetation in the montane conifer forests is extremely undiverse, with extensive areas dominated by pine-mat man- zanita (Arctostaphylos nevadensis Gray) in successional sites. The greatest herbaceous April 1986 SHAPIRO: BUTTERFLY BIOGEOGRAPHY 339 TABLE 1. Occurrence of species on Ball Mountain on collecting dates, 1983 and 1985. vi.12.85 vii.3.85 vii.15.83 viii.10.83 viii.10.85 ix.5.85 ix.7.83 Papilio zelicaon Lucas x Papilio rutulus Lucas x Papilio eurymedon Lucas Parnassius clodius Men. ssp. x Neophasia menapia Feld. & Feld. x x x Pontia beckerii Edw. x x ; Pontia occidentalis Reak. Pieris napi L. ssp. Pieris rapae L. Colias eurytheme Bdv. Colias philodice eriphyle Edw. Colias eurytheme X philodice Anthocharis sara sara Lucas Euchloe ausonides Lucas Coenonympha “tullia” eryngii H.Edw. x x x x Danaus plexippus L. Limenitis lorquini Edw. x x x x Adelpha bredowii californica Butl. Vanessa virginiensis Dru. Xx xX Vanessa cardui L. x x Vanessa annabella Field x x Precis coenia Hbn. Nymphalis californica Bdv. x Nymphalis milberti furcillata Say x Nymphalis antiopa L. x x Polygonia faunus rusticus Edw. Polygonia zephyrus Edw. x x Phyciodes campestris Behr x Phyciodes mylitta Edw. X x x x x x Chlosyne hoffmanni segregata B. & McD. x Euphydryas chalcedona nr. wallacensis Gund. x Euphydryas editha nr. edithana Strand x Boloria epithore Edw. x x x x Speyeria coronis Behr (snyderi-simaethis blend zone population) Speyeria zerene conchyliatus Comst. Speyeria callippe nr. rupestris Behr Speyeria egleis nr. oweni Edw. x Speyeria atlantis Edw. (dodgei- melanic endemic) x x x x x x x Speyeria mormonia Bdv. ssp. X Speyeria hydaspe purpurascens H.Edw. x x x x x Satyrium saepium Bdv. x Mitoura nelsoni Bdv. x Mitoura spinetorum Hew. x Incisalia fotis nr. mossii H.Edw. x Incisalia eryphon Bdvy. x x Lycaena arota Bdv. x X Lycaena heteronea gravenotata Klots Xx x Lycaena xanthoides Bdv. - editha Mead intergrades x Lycaena gorgon Bdv. x Lycaena hellodies Bdv. x Lycaena nivalis Bdv. ssp. Plebeius “idas” ricei Cross-anna Edw. intergrades Plebeius saepiolus Bdv. x Plebeius icarioides Bdv. ssp. x Plebeius acmon Westw. & Hew. Plebeius lupini Bdv. x x Agriades “glandon Prun.” ssp. Everes amyntula Bdv. x x x Glaucopsyche piasus Bdv. x Glaucopsyche lygdamus nr. columbia Skin. x x xX nl oc ~ OO! os Pd «MM Ox epdekd ~ KK ~“ x ~~ eK KKK OK Mm OK mes ee > ~ OM Oe Oe os act “ 340 Table 1 continued. GREAT BASIN NATURALIST Vol. 46, No. 2 vi.12.85 vii.3.85 vii.15.83 viii.10.83 viii.10.85 ix.5.85 ix.7.83 Celastrina argiolus echo Edw. x Ochlodes sylvanoides Bdv. x x x x x Polites sonora Scud. x x x Hesperia “comma complex” x x x Hesperia juba Scud. x x x Pyrgus ruralis Bdv. x Pyrgus communis Grote x Erynnis icelus Scud. & Burg. x Erynnis propertius Scud. & Burg. Xx X x x x Total species recorded’ 36 32 26 25 25 32 17 ‘Not counting Colias hybrids as a species. diversity seen is at Little Shasta Meadow, which has a great variety of Composites and other showy flowering species. It has a light, sandy soil and supports many of the species also found at the summit in the subalpine forest. There are no true bogs, but the wetter meadows are filled with sedge peat. California pitcher plant (Darlingtonia californica Torr.), which is characteristic of boggy meadows in the ultrabasic Trinities and Eddies, is absent. Because the meadows are grazed seasonally by livestock, some herbaceous species may have been lost. Most of the meadows do not display severe sequelae of overgrazing, how- ever. Important nectar sources for collecting oc- cur primarily along the roads and on the meadows. They are very spotty, resulting in high concentrations of butterflies in very small areas. On 5 September 1985, for exam- ple, over 30 species (several hundred individ- uals) were seen in a walk from Martin Dairy Campground to Kuck’s Cabin, but it was com- mon to see nothing but Speyeria zerene in 1-km stretches where no flowers were avail- able. Among the most important nectar sources are Monardella odoratissima, Haplo- pappus bloomeri, Eriogonum spp., Cirsium vulgare (Savi)Ten., Aster spp., and Chryso- thamnus spp. FAUNISTICS AND PHENOLOGY Table 1 presents a complete itemization of species seen on each of the three days in 1983 and four in 1985, when the mountain was collected thoroughly. Spring was late and cold in 1983, with very heavy and persistent snow pack. Spring was very early, warm, and dry in 1985, with snow completely gone by early June. The early summer was hot and dry and the late summer and autumn cold and wet, culminating in heavy snow to the 1,500 m level on 8 September. The two years of study thus embrace very different conditions and probably reflect accurately the amount of phe- nological variation to be expected in the Ball Mountain fauna. Most of the fauna is uni- voltine. The only species-definitely having at least two broods are Pontia occidentalis, Phy- ciodes mylitta, and Lycaena helloides among residents and Pieris rapae, Pontia beckerii, Colias eurytheme and philodice, Coeno- nympha “tullia” eryngii, and Plebeius acmon among species whose ability to overwinter on Ball Mountain is strongly in doubt. A single late individual of Papilio zelicaon has been taken that might represent a rudimentary sec- ond brood (there is a late season flight at low elevations, and this is a hilltopping species). The single record of Pyrgus communis may also represent a fly-up. Most of the summer univoltines have very long flights, those of most Speyeria including nearly the entire season. In S. coronis, males and females emerge early and mate; males then apparently die, but females disappear for several weeks in estivation then reappear; they may be common on yellow Composites in September. The hibernating Nymphalines (genera Nymphalis, Polygonia, and Vanessa) fly in both spring and autumn. Of them, only V. annabella may be partially double-brooded on the mountain. Hesperia juba has the same phenology as the Nymphalines and is sus- pected of hibernating as an adult also (Shapiro 1979). The meadow flora and fauna peak early, with many species disappearing by mid-Au- April 1986 gust. On Little Shasta Meadow the only plants in flower by early September are Polygonum douglasii var. austinae Jones and a few Asters. The overall butterfly phenology is unusual, with little change in the number of species over the entire season but peaks at both ends and a trough in mid-summer. This pattern, albeit weak, is nearly the inverse of the mon- tane pattern shown in the Sierra Nevada (cf Donner Pass, Shapiro 1975: 189). Comments follow on the most unusual ele- ments in the fauna. Systematic order in Table 1 and the text follows Dornfeld, 1980, the geographically closest faunistic treatment in the literature. Pieris napi L. ssp.—One fresh male col- lected vi.12.85 at the edge of Little Shasta Meadow. It is very heavily marked and re- sembles first-brood marginalis Scud. from the north coast. There is an endemic ‘napi” in the Warner Mountains that has a similar first brood and a lightly marked “pallidissima” B. & McD. second brood and occurs on boggy meadows. The affinities of this isolated napi population need to be clarified, because the biogeography of the complex is very difficult in northern California (Geiger and Shapiro 1986a, in press). Colias .—Agricultural, alfalfa-based popu- lations as high as the end of York Road are hybrid swarms involving C. eurytheme Bdv. and C. philodice eriphyle Edw. Both species and hybrids are common on all the meadows and even to the summit most of the season. There is definitely breeding on Trifolium spp., but the phenotypes of spring animals suggest colonization from below each year, rather than overwintering in situ. By Septem- ber nearly all the clover on Little Shasta Meadow is senescent. Anthocharis sara sara Lucas.—Frequent along roadsides near Little Shasta Meadow. These butterflies have pure white: ground color and are indistinguishable from montane sara from the Trinity Alps; they are extremely distinct from both yellow Sierran stella Ed- wards and from the yellow-tinged race from the Warners, and genetically they are identi- cal to Trinity and North Coast Range sara (Geiger and Shapiro 1986b, in press). Euchloe ausonides Lucas.—These dense, montane-meadow populations behave as de- scribed by Dornfeld (p. 51) for Oregon ones, SHAPIRO: BUTTERFLY BIOGEOGRAPHY 341 flying low above sedges and forbs in the wettest parts of the meadows. This is quite unlike the usual behavior of the species in other parts of California. Phyciodes campestris Behr.—These are quite normal campestris , with no tendency to reduction of the pattern as in Sierran montana Behr. They thus resemble Trinity-Eddy spec- imens. Euphydryas .—Only one specimen of each species has been taken, so that subspecific assignments in both cases are very tentative and largely based on Dornfeld’s application of the names. The true identity of edithana Strand remains uncertain. Both subspecific assignments suggest affinity with the War- ners. Speyeri coronis Behr.—These are typical “blend zone” populations like those found in the Eddies, Scott Mountains, and Scott Valley as well as in much of southern Oregon. Speyeria callippe Bdv.—This is a scarce species on Ball Mountain, and the subspecific assignment is based on a short series and must be considered tentative. All our specimens are silvered, and on average they fall between rupestris Behr and topotypical juba Bdv. in facies. Speyeria atlantis Edw.—Common to abun- dant, flying all season. There is a remarkable, apparently endemic melanic form in both sexes—more extreme in the female—with a frequency of over 50% above the junction of roads 47N03 and 46N10, and over 70% at the summit. This form (Fig. 3a) intergrades to more or less normal dodgei Gunder. One specimen taken ix.5.85 has one hindwing aberrant (Fig. 3d). Speyeria mormonia Bdv. ssp.—Outside the Sierra Nevada, this species was previously known in California only from Deadfall Lakes and Mount Eddy (Shapiro, Palm, and Wcislo 1981) and from the Warners. The subspecific identities of these populations are not clear; Shapiro et al. treated the Deadfall-Eddy pop- ulation as an outlier of the Oregon Cascade erinna Edw., but neither it nor the somewhat paler Warner phenotypes precisely matches either Cascadian or Sierran material. Three specimens were collected ix.5.85 on flowers of Haplopappus bloomeri about 0.8 km below Little Shasta Meadow and on the lower por- tion of the meadow itself. They were among 342 Fig. 3. Same as for Figure 2, ventral surfaces. many other fritillaries of three species. All are very small (LFW 20-21.5 mm) and not a precise match for either Deadfall-Eddy or Warner spec- imens. Typical mormonia habitat (wet meadow) is abundant on Ball Mountain, but as in the Ed- dies the species appears to be rare and to emerge remarkably late in the season. The geography of Speyeria mormonia in far northern California promises to shed light on the history of biotic migrations in the Quaternary. Lycaena heteronea gravenotata Klots.— The status of this subspecies name is contro- versial. Ferris and Brown (1981) gloss over any pattern of geographic distribution for spotted and unspotted hindwings in the Rocky Mountains. Dornfeld (1980) finds a def- inite pattern of spotted colonies within a re- gion of generally unspotted ones. In Califor- nia, the spotted morph is known only from GREAT BASIN NATURALIST Ball Mountain (Little Shasta Meadow to sum- mit), nearby Goosenest, Warner Valley (just S. Mt. Lassen), and near Castella (J. F. Emmel, personal communication). These ap- pear to be pure spotted populations, sur- rounded by pure unspotted ones. The host plant on Ball Mountain has not been deter- mined, but a white-flowered Eriogonum that occurs at both Little Shasta Meadow and near the fire tower is suspected. (A member of the “E. nudum complex” is reportedly used at Warner Valley; J. F. Emmel, personal com- munication). There is no adult association with E. umbellatum, such as one sees consis- tently in the Trinities. Figures 4a,b and 5a,b show both sexes. Collectors should be aware of the possibility that further colonies exist near Mount Shasta. The possibility of sibling species cannot be discounted. April 1986 SHAPIRO: BUTTERFLY BIOGEOGRAPHY 343 Fig. 4. Northern California Lycaena, dorsal surfaces: a, L. heteronea gravenotata, male, Ball Mountain, viii. 10.83; b, same, female, Ball Mountain, viii. 10.85; c, L. h. heteronea (?), male, Dry Lake Lookout, Siskiyou Co., viii.9.83; d, same, female, Dry Lake Lookout, viii.9.83; e, L. h. heteronea, male, Winnemucca Lake, Alpine Co., viii.24.83; f, same, female, Winnemucca Lake, viii.24.83; g. Lycaena nivalis, male, Ball Mountain, vii. 15.83; h, same, female, Ball Mountain, vii. 15.83; i, L. nivalis, Cedar Pass, Modoc Co., vi.9.85; j, L. nivalis, male, Winnemucca Lake, Alpine Co., viii.24.83; k, L. nivalis “form 1,” male, Paradise Lake, Marble Mts., Siskiyou Co., vii.4.81; 1, same, female, Paradise Lake, vii.4.81. Lycaena xanthoides Bdv.-L. editha Mead.— Lycaena xanthoides is found in the Central Valley (usually near the Sacramento River), in the San Francisco Bay area, the Transverse Ranges and some areas of southern California, and apparently disjunctly as a series of mon- tane populations from Lake County through the Mendocino Pass—Anthony Peak area, and in the Willamette Valley in Oregon. Lycaena editha is found in the Rockies, the Sierra Ne- vada, and the Cascades as a montane-to-sub- alpine species. From Dunsmuir to Siskiyou Summit and from Gazelle to Ball Mountain and Iron Gate Reservoir occur a series of ap- parently intermediate populations, generally in agriculturalized valleys. The highest eleva- tion of these is the one on road 47N03 near Kuck’s Cabin, of which strays occur as high as Little Shasta Meadow. It also seems to be the easternmost of the blend-zone populations. 344 GREAT BASIN NATURALIST Vol. 46, No. 2 Fig. 5. Same as for Fig. 4, ventral surfaces. Lycaena helloides Bdv.—This is usually con- sidered a weedy lowland species, but here as elsewhere in northern California it seems to have permanent montane-meadow populations and has been seen laying on Polygonum dou- glasii var. austinae in September. Lycaena nivalis Bdv.—Dornfeld (1980: 97, maps 137-138) pointed out that two “forms” of the nominal species L. nivalis occur in Oregon— sometimes sympatrically, sometimes not. Both also occur in northern California. Trinity-Eddy populations (Shapiro, Palm, and Wcislo 1981) are 100% “form 1” (two-toned VHW), and Ball Mountain ones are 100% “form 2” (nearly unicol- orous VHW). Form | has not been rtounu cast us U.S. Highway 5 in California to my knowledge. Form 2 is similar, but not identical, to the form that occurs in the Sierra Nevada. Some individu- als show a tendency toward the endemic, heavily spotted Warner Mountains form. All these phe- notypes are shown in figures 4g-l, 5g-]. At Little Shasta Meadow eggs are laid on Polygonum dou- glasii var. austinae. Trinity-Eddy form 1 use P. spergulariaforme Meissn., a closely related spe- cies. As with L. heteronea, sibling species are strongly suspected. Agriades “glandon Prun.” ssp.—Con- specificity with the European taxa of this | April 1986 group is questionable. As noted by Shapiro, Palm, and Wceislo (1981), northern California populations are considerably darker and more heavily spotted beneath than Sierran podarce Felder (the usual usage, the type locality being simply “California’). The type of Boisduval’s nestos, from Oregon, should be examined to determine its consubspecificity with these popu- lations. Ball Mountain “glandon” average darker and more heavily spotted beneath than any other North American population known to me and seem to represent the extreme end of acline (Figs. 2,3h,i). Hesperia “comma complex.’ —Again, the use of the name comma L. seems questionable. On the other hand, it is unclear what strictly Nearc- tic names apply to our handful of Ball Mountain specimens, which vary in complex ways be- tween the taxa oregonia Edwards and harpalus Edwards; there is too little material (mostly from thistles along the roads, where Hesperia is enor- mously outnumbered by Ochlodes sylvanoides Bdv.) to say whether the phenotypes are altitu- dinally stratified as in the Trinities, or scrambled as on Mount Eddy. The range of variation is shown in figures 2, 3j-. DISCUSSION This fauna of 68 species is remarkably rich for a forested, isolated mountain area that is surrounded by unforested lowlands. The ma- jor elements of biogeographic interest are enumerated below. 1. Endemics.—These include the melanic forms of Agriades “glandon” and Speyeria at- lantis, and probably Speyeria mormonia; Ly- caena heteronea gravenotata is a near-en- demic (actually, most of the Speyeria species on Ball Mountain show local peculiarities, which, however, are less conspicuous than the melanism of S. atlantis). 2. Regional Rarities .—Species rare and lo- cal in northern California that occur in the study area include Incisalia fotis, Mitoura spinetorum, Lycaena arota, and Erynnis icelus (L. arota is relatively common in the Trinities but flies quite late, as here; it seems rare elsewhere in the region). Polygonia faunus rusticus is rare throughout its range; Glaucopsyche piasus nearly so. 3. Intermediate or Transitional Popula- tions.—Lycaena xanthoides—editha; Hespe- SHAPIRO: BUTTERFLY BIOGEOGRAPHY 345 ria “comma complex’; Colias hybrid swarms (common in agricultural alfalfa but not other- wise recorded in montane meadows region- ally). 4. Regionally Common Species, Rare on Ball Mountain.—These include Neophasia menapia, Phyciodes campestris, Chlosyne hoffmanni, Satyrium saepium, Plebeius “idas, Everes amyntula, Glaucopsyche lyg- damus, the Hesperia “comma complex,” and Pyrgus communis . 5. Range Extensions with Westward Affinities. —Anthocharis sara. 6. Range Extensions with Eastward (Warner Mountains) Affinities.—Possibly both Euphydryas ; possibly Lycaena nivalis . 7. Absences. These deserve special enu- meration; they are species that, on a regional basis, would be considered likely on Ball Mountain but have not been found. 7a. Alpine and Barren-Ground Species .— Suitable habitats are clearly not present for Papilio indra Reak., Parnassius phoebus ster- nitzkyi McD., Pontia sisymbrii Bdv., and Eu- chloe hyantis Edw. Callophrys lemberti Tilden and Philotes battoides Behr may also fit in this category, though potential host plants are present. 7b. Species of Special Habitats Apparently Absent or Too Isolated.—Lycaena mariposa Reak. is common at the edges of boggy mead- ows in the Lodgepole Pine zone farther west (Trinities and Eddies, Scott Mountains). Per- haps the meadows on Ball Mountain are insuf- ficiently boggy. The host plant remains un- published but is reported to be Vaccinium spp. (G. Pratt, in litt.) This plant is not recorded on Ball Mountain. Euphyes vestris Bdv. has been found in several isolated boggy meadows (e.g., Scott Mountain Summit Bog). Its host plant, Cyperus, occurs near Little Shasta Meadow, but the colony is perhaps too small and isolated to support the skipper. 7c. No Apparent Explanation .—These in- clude Chlosyne palla Bdv., Satyrium fuligi- nosum Edw., S. sylvinus Bdv., S. californica Edw., Incisalia iroides Bdv., Philotes enoptes Bdyv., Thorybes mexicana nevada Scud./ aemilia Skin., and Polites sabuleti Bdv. Sev- eral of these are regionally rare or local (fuligi- nosum, sylvinus, californica, iroides ). But Po- lites sabuleti is the single most common butterfly in the Trinities and Eddies (Shapiro, 346 GREAT BASIN NATURALIST Vol. 46, No. 2 TABLE 2. Composition by family of some regional butterfly faunas. Family Trinity Alps* Mt. Eddy* Crater Lake? Ball Mountain Papilionidae a 5 4 4 Pieridae 14 i fe) fe) Satyridae 4 3 4 1° Danaidae 1 1 1 1 Nymphalidae 31 25 29 24 Riodinidae 1 1 0 0 Lycaenidae 32 27 26 Pl Hesperiidae 25 11 11 8 Totals 115 80 80 68 “Shapiro, Palm & Wcislo 1981. Tilden & Huntzinger 1978. “Oeneis nevadensis may also occur. Palm, and Wceislo 1981). It occurs on the N ACKNOWLEDGMENTS slope of Mt. Shasta and, at low density, in the agricultural lands both east and west of Ball Mountian; there appears to be no population adapted to montane meadows in the Little Shasta Country. This is not surprising in one sense: there is no place known where both Ochlodes sylvanoides and Polites sabuleti are common in northern California; their abun- dances tend to be inversely correlated, and sylvanoides is the commonest skipper on Ball Mountain. 7d. Biennials.—Oeneis nevadensis Feld. & Feld. has a two-year life cycle in much of its range, flying only in even-numbered years. Thus, if it occurs (in the lower montane conif- erous forest) on Ball Mountain it would have been missed by sampling in 1983 and 1985. It is the only known biennial butterfly in the region. A final question is that of the faunal balance. As revealed in Table 2, the family breakdown of the Ball Mountain fauna is basically consis- tent with other regional faunas, though slightly impoverished in Hesperiidae. What does not emerge from these data is the enor- mous biomass of the genus Speyeria, whose members are overwhelmingly dominant over virtually all habitats on Ball Mountain. This is not the only such place in northern California: Anthony Peak (Mendocino County), for exam- ple, is another. But even the casual visitor cannot help but notice how all other butter- flies appear rare in comparison to the large fritillaries. Since all of them presumably feed on the genus Viola, which is neither unusu- ally conspicuous nor unusually diverse on Ball Mountain, their abundance poses an ongoing ecological problem. Some studies on Ball Mountain have been carried out under California Agricultural Ex- periment Station project CA-D*-AZO-3994- H, “Climatic Range Limitation of Phy- tophagous Lepidopterans.” I cordially thank my fellow collectors on the mountain: Bill Overton, Adam Porter, Virginia Pickles, Ce- cile La Forge, Hansjurg Geiger, and Iris Bug- mann, as well as Dr. John Emmel for informa- tion on Lycaena heteronea gravenotata and G. Pratt for information on L. mariposa. Mr. More, occupant of the last house on York Road, was kind enough to pull a University of California pickup truck out of a sand pit in 1985. Had he not done so, there would have been no 1985 data. LITERATURE CITED DoRNFELD, E. J. 1980. The butterflies of Oregon. Timber Press, Forest Grove, Oregon. 276 pp. Ferris, C. D., AND F. M. Brown. 1981. Butterflies of the Rocky Mountain States. University of Oklahoma Press, Norman. 442 pp. GEIGER, H. J.. AND A. M. SHapiro. In press. Elec- trophoretic evidence for taxonomic relationships within the western North American “Pieris napi’. J. Res. Lepid. ____. In press. Electrophoretic evidence for speciation within the nominal species Anthocharis sara (Pie- ridae). J. Res. Lepid. RUNDEL, P. W., D. J. PARSONS, AND D. T. GorpDon. 1977. Mon- tane and subalpine vegetation of the Sierra Nevada and Cascade ranges. Pages 559-600 in M. G. Barbour and J. Major, eds., Terrestrial vegetation of Califor- nia. John Wiley, New York. 1,002 pp. SHAPIRO, A. M. 1975. The temporal component of butter- fly species diversity. Pages 181-195 in M. L. Cody and J. M. Diamond, eds., Ecology and evolution of communities. Belknap Press, Harvard, Cam- bridge, Massachusetts. 545 pp. April 1986 SHAPIRO: BUTTERFLY BIOGEOGRAPHY 347 . 1979. Does Hesperia juba (Hesperiidae) hibernate — TILDEN, J. W., AND D. H. HuNTZINGER. 1978. The butter- as an adult? J. Lepid. Soc. 33: 258-260. flies of Crater Lake National Park, Oregon. J. Res. SHAPIRO, A. M., C. A. PALM AND K. L. WcIsLo. 1981. The Lepid. 16: 176-192. ecology and biogeography of the butterflies ofthe | WILLIAMS, F. X. 1909. The butterflies and some of the Trinity Alps and Mount Eddy, northern Califor- moths of the Mount Shasta region. Ent. News 20: nia. J. Res. Lepid. 18: 68-152. 62-75. COMPARATIVE HABITAT AND COMMUNITY RELATIONSHIPS OF ATRIPLEX CONFERTIFOLIA AND SARCOBATUS VERMICULATUS IN CENTRAL UTAH Jack D. Brotherson’, Lars L. Rasmussen’, and Richard D. Black’ ABSTRACT. —Thirty-four study sites were established in shadscale (Atriplex confertifolia [Torr. & Frem. | Wats.) and greasewood (Sarcobatus vermiculatus [Hoov. Torr. in Emory) communities bordering Utah Lake in central Utah. Differences in species composition, vegetation, and soil characteristics were assessed. Significant differences in soil factors between the two communities were found for sand, calcium, manganese, zinc, and copper. Soluble salts and sodium concentrations were generally higher in the greasewood type, but differences were not significant. Major differences were found in understory species, with burr buttercup (Ranunculus testiculatus Grantz) showing signifi- cantly greater cover in the shadscale community and cheatgrass (Bromus tectorum L.) showing significantly greater cover in the greasewood community. Shadscale (Atriplex confertifolia [Torr. & Frem.] Wats.) and Greasewood (Sarcobatus vermiculatus [Hook.] Torr. in Emory) are dominants of plant communities that cover vast areas of the Great Basin and are thus important components of our western range- lands. Recent research on these species has considered soil moisture relationships (Bran- son et al. 1976), evolution (Stutz 1978), phe- nology (roundy et al. 1981, Everett et al. 1980), faunal associates (Csuti 1979, Feld- hamer 1980), grazing effects (Fetcher 1981), physiology (Caldwell et al. 1977), production (Van Epps et al. 1982), and successional rela- tionships (Vasek and Lund 1980, Wallace and Romney 1980). However, there is a lack of information from central Utah concerning dif- ferences in community and habitat require- ments of shadscale and greasewood. The pur- pose of this study was to compare habitats of shadscale and greasewood dominated sites in central Utah. Such information is valuable when attempting to manage those species in relationship to their use as forage for sheep on our winter ranges. STUDY AREA Thirty-four study sites were sampled in shadscale and greasewood communities bor- dering Utah Lake, Utah (Fig. 1). Fifteen sites were studied in shadscale communities and nineteen in greasewood communities. Monee A) 4 A Greasewood sites @ Shadscale sites ie a” a ™ Study site EEE sce UTAH aie, | | Pee Fig. 1. Map showing the location of the 34 study sites near Utah Lake in central Utah. Shadscale communities are generally lo- cated on the west side of Utah Lake below the sagebrush zone and above or parallel to the ‘Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. Ellensburg District Office, Soil Conservation Service, Ellensburg, Washington 98926. 348 April 1986 greasewood zone. The overstory component of this community is dominated by shadscale. The understory is composed primarily of two introduced species, burr buttercup (Ranun- culus testiculatus) and cheatgrass (Bromus tectorum). Greasewood communities are found around the entire perimeter of Utah Lake but are best developed along the western shore. Overstory in this community is dominated by greasewood, and the understory is dominated by burr buttercup and cheatgrass. Both com- munity types serve as winter range and spring lambing areas for several thousand head of sheep each year. Climatic conditions at the study sites are characterized by hot dry summers and cold winters. Average annual precipitation varies from 190 to 290 mm, with 60% falling in the winter and early spring months. The hottest month of the year is July, with an average of 33 C; the coldest month is January, with an aver- age of 3 C. The frost-free period for the area ranges from 132 to 170 days (Swenson et al. 1979). METHODS Study areas were selected to represent a range of environmental conditions in shad- scale and greasewood communities of central Utah. Once a site was located, a 10 x 10 m macroplot (0.04 ha) was randomly located within each area. Elevation, percent slope, slope position, and soil erosion were noted for each site. Each plot was subsampled with 20 0.25m° quadrats (microplots) stratified across the macroplot in five rows of four quadrats each. Data were taken during May and June 1980. Total living cover, plant cover by life form, litter, exposed rock, and bare ground were estimated at each microplot following ocular procedures suggested by Ostler (1980). Cover of individual plant species encountered was also estimated using cover class categories suggested by Daubenmire (1959). All species occurring within a macroplot but not in any microplots were listed and given a percentage cover value of 0.01. All species encountered were classified as to life form, longevity, and whether native or introduced in the Utah flora. BROTHERSON ET AL.: PLANT COMMUNITY RELATIONSHIPS 349 Three soil samples were taken in each macroplot (from opposite corners'and the cen- ter) from the top 20 cm of soil and later com- bined for laboratory analysis. This depth was considered adequate based on Ludwig's re- sults (1969), which showed that the surface decimeter of soil yields 80% of the information useful in correlating plant response with con- centrations of essential mineral nutrients in the soil. Studies by Holmgren and Brewster (1972) also showed that greater than 50% of the fine roots are found in the top 15 cm of soil profiles in desert shrub communities in west- ern Utah. Soil samples were analyzed for texture (Bouyoucos 1951), pH, soluble salts, mineral composition, and organic matter. Soil pH was determined with a glass electrode pH meter. Soluble salts were determined with a Beck- man electrical conductivity bridge. Ex- changeable calcium, magnesium, potassium, and sodium were extracted from soils with DTPA (diethylene triamine-penta-acetic acid) (Lindsay and Norvell 1969). A Perkin Elmer Model 403 atomic absorption spectrophoto- meter was used to determine individual ion concentrations (Isaac and Kerber 1971). Phos- phorus was extracted with sodium bicarbon- ate (Olsen et al. 1954). Nitrogen analysis was made using macro-Kjeldahl procedures (Jack- son 1958). Organic matter was estimated from total carbon using methods described by Al- lison et al. (1965). Means, standard deviations, and coeffi- cients of variation were determined for each biotic or abiotic variable across the 34 sam- pling plots. Prevalent species were deter- mined following Warner and Harper (1972) on the basis of cover values. One-way analysis of variance was used to detect significant differ- ences between the two communities with ref- erence to 18 different soil variables. Student's t-test was used to detect significant differ- ences in site characteristics and biotic factors between the two communities. Taxonomic determinations for all plant species included in our study follow Arnow et al. (1980). RESULTS AND DISCUSSION Significant differences between factors of the shadscale and greasewood communities near Utah Lake existed for only two of the 390 GREAT BASIN NATURALIST Vol. 46, No. 2 TABLE 1. Means, standard deviations, and coefficients of variation of general site characteristics for shadscale and greasewood communities around Utah Lake (N = 34). Shadscale Greasewood Site characteristics Mean S.D. (CAVE Mean S.D. CAVE Elevation 4524.53 17.94 0.003 4530.75 45.75 0.01 Percent slope se 1.44 1.97 0.95 1.76 1.28 “Slope position* 1.47 0.83 0.57 2:15 0.93 0.43 >Erosion 0.00 0.00 0.00 0.05 0.22 4.40 Percent litter cover** 5.20 3.90 0.75 2.95 4.33 1.47 Percent exposed rock 0.00 0.00 0.00 0.02 0.07 3.39 Percent exposed soil 16.07 14.65 0.91 12.56 11.33 0.90 Slope position is defined as 1 = top of slope, 2 = midslope, 3 = bottom of slope. >The erosion index runs from 0 to 3, with 0 indicating no erosion and 3 heavy. *Significant differences between means at 0.10 level. **Significant difference between means at 0.05 level. TABLE 2. Means, standard deviations, and coefficients of variation of cover of prevalent species in shadscale and greasewood communities around Utah Lake (N = 34). Shadscale Greasewood Species Mean S.D. C.V. Mean S.D. C.V. Atriplex confertifolia 16.80* 8.71 0.52 0.24* 0.89 3.70 Bromus tectorum 10.77* 9.25 0.86 35.38* 27.69 0.78 Cardaria draba 0.00 0.00 0.00 5.68* 14.74 2.60 Ephedra viridis 2.01* 6.83 3.40 0.00 0.00 0.00 Halogeton glomeratus 1.89* 4.23 2.24 1.24 : 5.48 4.42 Hordeum leporinum 0.00 0.00 0.00 5.62* 14.74 2.62 Kochia americana 5.67* 8.05 1.43 0.00 0.00 0.00 Kochia scoparia oe 6.27 3.58 ean 13.13 1.76 Lepidium perfoliatum 8.71* 6.86 0.79 7.82* 8.45 1.08 Ranunculus testiculatus 60.77* 25.78 0.42 23.25* 25.99 1.11 Salsola iberica 0.01 0.04 4.00 3.26* 6.55 2.00 Sarcobatus vermiculatus 22.2% Salil 1.40 28.88* 14.62 0.51 Sitanion hystrix 0.99 1.56 1.58 2.39* 5.07 m2, Suaeda calceoliformis 0.01 0.03 3.00 3.75* 13.11 3.50 Suaeda torreyana 1.35* 2.40 Waa 2.69* 6.10 DoT *Prevalent species eight general site variables considered (i.e., slope position and percent cover, Table 1). Shadscale had greater litter cover and tended to occupy upper slope positions, whereas greasewood was found at midslope positions. Shadscale and greasewood communities had six prevalent species in common (Table 2). Cover of annual plants was 66% and 52% in the shadscale and greasewood communities, respectively (Table 3). These cover values represented 72% and 65%, respectively, of the total living cover of those two communi- ties. The cover values for burr buttercup and cheatgrass were of particular interest. The greasewood and shadscale communities con- sidered herein had been heavily impacted for many years by domestic grazing animals (Table 3). Such sustained overuse would open up areas within the community and allow these introduced species to invade and de- velop high cover values. The area is used as late winter and early spring sheep range, and sustained overuse is the suggested cause for deteriorated range condition at most of the sites (Brotherson and Evenson 1982). Grazing estimates were based on the condition of plants and soils and their relative responses to grazing (Stoddart et al. 1975). The shadscale community had significantly greater burr buttercup cover and significantly less cheatgrass cover than the greasewood community. Burr buttercup (Table 2) con- tributed 60% cover, whereas cheatgrass con- tributed only 10% in shadscale plots. Con- versely, the greasewood community con- tained significantly greater amounts of cheat- grass. Other annual forbs, (Belvedere sum- mer cypress, Kochia scoparia, Haloge- April 1986 BROTHERSON ET AL.: PLANT COMMUNITY RELATIONSHIPS 301 TABLE 3. Means, standard deviations, and coefficients of variation for biotic factors in shadscale and greasewood communities (N = 34). Shadscale Greasewood Mean S.D. C.V Mean S.D. C.V. Total living cover 79.4 14.50 0.18 80.7 16.12 0.20 % Shrub cover 14.1 6.00 0.43 20.3 10.13 0.50 % Subshrub cover** 8.4 12.87 1.54 2.9 8.71 2.93 % Perennial forb cover** 0.0 0.00 0.00 3.3 eit 2.23 % Perennial grass cover** 2.2 DIE) 4.91 5.3 8.05 1.53 % Annual grass cover*** 8.4 5.90 0.71 26.2 PAS) 0.81 % Annual forb cover*** 57.7 21.06 0.37 29.3 22.67 0.77 % Total annual cover*** 66.1 19.50 0.30 Oleg, 21.53 0.41 % Cryptogam cover* 9.4 6.61 0.71 6.8 6.85 1.01 Diversity 1.9 0.42 . 0.22 22) 0.48 0.22 Number of species/quadrat 0.5 0.12 0.28 0.5 0.16 0.31 Number of species/stand 9.1 2.71 0.30 10.3 3.21 0.31 Number of native species/stand 4.9 1.92 0.39 Dow 2.93 0.53 Number of introduced species/stand 4.2 1.42 0.34 4.8 1.70 0.36 Percent of flora: Native species 52.5 12.67 0.24 51.7 16.49 0.32 Introduced species 47.5 12.67 0.27 48.4 16.49 0.34 Percent of total cover: Native species** 27.9 19.08 0.68 34.6 Wed 0.50 Introduced species** 72.1 19.08 0.26 65.4 17.37 0.27 Grazing impact 2.5 0.74 0.29 2.5 1.00 0.40 *Significant difference between means at 0.10 level. **Significant difference between means at 0.05 level. ***Significant difference between means at 0.01 level. ton—Halogeton glomeratus, and clasping pep- perweed, Lepidium perfoliatum) showed dif- ferent patterns of distribution. Halogeton and clasping pepperweed were evenly distributed in both communities, whereas Belvedere summer cypress showed much greater cover in the greasewood type. Other grass species—rabbit barley (Hori- dum leporinum) and bottlebrush squirreltail (Sitanion hystrix)—showed patterns similar to cheatgrass. Reasons for these relationships are unknown. Edaphic factors may be par- tially responsible for site selection of the two annuals (burr buttercup and cheatgrass). This is shown in the distribution patterns of the two species when their cover values are plotted against the different soil factors (Fig. 2). As shown, the two species exhibit different pat- terns with respect to percent sand, percent silt, percent fines, pH, nitrogen, calcium, iron, zinc, and copper. The species patterns with respect to the textural classes appear as mirror images of each other; therefore, the relationship is probably due to species inter- actions rather than cause and effect with re- spect to the soil factor itself. The species dis- tribution patterns with respect to pH, nitrogen, calcium, iron, zinc, and copper are more disjunct and, therefore, may suggest cause and effect relationships. When we ex- amine these factors with respect to their dis- tribution between the two community types, the differences in calcium, zinc, and copper are shown to be significantly different. How- ever, the lack of significant differences be- tween the majority of the soil factors mea- sured and the rather large coefficients of variation attached to these same variables sug- gest that the exhibited differences in distribu- tion may be a function of which species first invaded and became established. Once estab- lished, species such as burr buttercup and cheatgrass may be highly competitive to other species. Competitive relationships between these two annuals would certainly follow inva- sion and may, therefore, be responsible for the observed distributions. Burr buttercup has been suspected to be allelopathic (an in- hibitor of seed germination of other species through release of harmful chemicals into the soil) (Buchanan et al. 1978). Further evidence of the competitive nature of burr buttercup can be seen by examining the prevalent spe- cies lists for the two community types (Table 2). The greasewood type contained six preva- lents that exhibited much greater cover values 302 % Cover Ranunculus % Cover Bromus % Cover Ranunculus % Cover Bromus % Cover Ranunculus % Cover Bromus 100 50 100 50 100 50 ty m ‘ 1" 7.94 8.95 9.95 pH 44 106 Mg (ppm X 10) 168 100 50 100 50 GREAT BASIN NATURALIST 28) fs) 7/3) % Silt 177 494 811 Sol. salt (ppm X 10) 100 100 50 100 100 50 50 ; 100 100 50 50 19 40 62 % Clay 12 N (%) 100 100 50 50 ke 137 369 602 Na (ppm X10) 113 240 368 K (ppm X 10) 100 50 100 50 58a 96 % Fines h . 100 50 100 P (ppm) 50 100 50 “AT tl 8-1) 16 Fe (ppm) Vol. 46, No. 2 100 50 -F FF Uk3 SC) SO) % OM 100 50 100 4 mn 77 100 123 Ca (ppm X 100) 50 100 50 -/-_ 16 = = 6 Mn (ppm) 100 50 1.6 44 Zn (ppm) @.7/ 100 16 3.2 49 Cu (ppm) Fig. 2. Cover of Ranunculus testiculatus and Bromus tectorum plotted against major soil gradients of study sites. April 1986 BROTHERSON ET AL.: PLANT COMMUNITY RELATIONSHIPS 303 TABLE 4. Means, standard deviations, and coefficients of variation of observed soil characteristics in shadscale and greasewood communities. Shadscale Greasewood Soil characteristics Mean S.D. C.V. Mean S.D. C.V. Percent sand* 16.53 8.43 0.51 27.33 16.15 0.59 Percent silt 43.20 8.58 0.20 39.90 15.48 0.39 Percent clay 40.27 10.50 0.26 32.78 12.38 0.33 Percent fines* 83.47 8.43 0.10 72.68 16.15 0.22 Percent organic matter 2.57 0.86 0.33 2.66 1.00 0.38 pH 8.06 0.90 0.11 8.21 0.60 0.07 Soluble salts (ppm) 614.20 259.03 0.42 1502.35 2156.21 1.44 *Significant difference at .05 in the means. TABLE 5. Means, standard deviations, and coefficients of variation of soil nutrient in shadscale and greasewood communities. Shadscale Greasewood Soil nutrient Mean S.D. C.V. Mean S.D. C.V. Percent nitrogen 0.11 0.04 0.37 0.13 0.09 0.72 Phosphorus (ppm) 13.70 5.62 0.41 18.60 Lit 0.61 Calcium (ppm)*** 11146.70 518.23 0.05 9062.30 1675.29 0.18 Magnesium (ppm) 786.90 355.39 0.45 607.80 409.31 0.67 Sodium (ppm) 613.10 331.78 0.54 1031.20 1566.45 1.52 Percent Na saturation 4.34 2.20 0.51 8.64 12.14 1.40 Potassium (ppm) 1364.30 772.01 0.57 921.00 598.08 0.65 Iron (ppm) 4.30 0.98 0.23 5.80 3.56 0.62 Manganese (ppm)* 6.60 1.56 0.24 8.70 3.54 0.41 Zinc (ppm)* 0.62 0.29 0.47 1.70 1.75 1.06 Copper (ppm)** 1.30 0.38 0.30 2.20 1.22 0.55 *Significant at .05 level in the means. ** Significant at .01 level in the means. ***Significant at .001 level in the means. on the greasewood sites than on the shadscale Mineral nutrient concentrations in the sites. The shadscale community, on the other hand, had only two prevalents that showed greater cover on the shadscale than on the greasewood sites. Both of these species were shrubs. If burr buttercup is as allelopathic to cheatgrass as it is to other grass species, then burr buttercup should have a competitive ad- vantage. However, further study is needed concerning the factors involved in the distri- bution of these two introduced annuals before the question can be fully answered. Soils from the greasewood community had significantly more sand than soils from shad- scale stands (Table 4). No significant differ- ences were found for percent silt or percent clay in these communities, but, when com- bined, silt and clay (fines) were significantly greater in the shadscale community. We sus- pect that the sifting effects of currents at high water levels of the lake in the distant past, may have been responsible for larger percentages of sands in the greasewood type in this study. shadscale stands were greater than have been previously reported (El-Ghonemy et al. 1980). Sodium, zinc, and nitrogen were ap- proximately 1.2 times greater in our study area, manganese was 3.4 times greater, and copper was 8.1 times greater. Even larger differences were found for phosphorus and iron concentrations, which were approxi- mately 26 times greater in our study area than in the Mojave Desert areas studied by El- Ghonemy et al. (1980). Calcium and magne- sium concentrations combined were 6.8 times greater in our study area. Potassium was the only ion reported to have greater concentra- tions in Nevada than in our study area in Utah. Comparable soil data were not found in the literature for comparison with our results from greasewood sites. Significant differences between shadscale and greasewood communities at the Utah Lake site were found for calcium, manganese, zinc, and copper (Table 5). Calcium was sig- 354 GREAT BASIN NATURALIST Vol. 46, No. 2 % Cover Sarcobatus (6) ro) fo) ro) oO ° ro) fo) a ° ro) ro) a ro) ro) fo) Q ° ro) ro) % 100 100 100 100 100 a fo) 50 50 50 50 50 14 33 52 29 53 78 19 40 62 58a 96: He Sk} SO) % Sand % Silt % Clay % Fines % OM 8 100 100 100 100 100 100 ro ze} fo} 1S) o 9) 3 50 50 50 50 50 50 oO x ss as 100 100 100 100 100 100 a =< o > fo) O50 50 50 50 50 50 794 8.95 9.95 177 494 811 aU .27 .42 15 ~35: 54 27 100 123 1G" 4Zal Ser pH Sol. salt (ppm X10) N (%) P (ppm) Ca (ppm X 100) Zn (ppm) 2 100 100 io ye} (eo) 2 oO 9) S 50 50 fe) Oo AS 100 100 100 100 100 a x o > [e) OF 50 50 50 50 50 44 106 168 137 369 602 113 240 368 5S IW I GQ} 1a 16 Mg (ppm X 10) Na (ppm X 10) K (ppm X 10) Fe (ppm) Mn (ppm) Fig. 3. Cover of Sarcobatus vermiculatus and Atriplex confertifolia plotted against major soil gradients of study sites. April 1986 BROTHERSON ET AL.: PLANT COMMUNITY RELATIONSHIPS 300 TABLE 6. Species preferences indices with respect to measured soil factors. Species Sarcobatus Atriplex Ranunculus Bromus Site factor vermiculatus confertifolia testiculatus tectorum Sand (%) 21.4 17.2 19.6 24.7 Silt (%) 42.5 42.7 44.4 40.2 Clay (%) 35.4 40.3 36.0 34.5 Fines (%) 78.0 83.1 80.4 74.7 Organic matter (%) 2.8 2.8 oS) ei pH 8.2 8.2 8.2 8.0 Soluble salts (%) 1,526.4 720.2 718.4 665.2 Nitrogen (%) 14 a2, ll 13 Phosphorus (ppm) 21.5 14.2 23.4 15.9 Calcium (ppm) 9,343.9 11,116.1 10,729.6 9,498.7 Magnesium (ppm) 649.0 852.7 664.9 620.0 Sodium (ppm) 1,057.8 730.0 631.2 457.1 Potassium (ppm) 987.7 1,333.4 1,384.8 910.3 Iron (ppm) 6.4 4.4 4.0 5.6 Manganese (ppm) 9.3 6.5 6.9 7.9 Zinc (ppm) 1.9 09 50S) 1.7 Copper (ppm) Dae 12 1S 12 nificantly greater in the shadscale commu- nity. Since the shadscale sites occurred at the top of slopes and greasewood sites occupied midslope areas (Table 1), Romney and Wal- lace’s (1980) contention that calcium tends to dominate the soluble cation complex at up- land positions is supported. We suspected that total soluble salts and concentrations of sodium would be significantly greater in the greasewood community (Fireman and Hay- ward 1952). However, significant differences were not observed, although salt concentra- tions were higher at some greasewood sites. Percent sodium saturation was also greater in the greasewood community, but differences between communities was not statistically sig- nificant. This is best explained in terms of slope position. As indicated by Romney and Wallace (1980), sodium should dominate only at the bottom of slopes in closed drainage basins. In our study sites greasewood was not located within a closed drainage basin, but instead at midslope positions (Table 1). Copper, zinc, and manganese concentra- tions were all significantly greater in soils from the greasewood community, which suggests that there may be greater availability of these nutrients to greasewood than to shadscale. However, since no data are available on the nutritional needs of greasewood and shad- scale, no judgement can be made as to the role different concentrations might play in the dis- tribution patterns of species within the two communities. In any event, these minerals accounted for detectable differences between shadscale and greasewood habitats in central Utah. To better understand the microhabitat vari- ation between greasewood and _shadscale, their percent cover was plotted against the measured soil factors (Fig. 3). Variation in each soil factor (percent sand, organic matter, calcium, etc.) was considered as a gradient. Measured values for all sites were ranked and the corresponding species cover values were plotted against the gradient. This procedure allowed visualization of the point along the gradient where each species was most impor- tant as measured by percent cover. With this approach, one is able to visualize those factors that may be important in niche separation. In our case differences existed between grease- wood and shadscale for soluble salts, nitrogen, phosphorus, calcium, zinc, sodium, iron, manganese, and copper. Where a species ex- hibits distinctive patterns with respect to a soil gradient, we may postulate some degree of cause and effect relationship. Conversely, if the species showed no patterns of restriction, but was randomly spread across the gradient, we postulate that the species distribution is probably not affected by that factor. It is possi- ble that strong patterns may exist along a gra- dient and yet not be due to a causal relation- ship. Rather, the pattern could be due to some other environmental factor closely cor- 356 related to the gradient factor in question. In some cases, patterns in species distribution may also relate to patterns in a complex of habitat factors rather than to any single factor. Shadscale showed rather narrow ranges of distribution with respect to the micronutri- ents (zinc, iron, manganese, and copper) but generally grew across the full extent of all the other gradients. Greasewood showed rather broad tolerance ranges across all the gradients (Fig. 3). To further elucidate the relationships asso- ciated with the observed distribution patterns of the four species (burr buttercup, cheat- grass, shadscale, and greasewood), we com- puted species preferences indicies (Skougard and Brotherson 1979) for these species in con- junction with all measured soil factors (Table 6). As shown, the species show different pref- erence indicies for soil factors, texture (sand and fines), soluble salts, phosphorus, calcium, sodium, potassium, iron, manganese, zinc, and copper. Whether these differences are of sufficient magnitude to be causal with respect to the observed differences in the distribution patterns of these species is not clear. How- ever, the differences are detectable and in some cases are of sufficient magnitude that, when coupled with the other evidences dis- cussed in this paper, they aid us in better understanding that differences do exist in the habitats of the two community types. LITERATURE CITED ALLISON, L. E. 1965. Organic Carbon. Pages 1320-1354 in C. A. Black, D. D. Evans, J. L. White, L. E. Ensminger, F. E. Clark, and R. C. Dinauer, eds., Agronomy Series No. 9, Part 2. Methods of soil analysis—chemical and microbiological proper- ties. American Society of Agronomy, Inc. Madison, Wisconsin. ARNOW, L., B. ALBEE, AND A. WYCKOFF. 1980. Flora of the central Wasatch Front, Utah. University of Utah Printing Service, Salt Lake City, Utah. 663 pp. Bouyoucos, G. J. 1951. A recalibration of the hydrometer method for making mechanical analysis of soils. J. Agron. 43: 434-438. BRANSON, F. A., R. F. MILLER, AND I. S. MCQUEEN. 1976. Moisture relationships in 12 northern desert shrub communities near Grand Junction, Colo- rado, USA. Ecology 57: 1104-1124. BROTHERSON, J. D., AND W. E. EVENSON. 1982. Vegetation communities surrounding Utah Lake and its bays. Utah Lake Vegetation Studies report prepared for the Division of Wildlife Resources, State of Utah and the U.S. Bureau of Reclamation, Provo, Utah. 401 pp. GREAT BASIN NATURALIST Vol. 46, No. 2 BUCHANAN, B. A., K. T. HARPER, AND N. C. FRISCHKNECHT. 1978. Allelopathic effects of burr buttercup tissue on the germination and growth of various grasses and forbs in vitro and in soil. Great Basin Nat. 38: 90-96. CALDWELL, M. M., C. B. OSMOND, AND D. L. Nott. 1977. Four carbon pathway photosynthesis at low temperature in cold tolerant Atriplex ssp. Plant Physiol. 60: 157-164. Cstut, B. A. 1979. 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Soil characteristics associated with a primary plant succession on a Mojave Desert, California, USA, dry lake. Ecol- ogy 61: 1013-1018. WALLACE, A., AND E. M. RoMNEY. 1980. The role of pio- neer species in revegetation of disturbed desert areas. Great Basin Nat. Mem. 4: 31-33. WARNER, J. H., AND K. T. HARPER. 1972. Understory char- acteristics related to site quality for aspen in Utah. Brigham Young University Sci. Bull., Biol. Ser. 16(2): 1-20. TREES USED SIMULTANEOUSLY AND SEQUENTIALLY BY BREEDING CAVITY-NESTING BIRDS Kevin J. Gutzwiller’? and Stanley H. Anderson’ ABSTRACT.—We characterize 14 trees used simultaneously and sequentially by breeding cavity-nesting birds in Wyoming cottonwood communities. Our descriptions can be used as a management resource to enrich the diversity of breeding assemblages of these bird species. During our three-season (1982-1984) study along the North Platte and Laramie rivers between Guernsey and Fort Laramie in Platte and Goshen counties, Wyoming, we observed various bird species nesting in decayed sub- strates (trees, limbs, or boles) that were simul- taneously or previously occupied by other breeding hole nesters. These observations highlight behavioral phenomena that increase species richness and density in breeding com- munities of cavity-nesting birds. Habitat patches with individual trees that are used simultaneously or sequentially by several cav- ity-nesting species have higher species rich- ness and density during the breeding season than patches without such substrates (Gutz- willer 1985). The cottonwood (Populus) com- munities we studied (and probably many other plant communities) would support fewer nesting individuals and species in the absence of repeated tree use. In this note, we characterize nesting substrates associated with the phenomena of simultaneous and se- quential nesting. Our purpose is to provide habitat information useful to those who wish to improve species richness and density of cavity-nesting birds in western riparian cot- tonwood communities. Only 14 of 173 (8%) active nest trees re- ceived repeated use during the three breed- ing seasons; 7 trees were used simulta- neously, 4 were used sequentially from year to year, and 3 were used sequentially within a single breeding season (Table 1). We searched for nests and observed behavior around nests for a total of approximately 720 h; yet we detected little intraspecific and interspecific aggression near nest cavities, despite consid- erable overlap in habitat use (Gutzwiller 1985) and the close proximity of many nests. Once we saw a pair of American Kestrels chase an adult male American Kestrel away from their cavity. Twice we observed individual Red- headed Woodpeckers enlarging the entrances of active Downy Woodpecker nests; the latter species (a pair in each case) attacked the for- mer continuously, and neither species nested in the cavities afterward. One other time we saw a pair of Red-headed Woodpeckers de- fending their nest hole by chasing several Eu- ropean Starlings that approached the nest en- trance or the branches of the nest tree. We attribute this general lack of aggression, in part, to an abundance of suitable cavities, not all of which were occupied each year. Most (159 of 173) (92%) of the nest substrates we examined were not used more than once or occupied by more than one species during our three-season study; those that were used more frequently (Table 1) were, presumably, more attractive to various species than other available substrates. This lack of aggression among individuals nesting in close proximity supports other reports of cavity nesters breed- ing harmoniously in similar circumstances (Hoyt 1957, Reller 1972, Jackson 1978). Short (1979: 25) explained that “Despite intense ag- gression between competitors for nesting sites, such competitors at least occasionally appear ‘satisfied’ once they have secured a nesting site [cavity], and there are many re- ports of nesting in proximity of usually aggres- sive nest-hole competitors. . . .” Another rea- son we detected little territorial aggression 'Wyoming Cooperative Fishery and Wildlife Research Unit, University of Wyoming, Laramie, Wyoming 82071. Present address: Department of Statistics, University of Wyoming, Laramie, Wyoming 82071. 398 April 1986 GUTZWILLER, ANDERSON: CAVITY-NESTING BIRDS 3909 TABLE 1. Structural and floristic attributes of 14 nest trees used simultaneously and sequentially by cavity-nesting birds in southeastern Wyoming, 1982-1984. Bird Substrate Tree Nest Diameter Diameter Nest Nest Nest Nest species* use? species® height at breast at nest locations entrance entrance — entrance (m) height height diameter orientation’ _bearing® (cm) (cm) (cm) (°) AMKE, EUST SIM PLCO 9.0, 11.9 133.6 —— limb® —, — A,A 243, 295 AMKE, EUST, SIM PLCO 16.0, 17.8, 935i ——> —— limb OS O AS BSB 2, 16; REWO 15.0 — 7.0 49 AMKE, REWO _ SIM UNKN 7.4, 8.2 49.6 38.1, 25.4 bole 5.0, 6.0 B,A PRIS PD NOFL, EUST SIM PLCO LOW Ol 37.3 25.4, 40.6 bole O05 onl eB YEH 214, 168 REWO, EUST SIM UNKN 15.4, 11.0 (ale? —, — limb 4.8, — H,A 18, 329 HAWO, HOWR_ SIM PEWI 4.5, 5.1 22.5 e8s loe2 bole BO, Gal 18,3} 159, 159 DOWO, NOFL_ SIM PLCO 16.4, 11.4 66.2 12.7, 25.4 limb® 32, 4:7 BB ay COGR, EUST SEQ(C) PLCO 18.3, 20.3 94.8 30.5, 30.5 limb NOP, Bee ANSI} 395, 12 DOWO, DOWO. SEQ(C) UNKN 5.0, 4.4 18.5 —, 15.2 bole 3.8, 3.6 BB 147, 148 DOWO, DOWO. SEQ(C) UNKN 3.7, 3.1 17.5 WAS ee bole 4.0, 3.3. B,B 283, 285 REWO, REWO SEQ(C) UNKN AT, Gil 5G BASO ie 5 =. AA Nie ee EUST, NOFL SEQ(S,H) PLCO 11.5 62.2 20.3 limb 8.3 B 200. DOWO, HOWR SEQ(S,H) PLCO 8.3 110.0 7.6 limb 2.0 B 339 DOWO, HOWR SEQ(S,H) NACO Toll 43.8 15.2 limb 3.0 B 269 “COGR = Common Grackle (Quiscalus quiscula), EUST = European Starling (Sturnus vulgaris), AMKE = American Kestrel (Falco sparverius), REWO = Red-headed Woodpecker (Melanerpes erythrocephalus), NOFL = Northern Flicker (Colaptes auratus cafer), DOWO = Downy Woodpecker (Picoides pubescens), HOWR = House Wren (Troglodytes aedon), HAWO = Hairy Woodpecker (P. villosus). Bird names are from American Ornithologists’ Union (1983). For sequential nesting, order of mnemonics reflects nesting sequence. SIM = simultaneous, SEQ(C) = sequential use during consecutive breeding seasons, SEQ(S,H) = sequential use of the same nest hole within a single breeding season. “PLCO = plains cottonwood (Populus sargentii Dode); UNKN = dead tree, species unknown; PEWI = peachleaf willow (Salix amygdaloides Anderss.), NACO = narrowleaf cottonwood (P. angustifolia James). Plant names are from Dorn (1977). 44 = entrance pointed above horizontal, H = entrance pointed horizontally, B = entrance pointed below horizontal. “Adjusted for 13° easterly declination. f__ — data not available. Enests occurred in different limbs. might be that dominance hierarchies were established through agonistic interaction be- fore we discovered species nesting together. Nevertheless, this probably does not account for the paucity of aggression we noticed over- all, unless dominance hierarchies were estab- lished quickly, and we happened to miss see- ing all of them. Sequential use of cavities and simultaneous and sequential use of nest trees are not rare events (e.g., Brewster 1893, Bent 1948, 1950, Lawrence 1967), presumably because hole nesters (particularly secondary cavity nesters) are versatile with respect to what constitutes a satisfactory nest substrate and because such substrates are typically limited. However, re- peated substrate use in our study area was not common (occurred only 8% of the time), prob- ably because of an abundance of suitable nest trees. Our note describes tree characteristics associated with simultaneous and sequential nesting, thus focusing on features found ac- ceptable, either synchronously or repeatedly, by as many as three species. Such acceptance of these traits increases species richness and evenness (overall diversity) in breeding com- munities of hole-nesting birds (Gutzwiller 1985). Our nest-tree descriptions can, there- fore, be used to attract and maintain greater diversity in breeding assemblages of these species. The Rocky Mountain Forest and Range Ex- periment Station of the U.S. Forest Service funded our study in cooperation with the Wy- oming Cooperative Fishery and Wildlife Re- search Unit. We thank D. M. Finch, R. L. Hutto, M. G. Raphael, and D. E. Runde for constructive criticism on an earlier version of this paper. Our note is based on data collected in partial fulfillment of the senior author's dis- sertation requirements at the University of Wyoming. LITERATURE CITED AMERICAN ORNITHOLOGISTS UNION. 1983. Check-list of North American birds, 6th ed. 877 pp. BENT, A. C. 1948. Life histories of North American nuthatches, wrens, thrashers, and their allies. U.S. Nat. Mus. Bull. 195, Washington, D.C. (Reprinted by Dover Publ., Inc., New York, 1964). 475 pp. _ 1950. Life histories of North American wagtails, shrikes, vireos, and their allies. U.S. Nat. Mus. Bull. 197, Washington, D.C. (Reprinted by Dover Publ., Inc., New York, 1964). 411 pp. 360 BREWSTER, W. 1893. A brood of young flickers (Colaptes auratus ) and how they were fed. Auk 10: 231-236. Dorn, R. D. 1977. Manual of the vascular plants of Wyo- ming. Vols. I and II. Garland Publ., Inc., New York. 1,498 pp. GUTZWILLER, K. J. 1985. Riparian-habitat use by breeding cavity-nesting birds in southeastern Wyoming. Unpublished dissertation, University of Wyo- ming, Laramie. 125 pp. Hoyt, S. F. 1957. The ecology of the Pileated Wood- pecker. Ecology 38: 246-256. GREAT BASIN NATURALIST Vol. 46, No. 2 JACKSON, J. A. 1978. Competition for cavities and Red-cock- aded Woodpecker management. Pages 103- 112inS. A. Temple, ed., Endangered birds, manage- ment techniques for preserving threatened species. University of Wisconsin Press, Madison. 466 pp. LAWRENCE, L. DE K. 1967. A comparative life-history study of four species of woodpecker. Ornithol. Monogr. 5. RELLER, A. W. 1972. Aspects of behavioral ecology of Red- headed and Red-bellied Woodpeckers. Amer. Midl. Nat. 88: 270-290. SHort, L. L. 1979. Burdens of the picid hole-excavating habit. Wilson Bull. 91: 16-28. NEW SPECIES OF MENTZELIA (LOASACEAE) FROM GRAND COUNTY, UTAH Barry A. Prigge’ ABSTRACT. —A new species, Mentzelia (Section Bartonia) shultziorum, of the Loasaceae (Mentzelioideae) is de- scribed. A close relationship with M. multicaulis (Osterh.) Darl. and M. argillosa Darl. is suggested based on the morphology of the flowers, leaves, and seeds. Recent field work in Grand County of southeastern Utah has turned up two popula- tions of an undescribed species of Mentzelia (sect. Bartonia). This section of Mentzelia demonstrates considerable morphological di- versity and adaptability and has radiated into many of the diverse and often isolated habitats resulting from the wide range of substrate, elevation, and precipitation of the Colorado Plateau Province of eastern Utah, western Colorado, northern Arizona, and western New Mexico. Many species of this section occur on unusual substrates that are com- monly unsuitable for most species because of textural properties or high concentrations of evaporites or minerals. Mentzelia is appar- ently able to exploit these habitats by escaping intense competition from species that occur on more suitable substrates. Edaphic factors and isolation are very important in their speci- ation and probably account for the many edaphically restricted, and often locally en- demic populations of Mentzelia in the Colo- rado Plateau Province. It is from such a small, isolated, unusual substrate that this new spe- cies was discovered. Within Mentzelia species recognition is based on subtle and technically difficult char- acters, and in the section Bartonia speciation has created many permutations of the avail- able character states. This has resulted in many taxonomic headaches that standard col- lecting and herbarium techniques have gen- erally been ineffective in solving. Scanning electron microscopy has greatly facilitated the delimitation of taxa by revealing the mi- crostructure of the seed coat (Hill 1976), thus providing two new characters and several character states (radial walls straight or wavy and number and shape of papillae on tangen- tial walls) that help elucidate the taxonomic problems. It is on the basis of micromorpho- logical characteristics of the seed and the stan- dard macromorphological characteristics that we recognize this new species. Mentzelia shultziorum Prigge, sp. nov. Fig. 1 Mentzelia multicaulis (Osterh.) Darl. affine, sed ab eo alis seminum 0.20—0.35 mm latis, foliis caulinis ovatis vel obovatis, dentibus non pro- fundis, diversum. Rounded, much. branched, herbaceous perennial 20-30 cm tall; branches generally arching upward; pubescent with glochidiate and pointed hairs. Cauline leaves obovate, ovate or elliptic in outline, 10-30 (33) mm long, 4-20 mm wide, shallowly toothed at margin with 3-4 teeth on each side, cuneate or broadly attenuate at base, rounded obtuse or acute at apex, densely pubescent on both surfaces with both glochidiate and pointed hairs, some of which have pustulate bases. Bracts linear, lanceolate or oblanceolate; 3.5-11.5 mm long; 1.2-2 mm wide; entire or with | or 2 short teeth along margin. Flowers with pedicels to 2.5 mm long; calyx lobes 5, 5.4-8.5 mm long, deltoid, accuminate, 2.2-3.0 mm wide at base, calyx tube 1-1.5 mm long; petals 5, yellow, 9.2-15.5 mm long, 2.7-5.2 mm wide, acute at apex, clawed, glabrous; the next whorl within the petals of 5 petaloid stamens, broadly obovate to oblance- olate, 6.5-9.0 mm long, 1.1-3.8 mm wide, with functional anthers; stamens numerous; grading from 9 mm long for the outermost ‘Department of Biology, University of California, Los Angeles, California 90024. 361 GREAT BASIN NATURALIST Fig. 1. Mentzelia shultziorum flowering branch: a, Fruit. b, Flower. {2 Vol. 46, No. 2 April 1986 PRIGGE: A NEW MENTZELIA 363 | | | | | ‘ | Fig. 2. Map of Utah and adjacent states showing the distribution of Mentzelia shultziorum (@ ) and some other species that also occur on similar clay substrates: M. multicaulis (©), M. argillosa (@), M. marginata (X), and M. pumila (A). Dashed line represents boundary of the Colorado Plateau Province. whorl to 3.8 mm long for the innermost whorl; _ style 1, 7-9 mm long. Capsules broadly urce- filaments narrow except sometimes for about _ olate, 4.0—5.5 (6.0) mm long and topped by a 3 in the outermost whorl which are up to 1.5 persistent calyx, 4.2-5.0 (5.2) mm diam. mm wide; anthers 0.7-1.0 mm long; pistil 1, | Seeds lenticular, dark brown or black, 2.3-2.8 364 GREAT BASIN NATURALIST Vol. 46, No. 2 10mm AN OO — d — f joes @ eee] Fig. 3. Floral diagram of Mentzelia shultziorum representing elements of the floral whorls: a, Sepal. b, Petal. c, Petaloid stamens. d, Outermost whorl of stamens. e, Stamens from outermost to innermost whorl. f, Style. | Fig. 4. Scanning electron micrograph: a, Whole seed (white bar = 1 mm). b, Seed coat cells showing straight radial | walls and 2-5 papillae on tangential walls (white bar = 0.1 mm). | mm long, 1.4—3.1 mm wide, with a narrow Fisher Valley, 11 air mi NE of Moab; Elev | wing 0.2—-0.35 mm wide, often ridged on one 5,200 ft; T24S, R24E, Sec 19., Shultz & Shultz | or both faces, seed coat cells with straight 2070. Holotype: UTC. Isotypes: RSA. 1 radial walls and with 2—5 papillae on tangen- SPECIMENS EXAMINED:—Utah. Grand Co.: © tial walls. Flowers from July to Sept. above Onion Creek, 5,200 ft., 7.1 mi E of | TyPE:—Utah. Grand Co.: 7.3 miS ofinter- State Hwy 128 on Fisher Ranch Rd (38° 42’ N;_ | section of Hwy 128 and Onion Creek Rd in 109° 15’ W), Prigge, Shultz, & Shultz 6644, — April 1986 (LA); Onion Creek drainage of Fisher Valley (11 air mi NE of Moab) 7.3 mi ESE of Hwy 128: Sec 22, T24S, R24E, 38° 42’N, 109° 15’ W, elev 5,200 ft, Shultz, Shultz, & Prigge 9030 (UTC, LA); 9 mi NE of Moab, NW above pass between Castle Valley and Porcupine Canyon, T25S, R23E, Sec 6 NW 1/4, 4,600 ft, Franklin 2201 (BRY). The specific epithet is in honor of John and Leila Shultz, students of the botany of the Great Basin and Colorado Plateau, who found this new population of Mentzelia. Mentzelia shultziorum is known from only two localities in Grand Co., Utah (Fig. 2). At the first locality it occurs near an old uranium and vanadium mine site on gray clay sub- strates of the Moss Back Member of the Chinle shale, which outcrops locally. It is re- stricted to steep, sparsely vegetated south- facing exposures on these substrates where alluvial and colluvial erosion rates are high. Associated species are: Oryzopsis hy- menoides, Fallugia paradoxa, Atriplex canes- cens, and Chrysothamnus nauseous. At the second locality it is on dark red clay of the Moenkopi Formation(?) with Atriplex, Eri- ogonum, and Ephedra. Mentzelia shultziorum possesses no charac- teristics that are not found in other species of Mentzelia (Sect. Bartonia) in Utah. However, the combination of characteristics is unique, and its relationships within the section Barto- nia are not entirely clear. Seed shape, which is lenticular with a narrow wing 0.2—0.35 mm wide (Fig. 4), is similar to that of M. pumila; the seed coat cells, which have straight radial walls and 2-5 papillae on tangential walls (Fig. PRIGGE: A NEW MENTZELIA 365 4), are like those of M. pumila (Nutt.) T. & G. as well as M. multicaulis (Osterh.) Darl. and M. argillosa Darl.; the floral parts, which are of 5 obovate, glabrous petals and 5 obovate, petaloid stamens (Fig. 3), are like those of M. multicaulis and M. argillosa (to a lesser ex- tent); and the leaves, which are obovate to ovate and shallowly toothed, are similar to M. marginata (Osterh.) Thompson & Prigge or M. pterosperma Eastw. Based on floral (petal shape, vestiture, and size; petaloid stamens presence/absence), seed (size, winged, seed coat cell radial walls straight or wavy and tan- gential wall papillae), and cauline leaf (shape and size) characteristics, Mentzelia shultzio- rum is phenetically close to M. multicaulis and M. argillosa (perhaps closer to the latter) and is probably closely related to these two species. Chromosome counts and hybridiza- tion studies will have to be done to confirm its relationship to these two species. ACKNOWLEDGMENTS I thank John and Leila Shultz for their help in collecting M. shultziorum, John Shultz for the Latin diagnosis, Henry J. Thompson for his helpful suggestions and discussions and for the use of his file of SEM micrographs of Mentzelia seeds, and Kaye Thorne for the illustration of M. shultziorum. LITERATURE CITED HILL, R. J. 1976. Taxonomic and phylogenetic signifi- cance of seed coat microsculpturing in Mentzelia (Loasaceae) in Wyoming and adjacent western states. Brittonia 28: 86-112. UTAH FLORA: JUNCACEAE Sherel Goodrich! ABSTRACT.— A revision of the rush family, Juncaceae is presented for the state of Utah. Included are 28 taxa in two genera. Keys to genera and species are provided, along with detailed descriptions, distributional data, and other comments. No new taxa or combinations are proposed. This paper is another in a series of works leading to a definitive treatment of the flora of Utah. The rush family as represented in Utah is rather small in comparison to some other families, but plants of the family are abundant throughout mesic and wet places of the state. Floral structures are reduced and uniform, and identification often entails observation of such minute features as tailed appendages on seeds. However, the taxa seem quite well marked and mistakes in identification do not seem so common as in some other families with reduced and uniform floral features, such as Salicaceae, Apiaceae, or Chenopodiaceae. Members of the family are more or less comparable in palatibility to grasses and sedges, and they are abundant enough to be of importance to the grazing of domestic live- stock and to big game animals. As in preceding parts of this series, there are two numbers at or near the end of the discussion of each taxon. The first number, in Arabic numerals, indicates the number of specimens from Utah seen in the preparation of this work. The second number, in Roman numerals, indicates the number of specimens collected by the author from the state. ACKNOWLEDGMENTS Appreciation is expressed to the curators of the following herbaria of Utah: Brigham Young University, Provo; Forest Service Herbarium, Ogden; Garrett Herbarium, University of Utah, Salt Lake City; Intermountain Herbarium, Utah State University, Logan. I appreciate the loan of specimens from each of these herbaria. These specimens are the basis of this work. JUNCACEAE Rush Family Perennial or annual grasslike herbs; stems terete or flattened, not jointed, caespitose or arising singly or few together from rhizomes; leaves sheathing, alternate or all basal, mostly 2—ranked, blades linear, sometimes much re- duced or lacking; inflorescence headlike to open paniculate, subtended by an involucral bract; branches, heads, and pedicels often subtended by bractlets; flowers bisexual (ours), sometimes subtended by bracteoles borne at pedicel apices, directly below perianth; perianth much re- duced, petals and sepals hardly if at all different and referred to herein as tepals, tepals membra- nous, rather scalelike, greenish or brownish, 6, an outer and inner set with 3 each; stamens (3) 6; pistal 1; ovary superior, with 1 or 3 chambers; fruit a capsule with 1 or 3 chambers. 11(1). Seeds numerous in each capsule; leaves glabrous, sheaths open; bracteoles subtending flowers entire onlacking: 254... .4eeeeeeee Juncus — Seeds 3 per capsule; leaf blades pubescent at least on margins near base except sometimes in L. parviflora, sheaths closed; bracteoles sub- tending flowers entire to lacerate ....... Luzula Juncus L. RusH Juncus Perennial or annual grasslike herbs; stems terete or flattened; leaf blades flat, strongly folded, or terete, when terete sometimes hol- low with cross membranes at intervals (sep- tate) or reduced to a bristle or lacking; flowers as described in family; seeds numerous, minute, usually apiculate or tailed. ‘Intermountain Research Station, Forest Service, U. $. Department of Agriculture, Ogden, Utah 84401. Present address: Vernal Ranger District, Ashley National Forest, Vernal, Utah 84078. 366 April 1986 Rlantsannuial ieee ree 2 Blantsiperennialgesperer sere erie 3 Plants 0.5-2 cm tall, subscapose; scapose stem with 1 flower; stamens 3; leaves not over Oloremilon Gurwen ein eet J. bryoides Plants 2-30 cm tall, not scapose; inflores- cence with 1—20 flowers; stamens 6; leaves ODO GINO Scocsc00s00n0b0 ox J. bufonius Flowers (1) 2-5 in solitary terminal head; leaves basal or nearly so, hollow, septate; plants densely tufted, without rhizomes, 3-19 cm tall, infrequent in Uinta Mountains Chou d EEe ens ncea Ee hae ar R J. triglumis Flowers either more numerous or not in a soli- tary terminal head or plants otherwise different from above Stems with 0-2 (rarely more) leaf blades; blades borne on lower 1/5 of plant, not hollow, not sep- tate, sheaths sometimes ending in a rudimentary bristle instead of a leaf blade; flowers not in heads but borne singly, each subtended by 2 hyaline bracteoles; rhizomes lacking or short and plants mostly caespitose except in J. arcticus ......... 5 Stems with 2 or more well-developed leafblades, at least uppermost blade borne above lower 1/3 of plant or else hollow and septate; flowers borne in 1 or more heads and not individually sub- tended by bracteoles; plants mostly with rhi- zomes (note: J. compressus and J. gerardii with leafy stems and solitary, bracteolate flowers are keyedjbothwways)s7nem aero a eee ee 12 Leaves all reduced to bladeless sheaths, upper ones sometimes with a bristle-tip, this not over 5 mm long; inflorescence with 5-75 or more flow- ers; seed not tailed At least uppermost leaf of most stems with a well-developed leaf blade well over 10 mm long, or, if leaves all reduced to bladeless sheaths (J. drummondii), inflorescence with only 1-3 flow- ers and seeds tailed ...................0005. fa Involucral bract about as long or longer than stem and inflorescence appearing at or below midlength of plant; stems seldom over 1 mm thick, somewhat tufted; plants of Uinta Moun- tains above 2,950 m, ratherrare .. J. filiformis Involucral bract mostly shorter than stem and inflorescence appearing above midlength of plant; stems often over 1 mm thick, mostly aris- ing singly or few together from robust dark rhi- zomes; plants widespread ........... J. arcticus Seeds tailed at each end, tails 1/2 as long to longer than body; inflorescence with 1-6 flow- ers; stem-leaves with blades lacking or reduced to a bristle or uppermost 1(2) with a well-devel- oped blade; plants mostly found above 2,620 m. Seeds apiculate but not tailed; inflorescence with (1)6-50 or more flowers; some of lower stem-leaves commonly with well-developed blades; plants commonly found below and above PALOPA iat lace one eee ee tetad neo ae oie iets aes eet ee eee 10 GOODRICH: UTAH FLORA, JUNCACEAE 10(8). 11(10). 12(4). 13(12). 14(12). 367 Stems with bladeless leaves, uppermost and often lower sheaths tipped with a bristle, this not over 1 cm long; tails of seeds equal to or longer than body; tepals 5-8 mm long; cap- sules blunt and more or less retuse, equal to or alittle shorter than tepals ... J. drummondii Most of stems with a well-developed leaf- blade on at least uppermost sheath, lower sheaths often tipped with a bristle; tails of seeds equal to or shorter than body; tepals andicapsuleswvaniouss seer eter 9 Capsules ovoid, retuse at apex; tepals 4—5 (5.5) mm long; anthers less than 1 mm long; filaments longer than anthers ........ J. hallii Capsules oblong, pointed; tepals 5-8 mm long; anthers 1.5-2 mm long; filaments only aboutaOkoimmylong ean ee ee J. parryi Tepals, at least outer ones, with incurved or hooded tips, rather obtuse, 1.5—2.5 mm long; uppermost leaf often borne above midlength Of SC rater SN ences ea he werner ences 13 Tepals with acute to acuminate erect tips, 3-5 mm long; uppermost leaf borne on lower LiStofistemens eee ene ae 11 Capsules retuse at apex, completely 3-loculed; panicles mostly less than 2 cm long; tepals with hyaline margins extending to apex of acute tip; plants montane, mostly VMAX COIN gavesoccvs odode nes J. confusus Capsules blunt but not retuse, incompletely 3-loculed; panicles various but often over 2 cm long; at least outer tepals with hyaline margins not extending to acuminate or acumi- nate-attennuate tip; plants found mostly be- lowi25380nmigers tee Dib me ehoe Cees J. tenuis Flowers borne singly, each subtended by 2 bracteoles; pedicels sometimes also sub- tended by bractlets; tepals with incurved or hooded tips, 1.5-2.5mm long............ 13 Flowers borne in 1-many heads, not sub- tended immediately by bracteoles; pedicels usually subtended by bractlets; tepals, at least outer ones, with erect or spreading tips, as short or longer than above ............. 14 Anthers about 3 times longer than filaments; capsule ellipsoid-ovoid, equal or slightly ex- ceeding tepals; plants sometimes over 40 cm tall, known in Utah from a single collection from a hot spring in Salt Lake County...... Anthers scarsely longer than filaments; cap- sule globose-ovoid, distinctly exserted; plants 20-40 cm tall, known from flood plains of Green and Colorado rivers...... J. compressus Leaf blades flat or strongly folded and appear- ing flat at least toward base, not terete, not hollow; sheaths with hyaline margins; cap- sules not exserted beyond tepals .......... 15 Leaf blades terete and hollow, if only toward tip, then sheaths without hyaline margins and capsules conspicuously exerted beyond (OIG covscguscsaanocaccoopuscamonce: 17 368 15(14). 16(15). 17(14). 18(17). 19(18). 20(19). GREAT BASIN NATURALIST Leaves strongly folded, narrow edge oriented toward the flattened stem; scarious margins of sheaths extending well beyond juncture with stem, gradually tapering to inconspicuous au- ricles or auricles lacking; margins of blade more or less united beyond scarious margins ROE Ae & SOI SS OER Oe J. ensifolius Leaf blades flat, flat surface oriented toward terete stem; scarious margins of sheaths not extending beyond juncture with stem ..... 16 Seeds tailed, tails as long or longer than body; tepals granular-papillate on back; heads sometimes with more than 10 flowers; plants known from Duchesne, Wasatch, and Salt Makecountiestirees ene sre J. regellii Seeds apiculate but not tailed; tepals smooth on back; heads with 3-10 flowers; plants WIGeSpreaduwcn sreeane tenn earn es J. longistylis Leaf blades folded to enrolled toward base, becoming terete and hollow distally; sheaths without hyaline margins; auricles lacking; capsules conspicuously exceeding tepals; seeds long-tailed; stamens 6; plants rare, from above timberline on Uinta Mountain J. castaneus Leaf blades terete and hollow from collar and outward, septate; scarious margins of sheaths projected into auricles; capsules not much if any longer than tepals and seeds not tailed or else stamens 3; plants widespread......... 18 Seeds tailed; stamens 3; capsules conspicu- ously exceeding tepals; plants known from one collection in Box Elder County. . . J. tweedii Seeds not tailed; stamens 6; capsules various; Plants widespread arr et eet tan eee 19 Capsules tapered almost from base into a mostly nondehiscent conspicuous _ stylar beak, often divergent in all directions in ma- ture globose or hemispheric head; heads rarely solitary, greenish or light brown; tepals acuminate or acuminate-subulate; rhizomes sometimes swollen and tuberous at nodes; plants found mostly below 2,320m........ 20 Capsules rather abruptly narrowed above into a dehiscent short or inconspicuous stylar bank, ascending to slightly spreading in heads, or, if spreading in all directions, then heads solitary; heads light or deep brown to blackish-purple; rhizomes not as above; plants from above and below 2,320m...... 21 Auricles 1.5—5 mm long; tepals 4—5 mm long, with rigid long-acuminate or subulate tips; mature heads 10-15 mm wide; capsules shorter or scarsely longer than tepals; stems toj6immithickean ene eee aoncr J. torreyi Auricles 0.25—1 mm long; tepals 2.5—3.5 mm long, acuminate, tips not so rigid as above; mature heads 5—12 mm wide; capsules to ca | mm longer than tepals; stems 1—2 mm thick .... J. ensifolius var. montanus Var. brunnescens (Rydb.) Crongq. [J. brun- nescens Rydb.; J. saximontanus f. brunnes- cens (Rydb) F.J. Hermann, J. tracyi f. utahen- sis (R. F. Martin) F. J. Hermann; J. utahensis R. F. Martin]. Along rivers, streams, ditch banks, around seeps, springs, ponds, lakes, and in hanging gardens, marshes, meadows, and bogs at 1,065 to 2,450 (2,740) m in Beaver, Duchesne, Emery, Garfield, Grand, Iron, Kane, Rich, San Juan, Sanpete, Sevier, Summit, Uintah, Utah, Washington, and Wayne counties, and intergrading into var. montanus in nearly all counties of state; nearly throughout range of var. montanus, but more common southward especially in Arizona and the only phase in Texas; 145 (v). See discus- sion under var. montanus. Var. ensifolius Wet places in mountains in Daggett, Salt Lake, Tooele, and Uintah coun- ties: Alaska to northern Mexico and east to Alberta and Arizona, 4 (i). The few specimens 372 from isolated stations in Utah with only 3 sta- mens per flower could be nothing more than odd specimens of var. montanus. However, to north of our area this is acommon phase. Var. montanus (Engelm.) C. L. Hitche. [J. xiphioides var. montanus Engelm.; J. saxi- montanus A. Nels.; J. tracyi Rydb. |. In mead- ows, along streams and rivers, about seeps and springs and other wet places at (853) 1,830 to 3,100 m in all counties of state; Alaska to southern California and east to Saskatchewan and New Mexico; 128 (v). In the study of numerous specimens seen from the state, the following trends were noted: (1) Most plants of the lower elevations in the Canyonlands Sec- tion are rather easily assigned to var. brunnes- cens, and they mostly have apiculate-tailed seeds.(2) Plants of the Great Basin are often referrable to var. montanus, and they have apiculate but rarely tailed seeds, (3) Through- out the plateaus and mountains that run through the center of the state and in the Uinta Mountains, there are plants of both va- rieties and numerous intermediate plants, and seeds are commonly with or without tails in both varieties as well as in intermediate plants. Color phases of the inflorescence (pale green to purplish black) are also found in both varieties and in intermediate plants. Perhaps outside Utah the picture is somewhat clearer, but Utah seems to be near the center of where the two varieties overlap. More than 25% of the Utah specimens examined appeared to be intermediate. Juncus filiformis L. Plants perennial, 5-40 cm tall; stems arising singly or in tufts from creeping rhizomes, terete, rarely over | mm in diameter; leaves reduced to bladeless sheaths, uppermost one often tipped with a tiny bristle, confined to lower 1/5 of plant; involucral bract 10-27 cm long, appearing as a continuation of stem, as long or to over 4 times longer than stem; inflorescence appearing lat- eral and on lower 1/2 to 1/10 of plant, 0.5—1(2) cm long, compact, with ca 5—20 flowers, flow- ers borne singly; bractlets subtending branches and pedicels scarious, lower ones sometimes aristate; pedicels nearly obsolete or to 4 mm long; bracteoles scarious, ovate or oblong; tepals 3-4.5 mm long, greenish, lanceolate, acute to acuminate, subequal or outer ones slightly longer; stamens 6, fila- ments ca 0.6 mm, anthers 0.4—0.6 mm long; GREAT BASIN NATURALIST Vol. 46, No. 2 styles and stigmas less than 1 mm long; cap- sules 3-loculed, ca 2-3 mm long, greenish, ovoid to obovoid, abruptly tapered to a short stylar beak; seeds 0.4—0.6 mm long, minutely winged-apiculate at both ends. Wet subalpine meadows and along streams in Uinta Moun- tains at 2,990 to 3,200 m in Summit, Uintah, and Wasatch counties; Alaska to Labrador and south to Utah and West Virginia; 10 (iv). Juncus gerardii Lois. Black Grass; Mud Rush. Perennial plants 15—80 cm tall; stems somewhat tufted on slender dark rhizomes; leaves rather scattered on stems, upper ones usually borne on upper 1/2 of stem, lower ones bladeless or with reduced blades, upper blades flat, 1.5-3 mm wide; inflorescence with several to many flowers; flowers borne singly and subtended by scarious bracteoles, nearly sessile to long-pediceled; tepals 2—-3.5 mm long, dark brown with a greenish midstripe, blunt and usually hooded at tip; stamens 6, anthers ca 1.5 mm long, ca 2-3 times longer than filaments; capsules ovoid to obovoid, rounded, about equaling but a little shorter than tepals; seeds 0.5—0.6 mm long, slightly apiculate at tapered end, nearly trun- cate-apiculate at other end. Becks Hot Spring in Salt Lake County (Flowers sn 24 Sept. 1924 UT). The population is not likely existing be- cause the area is now part of Interstate 15; Eurasia, Atlantic and Pacific coasts in North America, and sporadic inland; 1(0). Juncus hallii Engelm. Halls Rush. Peren- nial caespitose plants 20—40 cm tall, from fi- brous roots, rhizomes lacking; stems terete, to ca 1.5 mm thick; leaves basal and on lower 1/5 of plant, usually only uppermost cauline leaf bearing a well-developed blade, lower stem- leaves bladeless or tipped with a short bristle, innovations sometimes with well-developed blades; blades terete, upper side more or less channeled toward base, not or inconspicu- ously channeled toward tip, less than 1 mm wide; involucral bract 0.7—2.5 (7.5) cm long, scarious and caudate to awned, or elongate and leaflike, with scarious margins projected into auricles; inflorescence to 1.7 cm long, with (2) 3-6 flowers; flowers rather con- gested, but borne singly; bractlets subtending pedicels scarious, attenuate to caudate; pedicels 1-8 mm long; bracteoles subtending flowers hyaline, ovate to nearly orbicular; tepals 4—5(5.5) mm long, subequal or inner ———————————————————————— April 1986 ones a little shorter, lanceolate, acute, usually with greenish centers flanked by purple and with hyaline margins; stamens 6, filaments 1-1.5 mm long, anthers 0.5-0.7 mm long; styles and stigmas not over 1 mm long; cap- sules 3—loculed, equaling or ca 1 mm longer than tepals, triquetrous, retuse at apex, dark brown to purplish black; body of seeds 0.6—0.7 mm long, tailed at each end, tails ca 1/2 as long as body. Dry, wet, and boggy meadows, margins of ponds and lakes, and along streams at 2,956 to 3,350 m in Beaver, Daggett, Duchesne, Garfield, Summit, Uin- tah, and Wasatch counties; Montana to Colo- rado and Utah; 13 (xi). Juncus longistylis Torr. in Emory Longstyle Rush. Perennial plants 20-63 cm tall; stems arising singly or few together from creeping rhizomes, terete; leaves somewhat crowded on lower 1/2 of stem, but uppermost one often on upper 1/2—3/4 of stem, scarious margins of sheaths prolonged into auricles to ca 1 mm long, blades flat, not hollow, not septate, 1-3 mm wide; involucral bract 1—4 cm long, about equaling or shorter than inflo- rescence, mostly scarious, rarely leaflike, nar- rowly attenuate to caudate; inflorescence 1-7.5 cm long, usually with (1) 3-13 heads, heads with 3-10 flowers; bractlets subtending heads and pedicels scarious, acute to caudate; pedicels to ca 2 mm long, concealed in scari- ous bractlets; tepals (4)5—6 mm long, acute to acuminate, often purplish with greenish cen- ter and broad whitish or silvery hyaline mar- gins; stamens 6, filaments 0.5—1 mm long, anthers (1) 1.3-2 mm long; styles 1-2 mm long; stigmas ca 2 mm _ long; capsules 3—-loculed, shorter than or rarely equaling tepals, rather abruptly tapered to or retuse at stylar beak, brownish or purplish black; seeds ca 0.5 mm long, apiculate at each end. Wet meadows, along streams and rivers, about seeps and springs and other wet places, occa- sionally in alkaline places at 1,380 to 3,350 m in Box Elder, Cache, Daggett, Duchesne, Emery, Garfield, Grand, Kane, Millard, Morgan, Salt Lake, San Juan, Sanpete, Sevier, Summit, Tooele, Uintah, Utah, Wasatch, and Wayne counties; southern Canada, Washington to South Dakota and south to California and New Mexico; 100 (viii). Juncus mertensianus Bong. Mertens Rush. Perennial plants 13—42 cm tall; stems arising GOODRICH: UTAH FLORA, JUNCACEAE 373 singly or loosely to tightly clustered on long creeping or short rhizomes, rhizomes some- times short and stout and plants caespitose with numerous fibrous roots; leaves basal and on upper 1/4—3/4 or higher on stems, scarious margins of sheaths projected into ligulelike auricles 0.5=—2 mm long; blades terete, chan- neled above, hollow, septate, 0.5—2 mm wide when pressed; involucral bract 0.8—3.2 cm long, rarely leaflike, often caudate; inflores- cence 0.5—1.5 (3.5) em long, with 1 (2) head(s); heads with 5—40 or more flowers, to ca 1.5 cm thick; bractlets subtending heads and pedicels scarious, acute to caudate; pedicels toca | mm long; bracteoles lacking; tepals 2.5-4 mm long, acute to acuminate, blackish purple; sta- mens 6, filaments 1-1.3 mm long, anthers 0.5—1 mm long, shorter than filaments; styles to 1 mm long; stigmas to 1 mm long; capsules 1—chambered, triquetrous, slightly to con- spicuously shorter than tepals, abruptly ta- pered to or slightly retuse at stylar beak, often as blackish purple as tepals; seeds 0.5—0.7 mm long, apiculate but hardly tailed. Wet meadows, along streams, about seeps and springs, margins of lakes and ponds, and En- gelmann spruce-lodgepole pine, and tundra communities at 2,435 to 3,415 m in Box Elder, Cache, Duchesne, Garfield, Iron, Juab, Piute, Salt Lake, San Juan, Sanpete, Sevier, Summit, Uintah, Utah, and Wasatch coun- ties; Alaska and Yukon south to southern Cali- fornia and South Dakota; 83 (vi). Juncus nevadensis Wats. Nevada Rush. [J. badius Suksd.]. Perennial plants 12-35 (53) cm tall; stems more or less terete, arising singly or a few together from creeping rhi- zomes; leaves basal and on upper 1/4—3/4 or higher on stems, with well-developed blades, scarious margins of sheaths projected into ligulelike auricles 1.5-3 mm long, blades terete, hollow, septate, somewhat channeled above, 0.5-2 mm wide; involucral bract 1-3(8.5) cm long, bractlike to leaflike, seldom much exceeding inflorescence; inflorescence 1-8 cm long, with (1) 2-13 heads, heads with (3)6-13 or more flowers; bractlets subtend- ing, heads membranous, attenuate to cau- date; pedicels to ca 1 mm long; bracteoles lacking; tepals (3) 3.5-5 mm long, brown to purplish black; anthers 1-2 mm long, longer than filaments; styles to 3 mm long, stigmas 1-2 mm long; capsules 1—loculed, equal to or 374 conspicuously shorter than tepals, _ tri- quetrous, rounded or rarely slightly retuse at apex; seeds 0.5—0.6 mm long, apiculate but not tailed. Dry meadow, wet meadow, silver sagebrush-meadow, and lodgepole pine com- munities at 2,286 to 3,050 m in Box Elder, Cache, Daggett, Duchesne, Emery, Gar- field, Grand, Kane, Rich, Sanpete, Summit, Uintah, Wasatch, and Washington counties; southern British Columbia and Alberta south to California and New Mexico; 43 (xvii). The inflorescence is sometimes similar to those of J. ensifolius and J. longistylis, and plants of these taxa are sometimes confused. The leaves are different in each of these. Occa- sional plants with only 1 or 2 heads are rather easily mistaken for those of J. mertensianus . Juncus nodosus L. Jointed Rush. Perennial plants 17-58 cm tall; stems terete, 1-2 mm thick, arising singly to densely clustered on creeping rhizomes, rhizomes sometimes with small tuberlike segments; leaves usually ex- tending well up stems, those of stem with well-developed blades, those of innovations often without blades, scarious margins of sheaths prolonged into short auricles 0.25-1.0 mm long, blades terete or chan- neled above, hollow, septate, 0.5-1.5 mm wide when pressed; involucral bract 2.5-12 cm long, more or less leaflike; inflorescence 1.5—7 cm long, congested or rather open, with 3-12 globose or nearly globose heads; heads sessile or pedunculate, with (5) 10-25 flowers, 5-12 mm wide, flowers widely spreading to divergent; bractlets subtending heads scarious, acute to cuspidate; pedicels to ca 1 mm long; tepals 2.5-3.5 mm long, sub- equal, acuminate, acuminate tips shorter than and not so rigid as in those of J. torreyi; sta- mens 6, filaments ca 0.8 mm long, anthers 0.6—0.8 mm long; styles to about 3 mm long, stigmas ca 1 mm long; capsules incompletely 3—loculed, to ca 1 mm longer than tepals, slender, gradually prolonged into a tardily de- hiscent stylar beak, sharply triangular in cross section; seeds 0.4—0.5 mm long, apiculate. Wetlands along streams and rivers and in wet and boggy meadows at (1,250) 1,640 to 2,320 min Cache, Duchesne, Garfield, Piute, Rich, Summit, Uintah, Washington, and Wayne counties; southern Canada and Northern United States, south in West to California and Texas; 36 (xiii). GREAT BASIN NATURALIST Vol. 46, No. 2 Juncus parryi Engelm. Parrys Rush. Perennial caespitose plants (10)15—30 cm tall, from fibrous roots, lacking rhizomes; stems terete, about 1 mm thick; leaves basal and borne on lower 1/5 of stems, usually only up- permost one with a well-developed blade, lower sheaths usually tipped with a bristle or much reduced blade, and uppermost one sometimes reduced on a few of the stems, scarious margins of sheaths barely prolonged into auricles less than 0.5 mm long; blades less than 1 mm thick, terete, channeled to strongly involute below, obscurely channeled above, not septate; involucral bract 1.5—6(9) cm long, leaflike, terete, more or less simulat- ing a continuation of stem, often auriculate; inflorescence 0.7—2.2 cm long, with 1—4 flow- ers, flowers borne singly; bractlets subtend- ing pedicels scarious, acute to caudate, or one of them often similar to involucral bract but smaller; pedicels 1-20 mm long, abaxial ones often much longer than adaxial; bracteoles subtending tepals ovate to lance-ovate, rounded to acute or acuminate-attenuate; tepals 5-8 mm long, inner ones to | mm shorter than outer and somewhat less pointed and more scarious; stamens 6, filaments ca 0.3 mm long, anthers 1.5—2 mm long; styles ca 1.5mm long, stigmas to 3.5 mm long; capsules a little shorter to a little longer than tepals; body of seeds 0.6-0.7 mm long, with tails a little shorter than or to 0.1 mm longer than body. Engelmann spruce, lodgepole pine, meadow, and alpine communities, on wet to dry rocky ground, sometimes in rocky snowflush areas at 2,620 to 3,420 m in Cache, Duchesne, Iron, Juab, Salt Lake, Summit, Uintah, and Wasatch counties; British Co- lumbia to Alberta and south to California and Utah; 50 (v). Juncus regelii Buch. Regels Rush. [J. jones- ii Rydb.]. Perennial plants 10-60 cm tall; stems arising singly or few together from creeping rhizomes; leaves basal and extend- ing well up on stems, scarious margins of sheaths prolonged into inconspicuous or short auricles; blades flat, 2-4 mm wide, neither hollow nor septate; involucral bract 1-4 cm long; inflorescence with 1-5 globose or hemi- | spherical heads, heads 8-20 mm across; bractlets subtending heads scarious; bracte- oles lacking; pedicels ca 1-2 mm long; tepals 4—6 mm long, papillose-roughened on back, | April 1986 with a greenish midstripe flanked by dark brown and with scarious margins, inner ones slightly shorter and slightly less pointed than outer ones; stamens 6, anthers 1—1.5 mm long, subequal to filaments; capsules 3—loculed subequal to tepals, oblong-ovoid, truncate to retuse; body of seeds about 0.5 mm long, tailed at each end, tails about as long or longer than body. Meadows and along streams at 2,750 to 3,060 m in Duchesne, Salt Lake, and Wasatch counties; southern Wash- ington to northern California and east to Mon- tana and Utah; 8 (i). Much like J. longistylis but distinct in tailed seeds and more or less marked by papillose-roughened tepals. Juncus tenuis Willd. Poverty Rush. [J. dud- ley Wiegand; J. interior Wiegand; J. tenuis var. dudleyi (Wiegand) F. J. Hermann; J. tenuis var. congestus Engelm.]| Perennial, caespitose plants 22—65 cm tall, with fibrous roots; rhizomes lacking; stems terete, to 1.8 mm wide; leaves basal and cauline, borne on lower 1/5 of plant, those of stem mostly with well-developed blades, some of the basal ones with blades reduced to bristles, scarious mar- gins of sheaths projected into auricles to ca 0.75 mm long; blades flat but soon moderately to strongly involute, not hollow, not septate, to ca 2 mm wide; involucral bract 2—18 cm long, leaflike; inflorescence (0.7)1.5—8.5 cm long, congested to rather open, with (4) 10—50 or more flowers, flowers borne singly; bractlets subtending branches and pedicels scarious, caudate-acuminate or awned, or lower ones leaflike and similar to involucral bract; pedicels obsolete or to 5 mm long; bracteoles subtending tepals scarious, ovate to lanceolate, acute to caudate; tepals 4-5 mm long, subequal or outer ones a little longer than inner, outer ones narrowly acuminate or acuminate-attenuate with hyaline margins mostly not extending on to acuminate tip, inner ones mostly acute to slightly acuminate with hyaline margins often extending to tip; stamens 6, filaments 0.6—1 mm long, anthers 0.5—0.8 mm long; styles and stigmas ca 1.5 mm long; capsules imperfectly 3—loculed, 1-2 mm shorter than tepals, obtuse to trun- cate; body of seeds 0.3—0.4 mm long, with apiculate ends to about 0.1 mm long. Along streams, washes, ditchbanks, rivers, margins of ponds and reservoirs, about seeps and springs, and in meadows and hanging gardens GoopRICH: UTAH FLORA, JUNCACEAE 375 at 1,135 to 2,380 m in Cache, Daggett, Duch- esne, Garfield, Grand, Millard, Rich, San Juan, Uintah, Utah, Wasatch, Washington, and Wayne counties; widespread in North America and introduced in temperate regions elsewhere in world; 41 (iv). Three intergradi- ent phases-can be seen in our plants (var. dudleyi with cartilaginous, often yellow to brown auricles about 0.5 mm long; vars. con- gestus and tenuis with membranous, usually greenish or whitish auricles often over 0.5 mm long, the former with congested panicles mostly less than 3 cm long, the latter with open panicles mostly over 3 cm long). The morphological differences are minor at best, and the taxa are more or less sympatric. Vari- ety tenuis does seem to be more common in the southern half of the state. (Perhaps the recognition of these varieties serves more to waste time than for any important purpose. ) Juncus torreyi Cov. Torreys Rush. Peren- nial plants (10) 20-80 (100) cm tall; stems terete, to 6 mm in diameter near base, arising singly or a few together from robust creeping rhizomes, rhizomes often with swollen tuber- like segments; leaves well distributed up stem, scarious margins of sheaths prolonged into auricles, (1.5) 2-5 mm long; blades terete sometimes channeled on upper side, hollow, septate, 1-3 mm_ thick; involucral bract 1.5-17cm long, more or less leaflike; inflores- cence 1.5—7 cm long, congested, with (1) 3-13 more or less globose and sometimes bur- like heads, heads 6—15 mm across, with 10—50 or more flowers, flowers widely spreading, and some usually reflexed; pedicels short and hidden in compact heads; bracteoles lacking; tepals 4—5 mm long, or inner ones slightly shorter, long-acuminate and rigid at tip; sta- mens 6, filaments 0.7—1 mm long, anthers 0.5-0.8 mm long; styles ca 0.25 mm long, stigmas ca 1 mm long; capsules incompletely 3-loculed, slender, triquetrous, equal to or slightly longer than tepals, slender stylar beak tardily dehiscent; seeds 0.4—0.5 mm long, apiculate but not tailed. Along streams, riv- ers, washes, and ditchbanks, at margins of ponds and lakes, about seeps and springs, and in saline or alkaline moist to wet meadows, marshes, and swamps at 850 to 2,010 m in all Utah counties except Iron, Millard, Piute, Summit, and Wasatch; southern Canada to northern Mexico; 112 (iv). 376 Juncus triglumis L. Three flowered Rush. [J. albescens Fern. |. Perennial plants 4-19 cm tall, densely caespitose; leaves basal or on lower 1/4 of stems, those of stem with well-developed blades, scarious margins of sheaths projected into auricles to about 0.75 mm long; blades about 0.5 mm wide, hollow, sepatate; involucral bract 5-10 mm long, often purplish; inflorescence a solitary head, this 5-8 mm long, with (1)2-5 flowers; bractlets subtending pedicels similar to but somewhat smaller than involucral bract; pedicels to ca 1 mm long; tepals 3-4 mm long, acute, cream, yellowish, or greenish yellow, and often marked with purple; stamens 6, as long as tepals or ca 1 mm shorter, filaments to 2 mm long, anthers 0.5-0.8 mm long; stigmas and styles about 1 mm long; capsules shorter or about 1 mm longer than tepals, abruptly tapered to blunt or subtruncate at tip, blackish purple; trigonous-cylindric body of seeds ca 0.75-1 mm long, tailed at both ends, each tail ca 1/2 to as long as body, more or less flattened, scarious. Wet meadows and bogs at 2,800 to 3,810 m in Uinta Mountains in Daggett, Duchesne, Sum- mit, and Uintah counties; circumboreal, south in Rocky Mountains to New Mexico; 10 (v). Utah plants are referable to var. albescens Lange. Juncus tweedyi Rydb. [J. canadensis var. kuntzei Buchenau]. Perennial plants 20-40 cm tall, stems clustered, terete; rhizomes appar- ently lacking; leaves basal and cauline, scarious margins of sheath projected into auricles (0.5)1-2 mm long; blades 1—2.5 mm thick, terete or nearly so, hollow, septate; involucral bract shorter than or somewhat longer than inflores- cence; inflorescence with 2—8 heads, these with 3-8 flowers, 3-8 mm wide, brown; bractlets subtending heads and pedicels scarious; bracte- oles lacking; tepals 3-4 mm long gradually acute, inner ones equal to or a little longer than outer; stamens 3, anthers 0.5-0.7 mm long, shorter than filaments; capsule slightly longer than tepals, triquetrous, more or less acute, im- perfectly 3—loculed; seeds cylindrical with tails ca 1 mm long at each end. Near Corinne in Box Elder County (Kuntze 3133, NY). Wet places about hot springs, Yellowstone National Park, Wyoming; 0 (0). LuzULA DC. WOODRUSH Perennial grasslike herbs generally with long spreading hairs along margins of leaf GREAT BASIN NATURALIST Vol. 46, No. 2 blades, at least when young; leaves sheathing, sheaths closed, blades flat; inflorescence headlike or spicate to open paniculate; flowers subtended by bracteoles; tepals 6; stamens 6; capsules 1—loculed, with 3 seeds, dehiscent along midribs of carples; seeds with or without caruncles, sometimes comose with extremely fine hairs. 1. Inflorescence an open panicle, panicle sometimes drooping; leaves glabrous or nearly so at maturity, blades 3-11 mm wide; plants 27-77 cm tall ..... L. parviflora — Inflorescence of few to several congested or re- mote spikes or spikelike racemes, sometimes headlike; margins of leaves pubescent with long hairs especially near collar, 1-6 mm wide; plants 5-42. cm tall. oe eee 2 2. Flowers borne in a terminal compound spikelike or headlike inflorescence; leaves 1-3 mm wide; seeds without or with an inconspicuous caruncle; plants widespread = 4... a-ha aaa L. spicata — Inflorescence with 1 or more lateral spikes, some of laternal ones often borne on peduncles to 3(5.5) cm long; leaves 2-6 mm wide; seeds with a con- spicuous caruncle; plants known from Uinta Mountains) = ss) on teen L. campestris Luzula campestris (L.) DC in Lam. & DC. Hairy Woodrush. [Juncus campestris L.; L. multiflora (Retz.) Lej.; L. intermedia (Thuill.) A. Nels.] Plants 13-42 cm tall; stems more or less tufted; leaves basal and cauline, blades flat, 2-6 mm wide, margins with scattered to moderately dense villose hairs ca 2-7 mm long or longer; involucral bract 3-9 mm long, leaflike; some bractlets subtending branches and peduncles leaflike, others scarious; inflo- rescence 1.5—5 cm long, usually conspicu- ously branched, with 3-12 spikes; spikes 5-12 mm long, sessile or on peduncles to 3 (5.5) cm long, with 5-15 or more flowers; bracteoles subtending flowers hyaline, entire or ciliate to fringed; tepals 2-3.5 mm long, greenish to brownish, acute or acuminate to scarcely caudate; anthers ca 0.5—1 mm long, filaments about equal anthers; capsules equal to or shorter than tepals; seeds 1.4—1.7 mm long, with a whitish caruncle ca 0.3—0.6 mm long. Lodgepole pine, Engelmann spruce, and meadow communities at 2,440 to 3,110 m in Uinta Mountains in Daggett, Duchesne, Summit, Uintah, and Wasatch counties; widespread in temperate regions of world; 10 (vii). Utah plants apparently belong to var. multiflora (Ehrh.) Celak. April 1986 Luzula parviflora (Ehrh.) Desv. Millet Woodrush. [Juncus parviflorus Ehrh.; L. wahlenbergii Rupr. misapplied] Plants 27-77 cm tall; stems solitary or few together from short rhizomes and fibrous roots; leaves basal and cauline, glabrous or with a few scattered long-villose hairs especially near collar, blades flat, 2-11 mm wide; involucral bract scarious and as short as 1 cm or leaflike and to 4(7) cm long, shorter than inflorescence, sometimes sheathing at base for up to | cm; inflorescence 3.5—16 cm long, open-panicu- late, with slender flexuous branches; bractlets subtending branches mostly scarious, some- times frimbriate toward apex; flowers borne singly on slender pedicels to 10 mm long or 2-3 together on short pedicels; bracteoles subtending flowers hyaline, ovate-acute, en- tire to incised; tepals 1.5-2.5 mm _ long, brownish and partly hyaline, acute to acumi- nate, subequal; anthers 0.3—0.4, filaments ca 0.5 mm long; capsules slightly longer than tepals, blackish purple, shiny; seeds 1.2 mm long, with obsolete or inconspicuous carun- cles at each end. Ponderosa pine, aspen, lodgepole pine, spruce-fir, willow-stream- side, and wet meadow communities at 2,300 to 3,475 m in Beaver, Box Elder, Carbon, Daggett, Duchesne, Garfield, Grand, Iron, Juab, Piute, Salt Lake, San Juan, Sanpete, Sevier, Summit, Uintah, and Wasatch coun- ties; circumboreal, extending south in west- ern North America to California and New Mexico; 84 (v). GooDRICH: UTAH FLORA, JUNCACEAE 377 Luzula spicata (L.) DC. Spike Woodrush. Plants 5—40 cm tall: stems more or less caespi- tose from fibrous roots, rhizomes apprently lacking; leaves basal and cauline, blades flat and 1-3 mm wide or involute and to only 0.5 mm wide, margins with scattered to moder- ately dense long-villose hairs; involucral bract 0.7—4 cm long, bractlike or occasionally leaf- like, shorter than or equal to inflorescence or occasionally longer; inflorescence 1-3 cm long often nodding, of ca 4-10 or perhaps more sessile or subsessile heads or short spikes that are congested into a continuous or basally interrupted compound spike, individ- ual heads or spikes with few to several flowers; bractlets subtending spikes or heads scarious and bractlike, or rarely leaflike; bractleoles subtending flowers scarious, fimbriate-ciliate, acuminate-caudate; tepals 2-3 mm long, dark brown or partly hyaline, acuminate or acumi- nate caudate, inner ones a little shorter than outer ones; anthers 0.3—0.5 mm long, ca 1/2 as long or equal to filaments; capsules a little shorter than tepals; seeds 1—-1.3 mm long, caruncle obsolete or inconspicuous, not over 0.2 mm long. Lodgepole pine, spruce, fir, dry meadow, wet meadow, streamside-meadow, alpine tundra, and rarely aspen and oak-— ponderosa pine communities, also in talus and fell fields at 2,470 to 3,810 m in Daggett, Duchesne, Grand, Juab, Piute, Salt Lake, San Juan, Summit, Tooele, and Uintah coun- ties: circumboreal, south in western North America to California and Arizona; 65 (vi). ADVERTISEMENT CALL VARIATION IN THE ARIZONA TREE FROG, HYLA WRIGHTORUM TAYLOR, 1938 Brian K. Sullivan! ABSTRACT. —Advertisement call variation and male mating success was investigated in a population of the Arizona tree frog, Hyla wrightorum, from central Arizona. Dominant frequency of advertisement calls was significantly correlated (negatively) with male snout-vent length. Males found mating were not significantly larger than nonmating males, nor was there a significant correlation between sizes of males and females found in amplexus. These results are discussed in relation to Renaud’s (1977) work with H. wrightorum and in light of recent work with anurans in general. The Arizona tree frog (Hyla wrightorum) is primarily restricted to the Petran Montane Conifer Forest Biome (Brown and Lowe 1980) along the Mogollon Rim in central Arizona and extreme west central New Mexico, and in the Huachuca Mountains of southern Arizona (Steb- bins 1966, Renaud 1977). Other than discussions of taxonomy, to date little has been published concerning this anuran. In his description Taylor (1938) placed H. wrightorum in the H. eximia species group; subsequently a number of au- thors have considered H. wrightorum as a sub- species of H. regilla (Jameson et al. 1966, Jameson and Richmond 1971) or as conspecific with H. eximia (Duellman 1970). However, Re- naud (1977) documented that H. eximia, H. regilla, and H. wrightorum are distinct in their morphology (primarily size) and advertisement calls, and in light of their allopatric distributions warrant recognition as full species. Unfortu- nately Renaud’s work remains unpublished and, hence, largely overlooked. Here I present an analysis of advertisement call variation in a population of H. wrightorum from central Arizona. I compare my results with those of Renaud (1977) and hope to draw atten- tion to his valuable work. I also examine the relationship between male size and mating suc- cess and test the hypothesis that advertisement call parameters (frequency, pulse rate, and dura- tion) are predictably related to male snout-vent length. MATERIALS AND METHODS Breeding aggregations of H. wrightorum were observed near the towns of Pine and Strawberry along state route 87, Gila County, Arizona, on the night of 15 July 1984. A third population was observed breeding at Baker Lake near the intersection of forest route 300 and state route 87, Coconino County, Arizona, on the night of 21 July 1984. Calling males were recorded in the field using a Uher Recorder (4000 IC) and a Shure Unidyne IV (548) micro- phone. Immediately after recording a call, the cloacal temperature of the male was measured with a Schultheis Quick Recording Thermome- ter. Water and air temperatures were recorded at the calling site as well. All recorded males were collected and toe-clipped for permanent identification. A sample of amplexing pairs was collected at Baker Lake to test the prediction that large males were disproportionately suc- cessful in mating (Gatz 1981, Forester and Czarnowsky, 1985). Calls were analyzed with a Kay 6061 B Son- agraph. Dominant frequency of advertise- ment calls was measured to the nearest 100 Hz from a section taken at call midpoint (Fig. 1). It is important to note that in H. wrighto- rum the advertisement call is somewhat fre- quency modulated, increasing about 400 Hz from the beginning to the end ofa call. Rate of amplitude modulation was determined by counting the number of pulses in a 0. 15-sec- ond portion of a wide-band (filter = 300 Hz) audio-spectrogram of three calls for each male, and averaged to yield a mean pulse rate. Similarly, call durations were measured in seconds directly from audio-spectrograms for three calls of each frog, and a mean duration was subsequently calculated. ‘Department of Zoology, Arizona State University, Tempe, Arizona 85287. Present address: Department of Zoology, University of Texas, Austin, Texas 78712-1064. 378 April 1986 SULLIVAN: ARIZONA TREE FROG 379 Cl \y mth wen htiULNAadEU NGL ilitinsvaaui kHz a NO W Seconds Fig. 1. Audio-spectrogram (filter = 300 Hz) of Hyla wrightorum advertisement call. Body temperature = 18 C, snout-vent length = 41 mm. TABLE 1. Mean (x), standard deviation (SD) and extremes for advertisement call parameters, and snout-vent length of Hyla wrightorum from Baker Lake, Arizona (N = 20 for both samples). Présent study Renaud (1977) X SD Extremes x SD Extremes Snout-vent length (mm) 41.30 2.48 37.00— 47.00 37.30 2.64 31.00— 45.00 Dominant frequency (kHz) 1.86 0.20 1.70— 2.30 1.93 0.19 1.60— 2.20 Pulse rate (p/s) 118.10 17.66 83.00—156.00 98.70 12.30 77.50-123.30 Call duration (s) 0.18 0.03 0.13-— 0.24 0.17 0.02 0.12— 0.22 For analysis of variation in advertisement call parameters in relation to size, snout-vent length was measured to the nearest millime- ter witha plastic rule. Product-moment corre- lations between male snout-vent length and each of the call variables were calculated. RESULTS AND DISCUSSION A sufficient sample (N = 20) of calls for analysis was obtained only at Baker Lake on 21 July 1984. All calling males had body temper- atures of 18 + 1 C: hence, temperature-in- duced variation was considered negligible. Further, Renaud (1977) found no correlation between temperature and advertisement call variation in H. wrightorum. Variation in dom- inant frequency, pulse rate, and duration of advertisement calls of H. wrightorum from Baker Lake was within the range of variation documented by Renaud (1977) for this spe- cies. Renaud (1977) also recorded calls of a population of H. wrightorum at Baker Lake (Table 1). There was no significant difference in mean frequency (t = 1.11, P > 0.05) or mean duration (t = 1.24, P > 0.05) of adver- tisement calls recorded by Renaud compared with those of the present study. However, there was a significant difference in mean pulse rate (t = 3.93, P < 0.001). Variation in pulse rate of anuran advertisement calls is typically attributed to variation in body tem- perature (e.g., Gerhardt 1982); hence, these results suggest that pulse rate may be influ- enced by body temperature, contrary to Re- naud’s (1977) assertion. Unfortunately, tem- peratures are unavailable for his Baker Lake recordings. Only dominant frequency was significantly (r = —0.47, P< 0.04, N = 20) correlated with 380 2.0 38 GREAT BASIN NATURALIST Vol. 46, No. 2 42 46 SVL (mm) Fig. 2. Dominant frequency of advertisement calls against snout-vent length for 20 Hyla wrightorum from Baker Lake, Coconino County, Arizona. Body temperatures = 18 + 1C. snout-vent length (Fig. 2). Neither pulse rate (r = —0.24, P = 0.29, N = 20) nor call duration (r = 0.12, P = 0.60, N = 19) were significantly correlated with male size. These results are consistent with a number of studies of anuran vocalizations and support the hypothesis that dominant frequency is determined by compo- nents of the vocal tract directly influenced by male body size (Martin 1972). Many investigators of sexual selection in the Anura have documented a mating advantage for large males (see reviews by Kluge 1981, Ger- hardt 1982). These workers have argued that females might select large mates on the basis of call parameters that are correlated with male snout-vent length. Such discrimination would be possible for H. wrightorum since advertise- ment call frequency is significantly correlated (negatively) with male size, presuming that fe- males could detect small differences in fre- quency. However, males found in amplexus (x = 40.6 mm snout-vent length) were not signifi- cantly (t = 0.96, P > 0.30, N = 30) larger than nonmating males (x = 41.3 mm). Further, there was no correlation between the sizes of males and females in amplexus (r = —0.04, P = 0.90, N = 10). Hence, mating success of males appears unrelated to size, as has been documented for a number of other hylids (Fellers 1979, Gatz 1981, Kluge 1981, Gerhardt 1982, Forester and Czarnowsky 1985). My results, as well as the more substantial work of Renaud (1977), reveal that advertise- ment calls of H. wrightorum are distinct from H. eximia. Calls of H. wrightorum have a higher pulse rate and shorter duration than the majority of Mexican H. eximia (Duellman 1970). It is hoped that this brief report will foster a reassess- ment of the taxonomic status of this Arizona anuran. ACKNOWLEDGMENTS This study was supported by the Department of Zoology at Arizona State University. Eliza- beth Sullivan provided valuable field assistance. April 1986 LITERATURE CITED Brown, D. E., ANDC. H. Lowe. 1980. Biotic communities of the southwest. U.S. Department of Agriculture Forest Service General Technical Report RM-78. DUELLMAN, W. E. 1970. The hylid frogs of middle Amer- ica. Monograph of Kansas University Mus. Nat. Hist. 1: 1-753. FELLERS, G. M. 1979. Mate selection in the gray treefrog, Hyla versicolor. Copeia 1979: 286-290. FORESTER, D. C., AND R. CZARNOWSKY. 1985. Sexual selec- tion in the spring peeper, Hyla crucifer (Am- phibia, Anura): role of the advertisement call. Be- haviour. 92: 112-128. Gatz, A. J. 1981. Size selective mating in Hyla versicolor and Hyla crucifer. J. Herpetology 15: 114-116. GERHARDT, H. C. 1982. Sound pattern recognition in some North American treefrog (Anura: Hylidae): implications for mate choice. Amer. Zool. 22: 581-595. SULLIVAN: ARIZONA TREE FROG 381 JAMESON, D. L., J. P. MACKEY, AND R. C. RICHMOND. 1966. The systematics of the Pacific treefrog, Hyla regilla. Proc. California Acad. Sci. 33: 551-620. JAMESON, D. L., AND R. C. RICHMOND. 1971. Parallelism and convergence in the evolution of size and shape in holartic Hyla. Evolution 25: 497-508. KLUGE, A. G. 1981. The life history, social organization, and parental behavior of Hyla rosenbergi Boulenger, a nest-building gladiator frog. Misc. Publ. Mus. Zool. Univ. Michigan 160: 1-170. Martin, W. F. 1972. Evolution of vocalization in the genus Bufo. Pages 280-309 in W. F. Blair, ed., Evolution in the genus Bufo. University of Texas Press, Austin. RENAUD, M. 1977. Polymorphic and polytypic variation in the Arizona treefrog (Hyla wrightorum). Unpublished dissertation, Arizona State University, Tempe. STEBBINS, R. C. 1966. A field guide to western reptiles and amphibians. Houghton Mifflin Co., Boston. TAYLOR, E. H. 1938. Frogs of the Hyla eximia group in Mexico, with descriptions of two new species. Kansas Univ. Sci. Bull. 25: 421-445. av i! ' i ‘ ii) ’ 6 i , md os 2 i a5 > - try Pearl A f Slike Yay tof att i ee - ; mrs Ue Fe me 1 vom aif i ry ; 1 7 iver SS i* ify inf : i 4) ; r ] ‘ ; 7 | agp ris? 7 t ar: Tn ‘ 7 om Mist: <2 oF ‘ os - ; vt wks ” Scaili be ‘ uf Si vi tee reur ro RT CF ay, ' ' , ' ae itt 4 ale curt At P = i OWA TOD i r , Mf s i tiga $ : é 8 es PSA : ' : ex — ' j & x linked , 4 n 3 100) 7 : \ , 5 it , ‘ > : 4a rts rd > :, an vee ee 4 wer ra yeh oa it ‘ ' " in ’ ‘ are: ¥ i ; os eh PSs Ml i bet i ; Ene f r eee i 2} iL t ari i a scien ; f Pan) on ae i iii sa Mey ih a *, Vrs ; : us x4 " 2 - } } . Fe ee a / . > ) — =e y , " > een Fo = 7 = . i ie ey 7 \ i = ' 1 ‘ 2 (aD a IP LA NOTICE TO CONTRIBUTORS Manuscripts intended for publication in the Great Basin Naturalist or Great Basin Natural- ist Memoirs must meet the criteria outlined in paragraph one on the inside front cover. They should be directed to Brigham Young University, Stephen L. 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No. 6 The bark and ambrosia beetles of North and Central America (Coleptera: Scolytidae), | a taxonomic monograph. $60. ‘No. 7 Biology of desert rodents. $8. No. 8 The black-footed ferret. $10. TABLE OF CONTENTS Biology of Red-necked Phalaropes (Phalaropus lobatus) at the western edge of the Great Basin in fall migration. Joseph R. Jelly Pre .2)56 bes Ra hs i ole aa SS ce ee a etka 185 Some relationships of black-tailed prairie dogs to livestock grazing. Craig J. Knowles. ...................--. 198 Effect of excluding shredders on leaf litter decomposition in two streams. James R. Barnes, J. V. McArthur, and C.F: Cushing. | sie che ee hla ne Uthas oie Satcher dene iS. oa in) Gem eee rrr 204 Three-year surveillance for cestode infections in sheep dogs in central Utah. Ferron L. Andersen, Lauritz A. Jensen, H. Dennis McCurdy, and Craig R: Nichols: 0.5 5. 5002.0. eevee ee oes ee 208 Dam-raised fawns, an alternative to bottle feeding. Kathrin M. Olson-Rutz, Philip J. Urness, and Laura A. - Unrmesss of Sh dials wa ie ae RCE aw Bt A oreitigs ER en, raviagel dame eA UA Te oO rrr rr palTh Subspecific identity of the Amargosa pupfish, Cyprinodon nevadensis, from Crystal Spring, Ash Meadows, Nevada. Jack E. Williams and James E:Deacon:).. 4). 2.0...) ose os er 220 Two aberrant karyotypes in the sagebrush lizard (Sceloporus graciosus): triploidy and a “supernumerary oddity: Pamela Thompson and Jack W. Sites; Jr: ¢.......5- 22.4 6 ieee ce eee needa ee ee 224 Food habits of clouded salamanders (Aneides ferreus) in Curry County, Oregon (Amphibia: Caudata: Pletho- dontidae). John O. Whitaker, Jr., Chris Maser, Robert M. Storm, and Joseph J. Beatty. ................. 228 Wintering bats of the upper Snake River plain: occurrence in lava-tube caves. David L. Genter. ............. 241 Growth rates of mule deer fetuses under different winter conditions. Richard M. Bartmann. ................. 245 Denning habitat and diet of the swift fox in western South Dakota. Daniel W. Uresk and Jon C. Sharps. ....... 249 New taxa and combinations in the Utah flora. Stanley L. Welsh. ................ 000. cece ee eee cee eee eee 254 New taxa in miscellaneous families from Utah. Stanley L-Welsh. .......:2..:. 325.22). 22 eee 261 New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), Part XI. Stephen L. Wood. . 265 Energy and protein content of coyote prey in southeastern Idaho. James G. MacCracken and Richard M. anise nas 3 eet ke i eae Soe Ne tite WR oS on RA 274 Estimates of site potential for Douglas-fir based on site index for several southwestern habitat types. Robert L. Mathiasen, Elizabeth A. Blake, and Carleton B. Edminster. ............... 00 cee cece eee eee ee PUT. Wintering mule deer preference for 21 accessions of big sagebrush. Bruce L. Welch and E. Durant McArthur. . 281 Coleoptera of the Idaho National Engineering Laboratory: an cemsciete checklist. Michael P. Stafford, William F. Barr, and James:B. Johnson... 2... 2 sch yas Gales Pe Sh oa es ce dene ede te 287 Microhabitat affinities of Gambel oak seedlings. Ronald P. Neilson and L. H. Wullstein. .................... 294 | Feeding habits of metamorphosed Ambystoma tigrinum melanostictum in ponds of high pH (> 9). Brian T. | Miller and John H. Larsen, Jr. 2.0 0.0. 05 bee ori See ek oie a err 299 Notes on the Swainson’s Hawk in central Utah: insectivory, premigratory aggregations, and kleptoparasitism. Neil D. Woffinden®.)-. 0 0 PoE ee ws See aa rr 302 — Turkey vultures decline at a traditional roosting site. Daniel M. Taylor. ..........-...020eeeecseteveeeeees 305 | Barn ow! diet includes mammal species new to the island fauna of the Great Salt Lake. Carl D. Marti. ......... 307 | Distributional study of the Zion Snail, Physa zionis , Zion National Park, Utah. David Ng and James R. Barnes. . 310 Blockage and recovery of nitrification in soils exposed to acetylene. Steven J. Burns and Richard E. Terry. ..... 316 New thagriine leafhoppers from the Oriental Region, with a key to 30 species (Homoptera: Cicadellidae: Coelidiinae);.M. W.-Nielson..«. 02.004 ce by 2 Gaid es oe kee a elon oe OAs won ts eee 321 Genus Paralidia with descriptions of new species (Homoptera: Cicadellidae: Coelidiinae). M. W. Nielson. ..... 329 Montane insular butterfly biogeography: fauna of Ball Mountain, Siskiyou County, California. Arthur M. Shapiro 555 Pash wieace eee aes Gila ais ae aae eo ea ik Selgce Nee Midas ant Olt a rr 336 | Comparative habitat and community relationships of Atriplex confertifolia and Sarcobatus vermiculatus in central Utah. Jack D. Brotherson, Lars L. Rasmussen, and Richard D. Black. ...................-..00-- 348 Trees used simultaneously and sequentially by breeding cavity-nesting birds. Kevin J. Gutzwiller and Stanley | Hi: Andersone 3)....8nerihic va dons ae Vy ie Neyer c ae ines Higa 3 Sel a 358 | New species of Mentzelia (Loasaceae) from Grand County, Utah. Barry A. Prigge. ................00-00000, 361 | Utah flora: Juncaceae. Sherel Goodrich. +2. 62.632 04.0624 el we) oo Oe eee 366 | Advertisement call variation in the Arizona tree frog, Hyla wrightorum Taylor, 1938. Brian K. Sullivan. ....... 378 | | HE GREAT BASIN NATURALIST lume 46 No. 3 31 July 1986 Brigham Young University \ f } Shon Dent NARS =p = es \ GREAT BASIN NATURALIST Editor. Stephen L. Wood, Department of Zoology, 290 Life Science Museum, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Chairman, Botany; James R. Barnes, Zoology; Hal L. Black, Zoology; Stanley L. Welsh, Botany; Clayton M. White, Zoology. All are at Brigham Young University, Provo, Utah 84602. Ex Officio Editorial Board Members. Bruce N. Smith, Dean, College of Biological and Agricultural Sciences; Norman A. Darais, University Editor, University Publications. Subject Area Associate Editors. Dr. Noel H. Holmgren, New York Botanical Garden, Bronx, New York 10458 (Plant Taxonomy). Dr. James A. MacMahon, Utah State University, Department of Biology, UMC 53, Logan, Utah 84322 (Vertebrate Zoology). Dr. G. Wayne Minshall, Department of Biology, Idaho State University, Pocatello, Idaho 83201 (create Biology). Dr. Ned K. Johnson, Museum of Vertebrate Zoology and Department of Zoology, University of California, Berkeley, California 94720 (Ornithology). Dr. E. Philip Pister, Associate Fishery Biologist, California Department of Fish and Game, 407 West Line Street, Bishop, California 93514 (Fish Biology). Dr. Wayne N. Mathis, Chairman, Department of Entomology, National Museum of | Natural History, Smithsonian Institution, Washington, D.C. 20560 (Entomology). Dr. Theodore W. Weaver III, Department of Botany, Montana State University, Boze- man, Montana 59715 (Plant Ecology). The Great Basin Naturalist was founded in 1939 and has been nnbligeed from one to four times a year since then by Brigham Young University. Previously unpublished manuscripts in English of fewer than 100 printed pages in length and pertaining to the biological natural history of western North America are accepted. Western North America is considered to be west of the Mississippi River from Alaska to Panama. The Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natural history longer than can be accom- modated 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 for private individuals | is $16; for institutions, $24 (outside the United States, $18 and $26); and for student subscrip- tions, $10. The price of single issues is $6 each. All back issues are in print and are available for | sale. All matters pertaining to subscriptions, back issues, or other business should be directed | to Brigham Young University, Great Basin Naturalist , 290 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 foun | through a continuing exchange of scholarly publications should contact the Brigham Young | University Exchange Librarian, Harold B. Lee Library, Provo, Utah 84602. Manuscripts. See Notice to Contributors on the inside back cover. 10-86 650 25994 ISSN 017-3614 The Great Basin Naturalist PUBLISHED AT PROVO, UTAH, By BRIGHAM YOUNG UNIVERSITY ISSN 0017-3614 VOLUME 46 31 July 1986 No. 3 NORTH AMERICAN STONEFLIES (PLECOPTERA): SYSTEMATICS, DISTRIBUTION, AND TAXONOMIC REFERENCES B. P. Stark’, S. W. Szczytko’, and R. W. Baumann® ABSTRACT.—A list of 537 Nearctic stonefly species is provided and distributions of all species are given by U.S. state and Canadian province. The list includes a bibliography of systematic and biogeographic papers published since the Zwick (1973) catalogue. Periodically, it is useful to update catalog information on such well-studied groups as the nearctic Plecoptera. The most recent up- date by Baumann (1976) included distribu- tions of genera by regions and reported a total of 470 nearctic species; prior to that list, the Illies (1966) and Zwick (1973) catalogs pro- vided sources from which distributional data could be extracted. Unfortunately, not only are these works somewhat unavailable to workers, but also they are somewhat out of date. Consequently, we present this list of 537 species currently recognized (as of January 1986) from North America. Distribution records are reported by standard postal ser- vice abbreviation for U.S. states and Canadian provinces; Mexican records are given as MX, but only those species of nearctic origin are included because of the poor state of knowl- edge of the neotropical Anacroneuria. The list includes unpublished records provided by B. C. Kondratieff and R. F. Kirchner, along with a few from our personal collections, and the bibliography includes only those papers published since the Zwick (1973) catalog that contain new state or province records, or which reflect systematic changes for nearctic Plecoptera. ‘Department of Biology, Mississippi College, Clinton, Mississippi 39058. Since this list is likely to expand and un- dergo change as generic and specific limits are refined and phylogenetic affinities are postu- lated, we are maintaining the list in computer files. Currently we are using IBM and AT&T personal computers with Wordstar and Word- Perfect software and a fortran program devel- oped at Mississippi College for the HP 3000 mainframe academic computer. We suggest others with a vital interest in Plecoptera may also want to adopt this approach. LIST OF ABBREVIATIONS AB Alberta KS Kansas AL Alabama KY Kentucky AK Alaska LA Louisiana AR Arkansas LB Labrador AZ Arizona MA _ Massachusetts BC __ British Columbia MB Manitoba CA California MD _ Maryland CO Colorado ME Maine CT Connecticut MI Michigan DC _ District of Columbia MN _ Minnesota DE Delaware MO Missouri FL Florida MS Mississippi GA Georgia MT Montana IO Iowa MX Mexico ID Idaho NB Nebraska IL Illinois NC _ North Carolina IN Indiana ND North Dakota College of Natural Resources, University of Wisconsin, Stevens Point, Wisconsin 54481. 3Life Science Museum and Department of Zoology, Brigham Young University, Provo, Utah 84602. 383 GREAT BASIN NATURALIST 384 NF Newfoundland RI Rhode Island NH New Hampshire SC South Carolina NJ New Jersey SD South Dakota NM_ New Mexico SK Saskatchewan NS Nova Scotia ™N Tennessee NT Northwest Territories TX Texas NV Nevada UT Utah NW_ New Brunswick VT Vermont NY New York VA _ Virginia OH Ohio WA Washington OK Oklahoma WI ‘Wisconsin ON Ontario WV West Virginia OR Oregon WY Wyoming PA Pennsylvania YK Yukon QB Quebec ACKNOWLEDGMENTS E We thank B. C. Kondratieff for reviewing an early draft of the list and L. C. Whitlock for his expertise in developing the HP 3000 program. UHOLOGNATHA Capniidae Capniinae Allocapnia aurora Ricker brooksi Ross cunninghami Ross & Ricker curiosa Frison forbesi Frison frisoni Ross & Ricker frumi Kirchner fumosa Ross granulata (Claassen) harperi Kirchner illinoensis Frison indianae Ricker jeanae Ross loshada Ricker malverna Ross NU, YG, @A, MD, MS, NC, SC, TN, VA TN KY, TN KY, MD, NY, PA, VA, WV IL, IN, KY, OH, TN, WV KY, NY, OH, PA, TN, VA, WI, WV WV NC, TN, VA AL, AR, DG, IA, IL, IN, KS, KY, LA, MB, MD, MI, MN, MO, MS, NJ, NY, OH, OK, ON, PA, QB, TN, TX, VA, WI, WV VA, WV IL, IN, ME, MN, NY, OH, ON, QB, VA, WI IN, KY, NY, OH AR, OK TN, VA, WV AR, LA, OK, TX maria Hanson minima (Newport) mohri Ross & Ricker mystica Frison nivicola (Fitch) ohioensis Ross & Ricker ozarkana Ross pechumani Ross & Ricker peltoides Ross & Ricker perplexa Ross & Ricker E polemistis Ross & Ricker pygmaea (Burmeister) recta (Claassen) rickeri Frison sandersoni Ricker simmonsi Kondratieff & Voshell smithi Ross & Ricker stannardi Ross tennessa Ross & Ricker unzickeri Ross & Yamamoto virginiana Frison Vol. 46, No. 3 CT, MA, MD, ME, NH, NS, NW, NY, PA, QB, VA, VT, WV CT, IN, MA, ME, MI, MN, NF, NH, NS, NW, NY, ON, QB, VT, WI AR, MO, OK AL, AR, GA, IL, IN, KY, MO, MS, OH, TN, VA, WV NIL, (CA, IDYC, DE, IL, IN, KY, MA, MD, NC, NJ, NS, NW, NY, OH, PA, | QB, RI, TN, VA, | VT, WI, WV | IN, KY, NY,OH, | : AR NW, NY, QB | AR, OK | TN AL CT, DC, IN, KY, MA, MD, ME, MI, MN, MO, NH, NS, NW, NY, ON, PA, QB, RI, TN, VA, VT, WI, WY ALIGN DES DE, GA, IL, IN, KY, MA, MD, ME, MS, NC, NH, NS, NY, OH, ON, PA, QB, SC, TN, VA, VT, WI, WV AL, AR, DC, DE, GA, IA, IN, KS, KY, MD, MN, MO, MS, NC, NJ, NY, OH, OK,ON, | PA, TN, VA, WI, | San | AR, MO, OK VA AL, IL, KY, OH NC, TN, VA | AL, TN TN AL, DE,GA, | i | | MS, NC, SC, VA July 1986 vivipara (Claassen) warreni Ross & Yamamoto wrayi Ross zola Ricker Bolshecapnia gregsoni (Ricker) maculata (Jewett) milami (Nebeker & Gaufin) rogozera (Ricker) sasquatchi (Ricker) spenceri (Ricker) Capnia bakeri (Banks) barbata Frison barberi Claassen californica Claassen cheama Ricker coloradensis Claassen confusa Claassen cygna Jewett decepta (Banks) disala Jewett elevata Frison elongata Claassen ensicala Jewett erecta Jewett excavata Claassen fibula Claassen glabra Claassen gracilaria Claassen hornigi Baumann & Sheldon jewetti Frison lacustra Jewett licina Jewett lineata Hanson manitoba Claassen melia Frison nana Claassen nearctica Banks nedia Nebeker & Gaufin oregona Frison petila Jewett DG, IA, IL, IN, KS, KY, MD, MI, MN, MO, NY, OH, OK, ON, PA, QB, TN, VA, WI, WV NW, NY, OH, PA, TN, VA, WV AB, BC, MT AZ, CA AB, BC, MT, YK CO, ID, MB, MT, NM, SK, WY, YK AB, AK, BC, CO, AB, AK, BC, CO, ID, MT, OR, UT, AB, ID, MT, OR, WY STARK ET AL.: NORTH AMERICAN PLECOPTERA pileata Jewett promota Frison quadrituberosa Hitchcock scobina Jewett sextuberculata Jewett spinulosa Claassen sugluka Ricker teresa Claassen tumida Claassen uintahi Gautin umpqua Frison utahensis Gaufin & Jewett venosa (Banks) vernalis (Newport) wanica Frison willametta Jewett zukeli Hanson Eucapnopsis brevicauda Claassen Isocapnia abbreviata Frison agassizi Ricker crinita (Needham & Claassen) fraseri Ricker grandis (Banks) hyalita Ricker integra Hanson missouri Ricker mogila Ricker spenceri Ricker vedderensis (Ricker) Mesocapnia arizonensis (Baumann & Gaufin) autumna (Baumann & Gaufin) bergi (Ricker) frisoni (Baumann & Gaufin) lapwae (Baumann & Gaufin) oenone (Neave) ogotoruka (Jewett) porrecta (Jewett) projecta (Frison) variabilis (Klapalek) werneri (Baumann & Gaufin) yoloensis (Baumann & Gaufin) Nemocapnia carolina Banks 385 BC, OR BC, CA, OR, WA CA CA AB, MT, OR CA QB CA BC, CA, OR ID, NV, UT, WY CA, OR NV, UT OR, WA AB, BC, CO, ID, LB, MB, MN, AB, AK, AZ, BC, CA, CO, ID, MT, NM, NV, OR, UT, WY CA, OR BC, OR, WA CO, ID, MT, SK, UT BC AB, AK, BC, CA, CO, MT, OR, UT CO, ID, MT, UT AB, MT AB, ID, MT, SK, AZ, NM AK, BC, OR, WA AK CO, NM, TX, UT CA, OR, WA BC, CA, OR, WA AK, BC AZ, CA, NM CA, OR AL, AR, FL, IL, IN, MS, NC, QB, SC, VA 386 Paracapnia angulata Hanson opis (Newman) oswegaptera (Jewett) Utacapnia columbiana (Claassen) distincta (Frison) imbera (Nebeker & Gaufin) labradora (Ricker) lemoniana (Nebeker & Gaufin) logana (Nebeker & Gaufin) poda (Nebeker & Gaufin) sierra (Nebeker & Gaufin) tahoensis (Nebeker & Gaufin) trava (Nebeker & Gaufin) Leuctridae Leuctrinae Despaxia augusta (Banks) Leuctra alabama James alexanderi Hanson alta James baddecka Ricker biloba Claassen carolinensis Claassen cottaquilla James crossi James duplicata Claassen ferruginea (Walker) grandis Banks laura Hitchcock maria Hanson mitchellensis Hanson moha Ricker monticola Hanson nephophila Hanson GREAT BASIN NATURALIST CORCIEDE SIE MA, MB, MD, ME, MI, NC, OH, OK, PA, QB, SK, TN, VA, WI, WV, WY CT, ME, MN, NF, NY, ON, AB, AK, BC, CA, MT, OR, YK AB, ID, MT, WY OR, WA LB, QB CO, ID, NV, UT, CO, NM, UT, CO, MT, NM, UT, WY CA, NV AB, ID, MB, MT, SK AK, BC, CA, ID, MT, OR, WA KY, NC, SC, TN, VA, WV AL, GA, NC, ON, TN, VA NC, PA, SC, TN, AL, FL, MS ON, PA, QB, VA, WV AL, CT, DE, FL, IL, KY, MA, ME, MN, MS, NS, OH, PA, QB, SC, SK, VA, WI, WV MA, ME, NC, NC, SC. TN, VA NG, TN rickeri James sibleyi Claassen szczytkoi Stark & Stewart tenella Provancher tenuis (Pictet) triloba Claassen truncata Claassen variabilis Hanson Moselia infuscata (Claassen) Paraleuctra andersoni Harper & Wildman divisa (Hitchcock) forcipata (Frison) jewetti Nebeker & Gaufin occidentalis (Banks) purcellana (Neave) rickeri Nebeker & Gaufin sara (Claassen) vershina Gaufin & Ricker Perlomyia collaris Banks utahensis Needham & Claassen Zealeuctra arnoldi Ricker & Ross cherokee Stark & Stewart claasseni (Frison) Vol. 46, No. 3 AL, IL, KY, MS, OH CT, DE, IL, IN, KY, MA, MD, ME, NG, NY, OH, ON, PA, QB, SC, TN, VA, WI, WV LA CT, IN, LB, MA, ME, MN, MS, NJ, NS, NW, NY, ON, PA, QB, WI IMG INR Clk, DIE, IL, MA, ME, MI, MN, MO, MS, NJ, NS, NY, OH, OK, ON, PA, QB, SC, TN, VA, WI, WV FL, NC, NY, QB, SC, VA CT, ME, NY, PA, QB, VA, WV MA, ME, NH, VA, VT CA, OR AB, AK, BC, CA, ID, MT, OR CO, MT, UT AB, AK, BC, CA, CO, ID, MT, NM, NV, OR, UT, WY AB, BC, MT, OR, WY AK, ID, MT, NM, UT CE DE ke MA, ME, NC, NY, ON, QB, SC, TN, VA, WV AB, AK, BC, CA, CO, ID, MT, NM, NV, OR, SK, UT, WA, WY BC, CA, ID, OR AB, BC, CA, CO, MT, NM, NV, OR, UT, WA, WY TX AR, OK AR, IL, IN, KS, KY, MO, OK, TX, VA July 1986 STARK ET AL.: NORTH AMERICAN PLECOPTERA fraxina Ricker & Ross hitei Ricker & Ross narfi Ricker & Ross warreni, Ricker & Ross washita Ricker & Ross Megaleuctra Megaleuctrinae complicata Claassen flinti Baumann kincaidi Frison sierra Fields stigmata (Banks) williamsae Hanson Amphinemura Nemouridae Amphinemurinae apache Baumann & Gaufin banksi Baumann & Gaufin delosa (Ricker) linda (Ricker) mexicana Baumann mockfordi (Ricker) mogollonica Baumann & Gaufin nigritta (Provancher) puebla Baumann reinerti Baumann varshava (Ricker) venusta (Banks) wui (Claassen) Malenka bifurcata (Claassen) biloba (Claassen) californica (Claassen) coloradensis (Banks) cornuta (Claassen) depressa (Banks) IL, KY, OH, TN, WV TX AR, IL, MO, Ok, MT, WA SC, TN, VA AZ AZ, CO, ID, MT, SD, UT, WY AL, AR GAU IE IN, KS, KY, MI, MO, MS, OH, OK, ON, PA, QB, TN, VA, WI, WV AB, AK, BC, CO, LB, MB, MI, NT, ON, PA, QB, SK, WI MX TN AZ, NM, UT AL, CT, DE, FL, © Ties INS Kee LA, LB, ME, MS, NS, OH, PA, QB, SC, TN, TX, VA, WV WV AB, BC, CA, CO, ID, MB, MT, NV, SK, UT, WY AZ, CO, NM, SD, UT, WY BC, OR, WA CA, OR flexura (Claassen) marionae (Hitchcock) perplexa (Frison) tina (Ricker) wenatchee (Ricker) Nemourinae Lednia tumana (Ricker) Nemoura arctica Esben-Petersen normani Ricker rickeri Jewett spiniloba Jewett trispinosa Claassen Ostrocerca albidipennis (Walker) complexa (Claassen) dimicki (Frison) foersteri (Ricker) prolongata (Claassen) truncata (Claassen) Paranemoura perfecta (Walker) Podmosta decepta (Frison) delicatula (Claassen) macdunnoughi (Ricker) obscura (Frison) weberi (Ricker) Prostoia besametsa (Ricker) completa (Walker) 387 AB, CO, ID, MT, ID, MT, NV, OR, WA WA MT AB, AK, BC, CT, MA, ME, NH, NS, NY, ON, QB, VA CT, MA, ME, NY, ON, PA, QB, VA, WV BC, OR, WA BC, OR, WA DE, ME, NH, NW, QB, VA, WV CT, MA, MD, ME, NY, OH, ON, PA, QB, VA, WV CT, ME, NC, NS, NY, ON, PA, QB, VA, WV AB, AK, BC, CO, ID, MT, OR, UT, WA AB, BC, CA, CO, ID, MT, NM, NV, OR, SK, UT, WY LB, ME, NF, NS, QB OR, WA AK, YK ING, BIG, GA, CO, ID, MT, NM, NV, OR, UT, WY AL, AR, DE, IL, MA, ME, MI, MN, MO, MS, NC, NS, OK, ON, QB, SC, VA, WI, WV 388 hallasi Kondratieff & Kirchner similis (Hagen) Shipsa rotunda (Claassen) Soyedina carolinensis (Claassen) interrupta (Claassen) nevadensis (Claassen) potteri (Baumann & Gaufin) producta (Claassen) vallicularia (Wu) washingtoni (Claassen) Visoka cataractae (Neave) Zapada chila (Ricker) cinctipes (Banks) columbiana (Claassen) cordillera (Baumann & Gaufin) frigida (Claassen) glacier (Baumann & Gaufin) haysi (Ricker) oregonensis (Claassen) wahkeena (Jewett) Taeniopterygidae Brachypteryinae Bolotoperla rossi (Frison) Doddsia occidentalis (Banks) GREAT BASIN NATURALIST VA CT, DE, IN, KY, MA, ME, MI, MN, OH, QB, SC, VA, WI, WV AB, AK, AL, MB, MD, ME, MI, MN, NT, ON, QB, SC, SK, VA, WI DE, NC, TN, VA, WV BC, OR, WA CA, NV ID, MT BC, CA, OR, WA CT, IN, KY, ME, MI, NS, NY, OH, ON, PA, QB, VA, WI, WV CT, ME, NH, PA AB, BC, CA, ID, MT, OR, WA TN AK, BC, CA, CO, ID, MB, MT, NM, NV, SD, SK, UT, WY AB, AK, BC, CA, ID, MT, OR, UT, WA, WY CA, ID, MT, OR, WA AB, AK, BC, CA, CO, ID, MT, AB, AK, BC, CA, CO, ID, MT, NM, NV, OR, UT, WA, WY AB, AK, BC, CA, CO, ID, MT, NV, OR, WY, YK OR ME, NC, NH, QB, VA, WV AK, BC, CA, CO, ID, MT, NM, NV, OR, UT, WA, WY Oemopteryx contorta (Needham & Claassen) fosketti (Ricker) glacialis (Newport) vanduzeea (Claassen) Strophopteryx appalachia Ricker & Ross arkansae Ricker & Ross cucullata Frison fasciata (Burmeister) inaya Ricker & Ross limata (Frison) ostra Ricker & Ross Taenionema atlanticum Ricker & Ross Vol. 46, No. 3 CT, KY, MA, ME, NH, TN, VA, WV AB, CO, MB, MT, SK, UT CT, MN, NY, ON, QB, WI CA NC, TN, VA, WV AR, MO OK AL, CT, DE, IN, KS, KY, MB, ME, MN, MS, NC, ND, OH, OK, PA, QB, SC, WI NC, SC TN, VA AR, OK, TX CT, KY, LB, MA, MD, ME, NC, NE, NH, NY, QB, TN, VA, WV californicum (Needham & Claassen)CA grinnelli (Banks) nigripenne (Banks) oregonense (Needham & Claassen) pacificum (Banks) pallidum (Banks) raynorium (Claassen) Taeniopteryginae Taeniopteryx burksi Ricker & Ross lita Frison lonicera Ricker & Ross CA AB, AZ, BC, CA, CO, ID, MT, NM, NV, OR, UT, WA, WY OR, WA AB, AZ, BC, CA, CO, ID, MT, NM, OR, UT, WA, WY BC, CA, CO, MT, NM, OR, UT, WA CA AL, CT, DE, FL, IA, IL, IN, KS, KY, LA, MD, ME, MN, MO, MS, OH, OK, ON, PA, QB, TN, TX, VA, WI, WV AL, AR, FL, IL, IN, KY, LA, MS, NGINIAS@ xe VA ALS ED S@AION MD, MS, SC, TN, TX, VA July 1986 maura (Pictet) metequi Ricker & Ross nelsoni Kondratieff & Kirchner nivalis (Fitch) parvula Banks pecos Baumann & Jacobi robinae Kondratieff & Kirchner starki Stewart & Szczytko ugola Ricker & Ross SYSTELLOGNATHA Chloroperlidae Chloroperlinae Alloperla acadiana Harper aracoma Harper & Kirchner atlantica Baumann banksi Frison biserrata Nelson & Kondratieff caudata Frison chandleri Jewett chloris Frison concolor Ricker delicata Frison fraterna Frison furcula Surdick hamata Surdick idei (Ricker) imbecilla (Say) AL, AR, CT, DE, GA, IN, KY, MA, MD, ME, MN, MS, NC, OH, PA, SC, TN, TX, VA, WV AL, AR, IL, IN, KS, KY, OH, AB, CA, CT, DE, ID, IL, IN, MB, ME, MN, NY, ON, OR, PA, QB, SK, UT, WI AB, AR, CT, GA, IN, KY, MB, ME, MI, MN, MS, NY, OH, ON, PA, QB, SC, TN, VA, WI, WV NW WV CT, GA, IN, MA, MD, ME, MI, MN, NC, NH, NS, NW, NY, ON, PA, QB, SC, TN, VA, VT IL, ME, MI, NS, NY, ON, VA VA AR, CT, IL, IN, MA, ME, MO, AB, BC, CA, ID, MT, OR AK, BC, CA, OR, QB, VA IN, KY, NY, OH, QB, VA, WV STARK ET AL.: NORTH AMERICAN PLECOPTERA leonarda Ricker medveda Ricker nanina Banks natchez Surdick & Stark neglecta Frison ouachita Stark & Stewart pilosa Needham & Claassen roberti Surdick serrata Needham & Claassen severa (Hagen) usa Ricker voinae Ricker vostoki Ricker Bisancora pastina (Jewett) rutriformis Surdick Chloroperla ovibovis (Ricker) Haploperla brevis (Banks) chilnualna (Ricker) chukcho (Surdick & Stark) orpha (Frison) Neaviperla forcipata (Neave) Plumaperla diversa (Frison) spinosa (Surdick) Rasvena terna (Frison) Suwallia autumna (Hoppe) dubia (Frison) 389 ME, MI, MN, MO, QB AB, BC, ID, MT, WY, YK GA, NG, NY, SC, TN, VA NC, NF, NY, AB, AK, BC, ID, MT, WA, WY AB, AK, BC, CA, CO, ID, MT, NT, NV, OR, UT, WA, WY GA, SC, TN, VA, WV MA, ME, NS, NY, QB, VT NS, NY, PA AK, NT AB, AL, AR, BC, CT, DE, GA, IL, IN, KY, MA, MB, MD, ME, MI, MN, MO, NC, NJ, NS, NW, NY, OH, OK, ON, PA, QB, SC, SK, TN, VA, WI, WV BC, CA, OR, WA AB, AK, BC, MT, WA AB, AK, BC, CA, CO, ID, MT, NV, OR, UT, WA, YK CA NH, NY, TN, VT, WV AB, BC, CA, ID, MT, OR, WA, WY AB, AK, BC, CO, ID, MT, OR, UT, WN, WY 390 lineosa (Banks) marginata (Banks) pallidula (Banks) Sweltsa albertensis (Needham & Claassen) borealis (Banks) californica (Jewett) coloradensis (Banks) continua (Banks) exquisita (Frison) fidelis (Banks) gaufini Baumann hondo Baumann & Jacobi lamba (Needham & Claassen) lateralis (Banks) mediana (Banks) naica (Provancher) occidens (Frison) onkos (Ricker) oregonensis (Frison) pacifica (Banks) revelstoka (Jewett) tamalpa (Ricker) townesi (Ricker) urticae (Ricker) Triznaka pintada (Ricker) signata (Banks) GREAT BASIN NATURALIST BG, COMI: MT, OR, SK, UT, WA, WY LB, MA, ME, NY, PA, QB, VA, WV, WI AB, AK, AZ, BC, CA, CO, ID, MB, MT, NM, NV, OR, UT, WA, WY AB, ID, MT, WY AB, AK, BC, CA, CO, ID, MT, AK, BC, OR, WA AB, AK, BC, CA, CO, ID, NV, NM, OR, UT, WY CT, GA, MA, ME, NC, NH, PA, QB, SC, VA NG, CIRC, DN, TN, VA IN, LB, ME, NS, NY, PA, QB, VA BC, ID, MT, OR, WA DE SKA: ME, NG, NS, NY, OH, ON, PA, QB, VA AK, BC, OR, WA AK, BC, CA, ID, OR, WA, WY CA, NV NC, VA AZ, CA, CO, ID, NM, NV, SD, UT, WA, WY AK, BC, CO, ID, MT, NM, OR, SD, SK, UT, WA, WY Paraperlinae Kathroperla perdita Banks Paraperla frontalis (Banks) wilsoni Ricker Utaperla gaspesiana Harper & Roy sopladora Ricker Peltoperlidae Peltoperlinae Peltoperla arcuata Needham Sierraperla cora (Needham & Smith) Soliperla campanula (Jewett) fenderi (Jewett) quadrispinula (Jewett) sierra Stark thyra (Needham & Smith) tillamook Stark Tallaperla anna (Needham & Smith) cornelia (Needham & Smith) elisa Stark laurie (Ricker) lobata Stark maria (Needham & Smith) Viehoperla ada (Needham & Smith) Yoraperla brevis (Banks) mariana (Ricker) Perlidae Acroneuriinae Acroneuriini Acroneuria abnormis (Newman) Vol. 46, No. 3 AB, BC, CA, ID, MT, NV, OR, WA AB, AK, BC, CA, CO, ID, MT, NM, OR, SD, UT, WA, WY, YK AB, BC, CA, ID, MT, OR, WA, YK NH, PA, QB, WV AB, AK, BC, ID, MT, NV, UT, WY, YK KY, NY, PA, QB, TN, VA, WV CA, NV, OR GA, NC, PA, SC, VA, FL, GA, NC, SC NC, TN NC, NH, NY, _| PA, SC, TN, VA, | WV GA, NC,SC,TN_ | AB, BC, CA, ID, | MT, NV, OR, | WA, WY ; BC, CA, OR, WA | AB, AL, CO, CT, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, MB, July 1986 arenosa (Pictet) arida (Hagen) carolinensis (Banks) evoluta Klapalek filicis Frison flinti Stark & Gaufin internata (Walker) lycorias (Newman) mela Frison perplexa Frison petersi Stark & Gaufin Attaneuria ruralis (Hagen) Beloneuria georgiana (Banks) jamesae Stark & Szczytko stewarti Stark & Szcezytko Calineuria californica (Banks) Doroneuria baumanni Stark & Gaufin theodora (Needham & Claassen) MD, ME, MI, MN, MS, MT, NB, NC, NM, NW, NY, OH, ON, PA, QB, SC, SK, TN, UT, VA, WI, WV, WY AL, DC, DE, In, GA, uN, MD, ME, MS, NJ, PA, QB, SC, TX, VA, WV GA, NC, NJ, PA, TN CT, KY, MA, MB, ME, MN, NC, NJ, NY, OH, ON, PA, QB, SC, TN, VA, WV AR, IL, IN, KS, KY, MI, MO, OH, OK, ON, PA, TN, VA, WV IN, AVS, Gh, Uke AR, GA, IL, IN, KY, MI, MN, MO, OK, VA, WI, WV AR. Git, Wi, LO, MB, ME, MI, MN, NY, OH, ON, PA, QB, SK, TN, VA, WI, WV AL, AR, FL, GA, IL, IN, KS, MO, MS, OK, PA, TX AL, AR, DC, GA, iit, IN, OG MO, OH, OK, PA, TN, WV GA, TN AL, AR, DC, FL, GA, IA, IL, IN, KS, MB, MD, MN, MO, MS, NC, PA, SC, TN, VA, WI NG, SC, TN AB, BC, CA, ID; MT, OR, WA BC, CA, NV, OR, WA AB, BC, ID, MT, UT, WY STARK ET AL.: NORTH AMERICAN PLECOPTERA Eccoptura xanthenes (Newman) Hansonoperla appalachia Nelson Hesperoperla hoguei Baumann & Stark pacifica (Banks) Perlesta frisoni Banks placida (Hagen) Perlinella drymo (Newman) ephyre (Newman) fumipennis (Walsh) Anacroneurlini Anacroneuria wipukupa Baumann & Olson Perlinae Neoperlini Neoperla carlsoni Stark & Baumann catharae Stark & Baumann choctaw Stark & Baumann clymene (Newman) freytagi Stark & Baumann 391 AL, CT, DE, FL, GA, KY, MD, MS, NC, OH, PA, SC, TN, VA, WV MA, TN, WV CA AB, AK, AZ, BC, CA, CO, ID, MT, NM, NV, OR, SD, SK, UT, WA, WY NG, SC, TN EAR AZ ACI DC, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, MB, ME, MO, MS, NB, OH, OK, ON, PA, QB, SC, SK, TN, TX, UT, VA, WI, WV, WY AL, AR, CT, FL, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MO, MS, NY, OH, OK, QB, SC, TX; VA, WI, WV Ni, NR, GR, Ia, IN, KY, LA, MD, MN, MO, MS, NJ, OK, PA, SC, VA, WI, WV ING, GAP, [8k Ta, MS, SC AZ FL, LA, MS, OKGSCabx AR, OH, VA OK, WV Nit, Cit, IL, GA, IN, KS, LA, MS, NY, OK, ON, PA, QB, TX, VA, WI, WV AR, KY, ME, NY, OH, ON, SC, TN, VA 392 gaufini Stark & Baumann mainensis Banks stewarti Stark & Baumann Perlini Agnetina capitata (Pictet) Claassenia sabulosa (Banks) Paragnetina fumosa (Banks) ichusa Stark & Szczytko immarginata (Say) kansensis (Banks) media (Walker) Perlodidae Isoperlinae Calliperla luctuosa (Banks) Cascadoperla trictura (Hoppe) Clioperla clio (Newman) Isoperla acula Jewett adunca Jewett baumanni Szczytko & Stewart bellona Banks bifurcata Szezytko & Stewart GREAT BASIN NATURALIST IN, KY, OH ME, OH AR, KY, ME, MS, OH, VA, WI CR DCyDE FL, GA, IN, KS, KY, LA, MA, MB, MD, ME, MN, MO, MS, NC, NH, NY, OH, OK, ON, PA, QB, SC, TN, VA, WI, WV AB, AZ, BC, CA, CO, ID, MB, MT, NM, OR, QB, SD, SK, UT, WA, WY IN GIDYO), 1b GA, LA, MS, NCIS DXeAVA NC, SC, TN CIDENCAG MA, ME, NC, NH, NY, PA, QB, SC, TN, VA, WV Nig iby GAY Tt, IN, KS, LA, MS, SC CT, DE, IN, KY, MB, ME, MI, MO, NH, OH, ON, PA, QB, SK, VA, WI, WV CA, OR, WA BC, CA, ID, MT, OR, WA AL, AR, CT, DE, FL, GA, IL, IN, KY, MA, MD, MI, MO, MS, NC, OH, OK, ON, PA, SC, TN, VA, WI, WV GA, NC CA, ID, OR, WA bilineata (Say) burksi Frison conspicua Frison cotta Ricker coushatta Szczytko & Stewart davisi James decepta Frison decolorata (Walker) denningi Jewett dicala Frison distincta Nelson emarginata Harden & Mickel extensa Claassen francesca Harper frisoni Mlies fulva Claassen fusca Needham & Claassen gibbsae Harper gravitans (Needham & Claassen) holochlora (Klapalek) irregularis (Klapalek) jewetti Szczytko & Stewart katmaiensis Szczytko & Stewart lata Frison longiseta Banks major Nelson & Kondratieft marlynia (Needham & Claassen) Vol. 46, No. 3 CONCIMIIESIINE KS, KY, MA, MB, ME, MI, MN, MS, NB, NC, NF, NW, NY, OH, ON, PA, QB, SK, VA, WI, WV AR, IL, IN, KY, OH, WV MS, OK, TX, VA AL IL, IN, OH, ON AK, BC, MB, CT, FL, IN, MA, MB, ME, MI, MN, MO, ON, PA, QB, SC, TN, VA, WI, WV NC, SC, TN MN NY, QB BC, CT, DE, IN, MB, ME, MN, NS, NW, OH, QB, WI AB, AZ, BC, CA, CO, ID, MT, NM, NV, OR, UT, WA, WY AB, BC, ID, MT, OR, WA, WY, YK CT, NY, QB, WV OR, WA CT, DE, GA, KY, MA, ME, NC, NY, PA, QB, SC, TN, VA, WV MN, NS, ON, QB, TN, VA, WI, WV AB, BC, CO, IA, ID, IL, KS, MB, MN, MO, MT, IL, IN, KS, KY, MB, ME, MI, MN, NB, NH, NJ, NS, NW, OK, ON, PA, QB, SC, SK, VA, WI, WV July 1986 marmorata Needham & Claassen maxana Harden & Mickel mohri Frison montana (Banks) mormona Banks namata Frison nana (Walsh) orata Frison ouachita Stark & Stewart petersoni Needham & Christenson phalerata (Smith) pinta Frison quinquepunctata (Banks) raineri Jewett richardsoni Frison roguensis Szczytko & Stewart sagittata Szczytko & Stewart signata (Banks) similis (Hagen) slossonae (Banks) sobria (Hagen) sordida Banks tilasqua Szczytko & Stewart transmarina (Newman) CA, NV, OR, WA MN AR, TIL, ES, Thy MO, OK, PA CT, DE, ME, MN, NH, NS, NY, ON, PA, QB Wh, XC, GA, CO, AR, IN, KY, ME, MO, OH, PA, VA, WV IL, IN, KY, OH, ON, QB, WI CT, FL, ME, MN, NC, NH, NS, NW, NY, OH, PA, QB, SC, TN, VA, VT, WV AR, OK AB, AK, BC, CO, ID, MT, SK, UT, WY, YK CO, ID, NM, OR, SD, UT, WY AB, BC, CA, CO, ID, MT, OR, UT, WA, WY AB, BC, CA, CO, ID, MT, NM, NV, OR, SD, SK, UT, WY OR, WA CT, IL, KY, MN, PA, WI, WV CA, OR IDS CT, MB, ME, MI, MN, NS, NW, NY, OH, OK, PA, QB, VA, WI, WV CT, DE, KY, MA, MD, ME, NC, NH, PA, QB, SC, TN, VA, WV CT, ME, MI, MN, NH, NS, NW, NY, QB, VA, WI AB, AK, AZ, BC, CA, CO, ID, MT, NM, NV, OR, UT, WA, WY, YK AB, BC, CA, ID, MT, OR, WA OR AB, BC, CT, DE, KY, LB, MB, ME, MI, MN, NF, NJ, NW, NY, ON, PA, QB, SK, VA, WI, wv STARK ET AL.: NORTH AMERICAN PLECOPTERA Perlodinae Arcynopterygini Arcynopteryx compacta (McLachlan) Frisonia picticeps (Hanson) Megarcys irregularis (Banks) signata (Hagen) subtruncata Hanson watertoni (Ricker) yosemite (Needham & Claassen) Oroperla barbara Needham Perlinodes aurea (Smith) Setvena bradleyi (Smith) tibialis (Banks) wahkeena Stewart & Stanger Skwala curvata (Hanson) parallela (Frison) Diploperlini Baumannella alameda (Needham & Claassen) Cultus aestivalis (Needham & Claassen) decisus (Walker) pilatus (Frison) tostonus (Ricker) Diploperla duplicata (Banks) kanawholensis Kirchner & Kondratieff morgani Kondratieff & Voshell robusta Stark & Gaufin 393 AB, AK, BC, CO, ME, MT, NH, SK, WY BC, CA, NV, OR, WA BC, WA AK, BC, CO, ID, MT, NM, NV, UT, WY BC, ID, MT, OR, WA AB, BC, ID, MT CA, WA CA AB, CA, ID, MT, OR, WA, WY AB, BC, ID, MT BC, OR, WA OR BC, ID, MT, OR, WA, WY AZ, BC, CA, CO, ID, MB, MT, NM, NV, OR, SK, UT, WA, WY CA AZ, BC, CO, ID, MT, NM, UT, WY, YK CT, GA, IN, KY, ME, MI, NC, NY, OH, ON, PA, QB, TN, VA, WV BC, CA, ID, MT, OR, WA BC, CA, ID, MT, OR, WA, WY AL, DE, GA, MS, SC, TN, VA, WV WV VA, WV CAIN RYa OH: PA, VA, WV 394 Kogotus nonus (Needham & Claassen) modestus (Banks) Osobenus yakimae (Hoppe) Pictetiella expansa (Banks) Remenus bilobatus (Needham & Claassen) Rickera sorpta (Needham & Claassen) Perlodini Chernokrilus erratus (Claassen) misnomus (Claassen) venustus (Jewett) Diura bicaudata (Linnaeus) knowltoni (Frison) nanseni (Kempny) Helopicus bogaloosa Stark & Ray nalatus (Frison) rickeri Stark subvarians (Banks) Hydroperla crosbyi (Needham & Claassen) fugitans (Needham & Claassen) phormidia Ray & Stark Isogenoides colubrinus (Hagen) doratus (Frison) elongatus (Hagen) frontalis (Newman) GREAT BASIN NATURALIST BCACASTD Mile OR, WA, WY BC, CO, ID, MT, NM, UT, Wy BC, CA, OR, WA CO, ID, MT, UT, WY C15, ID Sy, , KY, NC, Aw AGS CesiNG VATA: , NV, OR, WA , OR NV, OR, SK, UT, WY, YK NH, QB FL, GA, LA, MS, 5c AR, IN, KS, MI, MO, OK TN GT, BL, KY ME, NC, ON, PA, QB, SC, TN, VA, WV AR, IL, IN, KS, MO, OK, TX AR, IL, IN, KS, NG EX FE, SEC AB, AK, AZ, BC, CA; CO, iD, MB, MT, NT, SK, UT, WY, YK IA, MI, PA, QB AB, AZ, BC, CO, ID, MB, MT, NM, UT, WA, WY LB, MB, ME, MI, MN, NF, NY, QB, SK, WI hansoni (Ricker) krumholzi (Ricker) olivaceus (Walker) varians (Walsh) zionensis Hanson Malirekus hastatus (Banks) Oconoperla innubila (Needham & Claassen) Yugus arinus (Frison) bulbosus (Frison) Pteronarcyidae Pteronarcyinae Pteronarcellini Pteronarcella badia (Hagen) ’ s regularis (Hagen) Pteronacrcyini Pteronarcys biloba Newman californica Newport comstocki Smith dorsata (Say) pictetii Hagen princeps Banks proteus Newman scotti Ricker Vol. 46, No. 3 CT, MA, MD, ME, NG, NS, NW, NY, PA, QB, VA, WV MB, MI, MN MI, MN, ON, QB, WI IL, IN, MI, MN, MS, SC, TN AK, AZ, CO, NM, UT GA, KY, ME, NC, NY, QB, SC, TN, VA, VI, WV NC, SC, TN GA, NC, PA, SC, TN, VA, WV GA, NC, PA, SC, TN, VA, WV AB, AK, AZ, BC, CO, ID, MT, NM, NV, OR, SK, UT, WY AB, AK, CA, NV, OR, WA Nb, Ci GA, MA, ME, NC, NH, NS, NY, PA, QB, SC, VA, WV AK, AZ, BC, CA, CO, ID, MT, NM, OR, UT, WA, WY ME, NW, NY, PA, VA, WV AL, AB, AK, BC, Filly by, IS, TOY, LA, LB, MB, ME, MN, MS, MT, NB, NY, OH, PA, QB, SC, SK, TN, VA, WI, WV, WY Ce TI INKS? 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The genus Isoperla (Plecoptera) of western North America; holomorphology and systematics, and a new stonefly genus Cascadoperla. Mem. Amer. Entomol. Soc. 32: 1-120. . 1981. Reevaluation of the genus Clioperla. Ann. En- tomol. Soc. Amer. 74: 563-569. . 1984. Descriptions of Calliperla Banks, Rickera Jew- ett, and two new western nearctic Isoperla species (Plecoptera: Perlodidae). Ann Entomol. Soc. Amer. 77: 251-263. TarTer, D. C., D. A. ADKINS, AND C. V. COVELL, JR. 1984. A checklist of the stoneflies (Plecoptera) of Kentucky. Entomol. News 95: 113-116. TarTER, D. C., AND R. F. KIRCHNER. 1980. List of stoneflies (Plecoptera) of West Virginia. Entomol. News 91: 49-53. TarTeR, D. C., M. L. Littce, R. F. KIRCHNER, W. D. WATKINS, R. G. FARMER, AND D. STEELE. 1975. Distribution of pteronarcid stoneflies in West Virginia (Insecta: Ple- coptera). Proc. West Virginia Acad. Sci. 47: 79-85. TKac, M.A., ANDB. A. Foore. 1978. Annotated list of stoneflies (Plecoptera) from Stebbins Gulch in northeastern Ohio. Great Lakes Entomol. 11: 139-142. UNZICKER, J. D., AND V. H. MCCASKILL. 1982. Plecoptera. In: Aquatic insects and oligochaetes of North and South Carolina. Midwest Aquatic Enterprises, Mahomet, Illinois. 837 pp. Wutt_, D. 1974. The distribution of stoneflies (Insecta: Ple- coptera) of the Salt River, Kentucky. Trans. Ken- tucky. Acad. Sci. 35: 17-23. Zwick, P. 1984. Notes on the genus Agnetina (Phasganophora) (Plecoptera: Perlidae). Aquatic In- sects 6: 71-79.4 THREE NEW RECORDS FOR DIATOMS FROM THE GREAT BASIN, USA Samuel R. Rushforth’, Lorin E. Squires’, and Jeffrey R. Johansen” ABSTRACT.—Three diatom species recently collected from Great Basin localities represent new records of these taxa from this region of western North America. Cocconeis scutellum Ehr. and Melosira dubia Kuetz. were collected from a thermal spring in Tooele County, Utah. Nitzschia hustedtiana Salah was collected from newly flooded marshes at the south end of the Great Salt Lake, Tooele County, Utah. We have recently studied the algal floras from several regions of western North Amer- ica. In particular, during the past few years we have examined the algae from the Great Salt Lake (Felix and Rushforth 1977, 1979, 1980, Rushforth and Felix 1982) and several thermal systems in the Great Basin Desert (Kacz- marska and Rushforth 1983a, 1983b, St. Clair and Rushforth 1977). These systems have proven to contain taxa that are unusual in comparison to other habitats in the area. For instance, Blue Lake Warm Spring contains 41 taxa that were new records for the state of Utah, 17 of which were also new records for North America (Kaczmarska and Rushforth 1984). As a part of ongoing ecological studies, we have examined the diatoms from a previously unstudied thermal system in Tooele County, Utah. We have also studied newly inundated marshlands created by the flooding of lands at the south end of the Great Salt Lake due to unusually high water during the past few years. While studying these samples, we en- countered three taxa unusual in this region that represent or confirm new records for Utah. This paper is a report and discussion of these taxa. METHODS Several periphyton samples were collected during 1983 and 1984 from a small thermal spring in Tooele County, Utah (T1S, R7W, Sec. 25), near the Genstar Dolomite Plant. These samples were collected by placing small amounts of attached algae and debris into vials. Similar samples were collected dur- ing July 1984 from ephemeral marshy pools at the south end of the Great Salt Lake, Tooele County, Utah (T2S, R4W, Sec. 4). All samples were returned to our laboratory and examined immediately. Samples were then cleared fol- lowing standard procedures using boiling ni- tric acid. Strewn mounts were prepared using Naphrax high resolution mountant, and re- sulting slides were examined using Zeiss RA microscopes equipped with bright field and Nomarski optics. RESULTS AND DISCUSSION The diatom floras of the two study sites were interesting in their floristic composition. At the Genstar locality, two of the dominant taxa were Melosira dubia Kuetz. and Coc- coneis scutellum Ehr. M. dubia represents a new record for Utah, whereas C. scutellum confirms a single questioned report by Patrick (1936). Both of these taxa are typically coastal in distribution. The third taxon reported in the present paper, Nitzschia hustedtiana Salah, was collected from the Great Salt Lake locality. This diatom is similar to specimens previously reported as Nitzschia punctata (Wm. Smith) Grun. from Utah Lake, and N. species (Patrick 1936) from the Great Salt Lake. These three taxa are described and dis- cussed below. Cocconeis scutellum Ehrenberg 1838. Figs. 1-5, 8-9. Valve elliptical, 16-21 pm wide by 20-27 tm long; rapheless valve striae radiate, 8-10 in 10 pm, proliferating to 2-3 rows of small punctae near valve margin; Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. “Department of Microbiology, Washington State University, Pullman, Washington 99164-4340. 398 RUSHFORTH ET AL.: DIATOM RECORDS July 1986 \\ \\ M : bs : ; / d ia tellum rapheless valves. 6-7, Nitzsch us SCU tellum raphe valves, 2-3, 5, Coccone Ss ss a a] o Oo Sa Sa 9 oO eI O &p =| to — rl Melos 1 X4250. > apheless valve, X1600 alve ide of v ms is scutellum SEM of r ia SEM of Coccone ira dub > 8 , Melos 11 Figs. 8— X3700. 10 by a hyaline ; raphe central area small, ar margin punctae round, 16-18 in 10 pm axial area narrow; >) oO (=| ao} oO — Q, f=} = © as a) fas She) we SaaS gE S i fae} eer ia} E50 Be at & OO Ss aos a fl — 243 HSE Sa Bs te) ne} Ke esa Sones ae ob 2s. im des 3 oO iS) on Sas a Po 2) Sis fey 9) (a July 1986 RUSHFORTH ET AL.: DIATOM RECORDS 401 Figs. 12-16. Melosira dubia: 12-14, Light micrographs of girdle views. 15, Light micrograph of valve view. 16, SEM of girdle view. All figures are X2000. circular. Our specimens are small, although within the size range of 12-40 um wide by 20-60 um long given by Hustedt (1959) for Cocconeis scutellum. The striae are finer than those often reported for this species, and the raphe valve striae are finer than the rapheless valve striae. The fine striae are undoubtedly related to the small size of the valves, a characteristic also ob- served by Mizuno (1982), who found that striae density on both valves of Cocconeis scutellum var. ornata Grun. increased with decrease in valve length. His specimens also had finer striae on raphe valves than rapheless valves. Cocconeis scutellum occurred as an epiphyte on the green alga Rhizoclonium species and on the diatom Pleurosira laevis (Ehr.) Com- pére (= Biddulphia laevis Ehr.) in water with salinity approximately 5 0/00. It is a cosmo- politan polyhalobous species that is fre- quently reported from coastal regions. 402 Melosira dubia Kuetzing, 1844. Figs. 10-16. Cells cylindrical, 13-25 pm in diame- ter by 14-30 ym in height, attached to one another by mucilage pads to form chains often more than 15 cells long; valves convex, with a flattened apex and distinctive corona, lacking collars; valve surfaces warty, with several evi- dent processes; striae generally not resolved in the light microscope. Melosira dubia is sim- ilar to Melosira nummuloides, Melosira arc- tica, and Melosira moniliformis. It differs from the former two especially in the absence of the collar on valve surfaces. It differs from Melosira moniliformis in several features, es- pecially by being smaller and having an evi- dent corona. Nitzschia hustedtiana Salah 1952. Figs. 6-7. Valves elliptical to elliptical-lanceolate, with rostrate-apiculate ends, a longitudinal fold and indistinct keel, 6-13 wm wide by 15-24 wm long; striae 16-19 in 10 pm, nearly parallel near midvalve, strongly curved radi- ate near ends, distinctly punctate; punctae 16—20 in 10 wm. Nitzschia hustedtiana is simi- lar to Nitzschia punctata (Wm. Smith) Grun., which is commonly reported from coastal ar- eas. It differs by being smaller and having finer striation. In addition, the longitudinal fold and keel may be less distinct. N. hustedtiana was discussed by Archibald (1983), who incorporated into this taxon other small Nitzschia punctata—like diatoms, in- cluding Nitzschia punctata f. minores Hustedt (1937-1938) and Nitzschia subpunc- tata Cholnoky (1960). Archibald also ex- panded the original size description of Nitzschia hustedtiana to 12.5—20.9 wm long by 5.5—-8.0 tm wide based on collections from South African rivers. The Great Salt Lake specimens expand the size range of this taxon to 24 um long and 13 wm wide, which inter- grades with the size range of Nitzschia punc- tata. However, the striae density (16-19 in 10 wm) for Nitzschia hustedtiana is distinct from the 7-10 in 10 um striae range typically re- ported for Nitzschia punctata. A diatom described as Nitzschia punctata has been reported from Utah Lake phyto- plankton (Rushforth et al. 1981, Rushforth and Squires 1985), bottom sediments (Grimes and Rushforth 1982), cores (Bolland 1974, Javakul and Rushforth 1983), and the stems of dead Phragmites plants on the shoreline GREAT BASIN NATURALIST Vol. 46, No. 3 (Grimes et al. 1980). These diatoms were 7-8 wm broad by 15-26 pm long, with 14-17 punctate striae in 10 pm and about 22 punctae in 10 pm. Utah Lake frustules differed from Great Salt Lake specimens by being linear-el- liptical in shape, resembling closely the frus- tule illustrated by Cholnoky (1960) as Nitzschia subpunctata, which is now con- specific with Nitzschia hustedtiana. Thus, these Utah Lake specimens seem to be better identified as Nitzschia hustedtiana than Nitzschia punctata. Nitzschia hustedtiana was rare in water with salinity approximately 56 0/00. LITERATURE CITED ARCHIBALD, R. E. M. 1983. The diatoms of the Sundays and Great Fish rivers in the Eastern Cape Province of South Africa. Bibliotheca Diatomolog- ica 1: 1-362, 34 plates. BOLLAND, R. F. 1974. Paleoecological interpretations of the diatom succession in Recent sediments of Utah Lake. Unpublished dissertation, University of Utah, Salt Lake City. 100 pp. CHOLNOKY, B. J. 1960. Beitrage zur Kenntnis der Di- atomeenflora von Natal. Nova Hedwigia 2: 1-128. EHRENBERG, C. G. 1838. Die Infusionstheirchen als voll- kommende Organismen. Ein Blick in das tiefere organische Leben der Natur. Leopold Voss, Leipzig. 548 pp. FELIX, E. A., AND S. R. RUSHFORTH. 1977. The algal flora of the Great Salt Lake, Utah: a preliminary report. Pages 385-392 in D. Greer, ed., Desertic termi- nal lakes. Utah Water Res. Lab. Pub. . 1979. The algal flora of the Great Salt Lake, Utah, USA. Nova Hedwigia 31 (12): 163-195. . 1980. Biology of the south arm of the Great Salt Lake, Utah. Utah Geol. Miner. Surv. Bull. 116: 305-312. Grimes, J. A., L. L. St. CLairn, AND S. R. RUSHFORTH. 1980. A comparison of epiphytic diatom assem- blages on living and dead stems of the common grass Phragmites australis. Great Basin Nat. 40 (3): 223-228. GRIMES, J. A., AND S. R. RUSHFORTH. 1982. Diatoms of Recent bottom sediments of Utah Lake, Utah, USA. Bibliotheca Phycologica 55: 1-179. Hustepr, F. 1938. Systematische und 6kologische Un- tersuchungen tber die Diatomeen-Flora von Java, Bali und Sumatra. Arch. Hydrobiol. Suppl. 15: 393-506. HustTeptT, F. 1959. Die Kieselalgen. In L. Rabenhorst, Kryptogamen-Flora von Deutschlands, Osterre- ichs und der Schweiz, v. 7, pt. 2. Reprint, Johnson Reprint Corp., New York. 845 pp. JAVAKUL, A., AND S. R. RUSHFORTH. 1983. Diatoms in sediment cores in Utah Lake, Utah, USA. Hydro- biologia 98: 159-170. KACZMARSKA I. AND S. R. RUSHFORTH. 1983a. Notes on a rare inland Hyalodiscus. Bacillaria 6: 157—164. July 1986 RUSHFORTH ET AL _____. 1983b. The diatom flora of Blue Lake, Tooele County, Utah. Biblictheca Diatomologica 2 (1): 1-123. _____. 1984. New records of diatoms from Blue Lake Warm Spring, Tooele County, Utah. Great Basin Nat. 44 (1): 120-124. KuETZING, F. T. 1844. Die Kieselschaligen Bacillarien oder Diatomeen. Nordhausen. 152 pp. Zweite Auflage, Nordhausen 1865. 152 pp. Mizuno, M. 1982. Change in striation density and systemat- ics of Cocconeis scutellum var. ornata (Bacillario- phyceae), Bot. Mag. Tokyo 95: 349-357. Patrick, R. 1936. Some diatoms of Great Salt Lake. Bull. Torrey Bot. Club 63(3): 157-166. RusHFoRTH, S. R., L. L. St. Carr, J. A. GRIMES, AND M. C. Wuitinc. 1981. Phytoplankton of Utah Lake. Great Basin Nat. Mem. 5: 85-100. .. DIATOM RECORDS 403 RUSHFORTH, S. R., AND E. A. FELIX. 1982. Biotic adjust- ments to changing salinities in the Great Salt Lake, Utah, USA. Micro. Ecol. 8: 157-161. RUSHFORTH, S. R., AND L. E. SQuireEs. 1985. New records and comprehensive list of the algal taxa of Utah Lake, Utah, USA. Great Basin Nat. 45 (2): 237-254. SALAn, M. M. 1952. Diatoms from Blakeney Point, Nor- folk. New species and new records from Great Britain. J. Roy. Microsc. Soc. Ser III, 72: 155— 169. p ST. Ciair, L. L., AND S. R. RUSHFORTH. 1977. The diatom floras of the Goshen warm spring ponds and wet meadows, Goshen, Utah, USA. Nova Hedwigia 28: 353-425. MOVEMENTS BY SMALL MAMMALS ON A RADIOACTIVE WASTE DISPOSAL AREA IN SOUTHEASTERN IDAHO Craig R. Groves' and Barry L. Keller!” ABSTRACT.—Average linear movement by populations of Dipodomys ordii, Microtus montanus, Perognathus parvus, and Peromyscus maniculatus was investigated over a 15-month period by live trapping on a low-level, radioactive waste disposal area in Idaho. No significant differences in movement among habitats were observed seasonally, excepting M. montanus in spring. Average linear movements within habitats ranged from 20 to 70 m for all species, but some patterns varied seasonally and among age classes for individual species. Although predation on contaminated small mammals from the disposal area is a vector of radionuclide transport, local movements by these rodents do not appear to be of sufficient magnitude to contribute significantly to redistribution of radioactive particles. The measurement of movements by small mammals has received considerable attention in ecological studies dealing with life history and estimation of density (Sanderson 1966). Recently several studies have focused on con- tamination of small mammals with chemical residues or radionuclides (e.g., Jefferies et all. 1973, Halford and Markham 1978), but only one investigation has considered movements by small mammals in the vicinity of a contami- nated area (Hedlund and Rogers 1980). Pre- liminary studies on a radioactive waste dis- posal area in southeastern Idaho indicated that deer mice (Peromyscus maniculatus ) tis- sues collected adjacent to the disposal area had higher concentrations of some radionu- clides than tissues from control areas (Mark- ham 1978, Markham et al. 1978). These data suggested that small mammals had access to contaminated soil areas near waste or were in direct contact with waste. Thus, small mam- mals could affect radionuclide distribution during their burrowing activities and move contaminated material in their gut, hide, or lungs. On the basis of these observations, we un- dertook a study to examine the ecology and radioecology of small mammals inhabiting the waste disposal area. One objective of this study was to determine average linear move- ments by small mammals whose activity could affect the spread and redistribution of ra- dionuclides, particularly via predation. The purpose of this paper is to report movements by small mammals on and adjacent to the disposal area and compare these movements among habitats and seasons and between sexes and age classes. Data on species diver- sity, biomass, population dynamics, and re- production of small mammals on the disposal area (Groves and Keller 1983a), as well as radiation doses and radionuclide contamina- tion to small mammals on the study area (Arthur et al. in press, 1986) have been re- ported previously. METHODS Our study was conducted at the Subsurface Disposal Area (SDA) of the Idaho National Engineering Laboratory (INEL) Radioactive Waste Management Complex. The INEL Site, a nuclear reactor testing facility under the jurisdiction of the U.S. Department of Energy, occupies 231,300 ha of sagebrush desert in southeastern Idaho. Since 1952 ap- proximately 9.9 x 10* m° of radioactive wastes have been placed in pits and trenches at the SDA, a 36 ha portion of the complex used for disposal of radioactive waste. Details on the types of waste disposed at the SDA and waste disposal practices are provided in Arthur et al. (1986). Vegetation on the SDA was dominated by seeded crested wheatgrass (Agropyron cristatum), with Russian thistle (Salsola kali) growing over more recently disturbed and ‘Department of Biological Sciences, Idaho State University, Pocatello, Idaho, 83209. Present address: Idaho Natural Heritage Program, The Nature Conservancy, 4696 Overland Rd. #576, Boise, Idaho 83705. 404 July 1986 GROVES, KELLER: IDAHO MAMMALS 405 | | awa DRAINAGE CHANNEL el \ Fig. 1. Location of Grids A-D, M, and perimeter trap lines G, H, J, and K on and adjacent to the Subsurface Disposal Area. unseeded areas. Native flora surrounding the disposal area is primarily big sagebrush (Artemisia tridentata )/bluebunch wheatgrass (Agropyron spicatum) steppe. Further details on the study area are provided in Groves and Keller (1983b). Four trapping grids (A, B, C, D), each 0.6 ha and containing 100 Longworth live traps placed at 9 m intervals in a 5 x 20 configura- tion, were established on the SDA between May and July 1978 (Fig. 1). A fifth grid (M) was established in native vegetation 150 m north of the SDA during October 1978 and con- sisted of 160 live traps in a 5 x 32 configura- tion. On the perimeter of the SDA, a dry drainage channel lies between a dike and fence that surround the disposal area. Rodent populations in this area were studied by plac- ing Longworth live traps at 9 m intervals around the entire perimeter (areas G, H, J, Our study was conducted from May 1978 to July 1979. Perimeter lines and grids were trapped weekly on a staggered basis (i.e., grids on odd weeks, lines on even weeks) from May through October 1978, and monthly thereafter. A trapping session consisted of two days of trapping per grid or line. During this time traps were baited and set in late after- noon, followed by removal and examination of animals the following morning. Captured ani- mals were eartagged with fingerling fish tags. Data obtained for each captured animal in- cluded species, trap location, weight, sex, and reproductive condition. Three measures of movement (Brant 1962) were used to estimate movements by individ- ual small mammals: (1) D, the average dis- tance between captures from one trapping period to the next, (2) M, the maximum dis- tance between captures from one trapping period to the next, and (3) S, average distance between captures within a two-day trapping session. D and M were employed to estimate how far an animal moved between trapping sessions, whereas S was used to estimate short-term movement, that is, movement from one day to the next within a trapping period. Following individual calculations, data were pooled to estimate average move- ment values for individual species. Movement data were first separated into four seasonal periods: winter (December— February), spring (March—May), summer (June-August), and autumn (September— November). These periods corresponded well with changes in temperature and snowfall, as well as changes in population density and re- 406 TABLE 1. Seasonal estimates of the average distance (m) (+S.E. (n GREAT BASIN NATURALIST Vol. 46, No. 3 = number of animals)) between successive captures from one trapping period to the next (D) and within a trapping period (S) for Peromyscus maniculatus in different habitats on and adjacent to the Subsurface Disposal Area. Blanks in the table indicate sample size was less than or equal to five animals. Habitat Statistic | Winter Spring Summer Autumn Crested Wheatgrass D Q8rA 5.2 (QU) 2525) == 1552/46) les) == So) (04D) BG se SI (G8) (A, B, D) S 9.0 + 2.8 (6) 16.2 + 4.7 (35) 17.8 + 4.0 (94) 33.7 + 12.0 (73) Russian Thistle D — 43.2 + 18.1 (8) 45.1 + 16.3 (20) All se BD C4) (C) S — — 16.4 + 2.0 (15) 16.9 + 4.1 (26) Sagebrush D —- — 43.8 +_ 7.0 (8) _ (M) S = -- 19.6+ 4.1 (6) _- Fence line D 211 = 479 (50) 34:7 = 4.7 (149)! 36758725 Gal (195) OANA (all) (G, H, J, K) S AT Al ==) 3°3(83)) 2652 42 75:3) (122) 2036) 255 229165) nl An Sena Clo?) TABLE 2. Seasonal estimates of the average distance (m) (+S.E. (n number of animals)) between successive captures from one trapping period to the next (D) and within a trapping peroid (S) for Dipodomys ordii and Microtus montanus in different habitats on the Subsurface Disposal Ar than five animals. rea. The blank in the table indicates a sample size was less Dipodomys ordii Habitat Statistic Summer Autumn Crested Wheatgrass D 55.3 + 25.5 (29) 37.5 + 15.0 (30) (A, B, D) S 18.6 + 3.8 (15) 9273)== Ss 610N(25) Fence line D 51.2 + 23.6 (38) 29.1+= 5.1 (90) (G, H, J, K) S 33.2 + 15.5 (30) ss 21.3 + 5.9 (74) Microtus montanus Habitat Statistic Spring Summer Crested Wheatgrass D 21.0 + 3.1 (42) 68.4 + 22.3 (16) (A, B, D) S 14.5 + 2.9 (20) S84! se INL (@) Fence line D 54.0 + 16.0 (75) 77.2 + 28.6 (13) - (G, H, J, K) S 21.2 + 7.2 (37) = production of small mammals on the study area (Groves and Keller 1983a). Next, move- ment data were tested for differences among habitats within a season. Lastly, data were pooled among habitats to test for differences in movement between sexes and between age classes within a season. Within any season, an animal was recap- tured in no more than six trapping periods; the majority of animals were recaptured four times. Only animals that remained ona grid or trapline during their recapture history were included in the movement analysis, except those animals that moved the short distance between grids and adjacent traplines (see Fig. 1). Movements by small mammals from one trapping line or grid to another within the SDA covered distances of more than 200 m, a length well beyond movements previously re- ported for any species on our study area (Brant 1962, Ramsey 1969). We considered such movements to be outside the normal home range of an animal and classified such individ- uals as dispersers. Less than 1% of the small mammals marked on an individual grid or trapline dispersed to another grid or trapline within the SDA. These animals, as well as those small mammals that dispersed off the SDA, will be reported on elsewhere. Skewness and kurtosis values indicated that the movement data were not normally dis- tributed. Additionally, the assumption of ho- moscedasticity among groups of movement data within each species was violated. Thus, nonparametric procedures, available as SPSS programs (Nie et al. 1975, Hull and Nie 1979), were employed with a = 0.05. RESULTS A total of 20,689 live-trap nights produced 9,318 captures of 10 species of small mammals July 1986 during the 15-month study (Groves and Keller 1983a). Sufficient data were obtained to estimate average movements for three spe- cies: (1) deer mice (Peromyscus maniculatus ), montane voles (Microtus montanus), and (3) Ord’s kangaroo rats (Dipodomys ordii). Addi- tionally, limited movement data were ob- tained for Great Basin pocket mice (Per- ognathus parvus ). Kruskal-Wallis (K-W) analyses of variance or Mann-Whitney U tests were used to ana- lyze differences in movement among habitats by P. maniculatus in each season (Table 1.). No significant differences in movement (D, S, or M) among habitats were detected for P. maniculatus’. Subsequently, data from differ- ent habitats were pooled to produce a single estimate of movement in each season for D, S, and M. For D. ordii and M. montanus, movement data by habitat were only estimated for two seasons because of insufficient sample sizes in other seasons. Mann-Whitney U tests were used to analyze differences in movement be- tween crested wheatgrass and fenceline habi- tats for both D. ordii and M. montanus (Table 2). No significant differences in movement between fenceline and crested wheatgrass habitats were detected for D. ordii in either summer or autumn by any movement statistic (D, S, or M). In spring, M. montanus moved significantly (P < .05) greater distances in fenceline habitat than crested wheatgrass habitat as indicated by D and M. No signifi- cant differences in movement between habi- tats were found for M. montanus in summer. Subsequently, data from different habitats were pooled for both D. ordii and M. mon- tanus to produce a single estimate of move- ment in each season for D, S, and M. A K-W analysis of variance followed by mul- tiple range tests indicated that P. maniculatus moved longer distances in spring and summer than other seasons, as estimated by D.or M (P = .05, Fig. 2a). Seasonal estimates of D ranged from approximately 22 m in autumn and winter to 32 m in spring and summer. Estimates of D and M were significantly dif- ferent (P S .05) among spring, summer, and autumn periods for D. ordii (Fig. 2b). There 3Because estimates, errors, and sample sizes for M were similar to D for all species, these data were excluded from the text but are available from the authors. GROVES, KELLER: IDAHO MAMMALS 407 60; 9) B maniculatus € lod Zz uJ = uJ > ro) c) M. montanus es = 20 100 (29) WINTER SPRING SUMMER AUTUMN Fig. 2. Seasonal estimates of D and S for P. manicula- tus (a), D. ordii (b), and M. montanus (c) on and adjacent to the Subsurface Disposal Area. Closed circies = D; open circles = $; bars = S.E.; sample sizes are in paren- theses. was a trend of decreasing range of movement from spring to autumn, with D ranging from approximately 72 to 35 m in these periods. Because only a few D. ordii were captured in winter (Groves and Keller 1983a), movements could not be estimated for this season. Both D and M were significantly greater (P = .01) in winter and summer compared to spring and autumn for M. montanus (Fig. 2c). Estimates of D ranged from approximately 70 m in win- ter and summer to 37 m in spring and autumn. 408 GREAT BASIN NATURALIST Vol. 46, No. 3 TABLE 3. Estimates of movement (m)(+ S. E. (n = number of animals) for male and female Microtus montanus and Dipodomys ordii on and adjacent to the Subsurface Disposal Area. All pairs of male and female movements shown in the table were significantly different (P < .05). Species Season Statistic M. montanus Spring D Autumn S D. ordii Spring D Summer D Short-term movements (S) were not signifi- cantly different among seasons for any spe- cies, but they were generally less than D for all species. A series of Mann-Whitney U tests was used to detect differences in movement (D and S) between males and females within each sea- son. No significant differences in movement were found between male and female P. maniculatus in any season. In spring male M. montanus and male D. ordii moved greater distances than females, as indicated by D (Table 3). In fall male M. montanus moved greater distances within trapping sessions (S ) than females (41 m vs. 6 m), although sample sizes were small for this comparison. Lastly, female D. ordii moved greater distances than males in summer. Mann-Whitney U tests were also used to detect differences in movement (D and S) between age classes for P. maniculatus and M. montanus. The mean weight at sexual matu- rity was used to separate juveniles from adults (Groves 1981). No distinctions were made be- tween juvenile and adult D. ordii because of difficulties in assessing external sexual charac- teristics. No significant differences in move- ment between age classes of P. maniculatus were observed in any season. For M. mon- tanus, juveniles moved significantly greater (P S .05) distances between trapping sessions (D) than adults in spring (55.2 + 20.2 m (n = 10) vs. 15.6 + 2.5 m (n = 36)). Additionally, juvenile M. montanus moved greater (P =< .05) distances within a trapping session (S) than adults (53.8+ 29.6 m (n = 4) vs. 14.8 + 9.9 m (n = 8)), although sample sizes were small for comparative purposes. Although sufficient sample sizes were not available to estimate seasonal movements by P. parvus, D and M were estimated by pool- ing data from spring and summer. The major- ity of these values were determined from re- capture records on Grids A and B in crested Male Female 36.6 = 6.0 (40) 26.6 + 10.2 (56) 41.4 + 16.7 (8) 6.0+ 2.1 (11) 44.2+ 6.0 (31) 21.5+ 4.2 (20) 37.8 + 5.0 (36) 43.2 + 29.4 (35) wheatgrass stands and Grid M in sagebrush habitat. Average distance between successive captures (D) + S.E. for P. parvus was 45.0 + 15.8 m (n = 14); maximum distance between successive captures (M) was 60.8 + 26.3 m (n — 14) DISCUSSION Local movements by small mammals have received considerable attention in the ecolog- ical literature. Most studies published to date have concentrated on home ranges (expressed in areal terms) of individual species as deter- mined by live trapping on-grids for short peri- ods of time. Because our primary objective was to determine the distance that a small mammal could transport contaminated mate- rial, we abandoned the concept of home range in favor of data on the magnitude of average linear movements by small mammals occupy- ing the Subsurface Disposal Area. On the SDA, P. maniculatus showed signif- icantly reduced movements in autumn and winter compared to spring and summer. Colder temperatures and a concomitant re- duction in activity may have been responsible for the lesser movements in these seasons. Stebbins (1971) has documented periods of torpor for P. maniculatus in Canada during periods of snow and cold. Similarly, D. ordii moved significantly lesser distances in au- tumn compared to spring and summer. This observation, coupled with the fact that we captured few D. ordii in winter (Groves and Keller 1983a), suggests that cold weather may also affect activity in this species. O Farrell (1974) previously reported that D. ordii may enter periods of torpor during cold weather. No significant differences were found be- tween movements of male and female P. maniculatus on the SDA, although adults did move greater lengths than juveniles in sum- mer. Both of these results are in agreement : tl i i if July 1986 with the findings of Brant (1962). Stickel (1968) noted that immature Peromyscus re- main near the natal site until the dispersal period that coincides with sexual maturity. Consequently juvenile P. maniculatus on the SDA could be expected to show reduced lin- ear movements when compared to adults. In spring juvenile M. montanus moved signifi- cantly greater distances than adults. These longer movements may have resulted from juveniles dispersing from an increasing M. montanus population (Groves and Keller 1983a), a phenomenon reported by several authors for microtine populations (Myers and Krebs 1971). No data on movements have been pub- lished for P. parvus. Our data indicated that this species moved approximately 45 m be- tween successive captures (D) in crested wheatgrass habitat during spring and sum- mer. Thus, P. parvus exhibited linear move- ments slightly less than D. ordii, a larger rodent in the same family (Heteromyidae). In addition to the grids and traplines lo- cated on the SDA for assessing rodent popula- tions there, dispersal from the area was esti- mated with subsampling systems used _ to enumerate the fraction of the populations that permanently leave the SDA. Although a vari- ety of factors affect the degree of accuracy of such estimates (Keller 1978), our data suggest that only 22% of the small mammals occupy- ing the SDA dispersed on an annual basis. Thus, the majority of movements by small mammals occupying the disposal area were found to occur within its boundaries. An obvi- ous corollary is that the majority of contami- nated small mammals also remain within the SDA during their movements. Data from the radioecology aspects of our study indicated that some P. maniculatus and __D. ordii on the SDA received radiation doses significantly higher than animals from control areas (Arthur et al. 1986). In addition, concen- trations of several radionuclides in P. manicu- latus tissues from the SDA were significantly higher than those from control areas (Arthur et al. in press a). Coyote fecal samples col- lected adjacent to the SDA boundary con- tained elevated concentrations of one ra- dionuclide, presumably from uptake of contaminated small mammals (Arthur and Mark- ham 1982). Because average linear movements GROVES, KELLER: IDAHO MAMMALS 409 by small mammals on the SDA range from 20 to 70 m, it is likely that most of the primary redistri- bution of contaminated material by small mam- mals via predation occurs within this range from the point of contamination on the SDA. This type of information should be helpful to waste management personnel in implementing a biotic monitoring plan. The environmental consequences of radia- tion doses and radionuclide uptake by small mammals on the SDA are likely minimal be- cause the overall amount of radioactivity transported by small mammals off the SDA is small (Arthur et al. in press, 1986) and no adverse impacts to small mammals on the SDA have been observed. Beyond the practi- cal application of these movement data, this study has also provided new information on linear movements by small mammals in crested wheatgrass, Russian thistle, and sage- brush habitats, all common in the Great Basin. Prior to this study, no information was available on movements by P. parvus; data on linear movements by P. maniculatus, D. or- dii, and M. montanus were not previously reported for any of the above habitats. There- fore, our movement data contribute new in- formation to the natural history of these four small mammal occupants of the Great Basin. ACKNOWLEDGMENTS We thank C. Levesque for field assistance and C. Nimz for extensive aid in computer programming. O. D. Markham and J. E. An- derson critically reviewed the manuscript. This research is a contribution from the INEL Site Ecological Studies Program, supported by the Office of Health and Environmental Research and the Nuclear Fuel Cycle and Waste Management Division, U. S. Depart- ment of Energy. B. L. Keller also received support from the Faculty Research Commit- tee (Grant 412), Idaho State University. LITERATURE CITED ARTHUR, W. J., AND O. D. MarKHAM. 1982. Radionuclide export and elimination by coyotes at two radioac- tive waste disposal areas in southeastern Idaho. Health Physics 43: 493-500. ARTHUR, W. J., O. D. MarKHAM, C. R. GROVES, AND B. L. KELLER. In press. Radionuclide export by deer mice at a solid radioactive waste disposal area in southeastern Idaho. Health Physics. 410 ARTHUR, W. J., O. D. MarkHam, C. R. Groves, B. L. KELLER, AND D. K. HALFoRD. 1986. Radiation dose to small mammals inhabiting a solid radioactive waste disposal area. J. Appl. Ecol. 23: 13-26. BRANT, D. H. 1962. Measures of the movements and population densities of small rodents. Univ. of California Publ. in Zool. 62: 105-184. GROVES, C. R. 1981. The ecology of small mammals on the Subsurface Disposal Area, Idaho National Engi- neering Laboratory Site. Unpublished thesis, Idaho State University, Pocatello. 87 pp. GROVES, C. RB., AND B. L. KELLER. 1983a. Ecological char- acteristics of small mammals on a radioactive waste disposal area in southeastern Idaho. Amer. Midl. Nat. 109: 253-263. _____. 1983b. Population ecology of small mammals on the Radioactive Waste Management Complex, Idaho National Engineering Laboratory. Pages 21-46 in O. D. Markham, ed., Idaho National Engineering Laboratory Radioecology and Ecol- ogy Programs 1983 Progress Report. DOE/ID- 12098. National Technical Information Service, Springfield, Virginia. 434 pp. HALForD, D. K., AND O. D. MarkHAM. 1978. Radiation dosimetry of small mammals inhabiting a liquid radioactive waste disposal area. Ecology 59: 1047-1054. HEDLUND, J. D., AND L. E. Rocers. 1980. Great Basin pocket mice (Perognathus parvus) in the vicinity of radioactive waste management areas. North- west Sci. 54: 153-159. HULL, C. H., ANDN. H. Nig. 1979. Statistical packages for the social science update—new procedures and facilities for releases 7 and 8. McGraw Hill, New York. JEFFERIES, D. J., B. STAINSBY, AND M. C. FRENCH. 1973. The ecology of small mammals in arable fields drilled with winter wheat and the increase in their dieldrin and mercury residues. J. Zool. 171: 513-539. GREAT BASIN NATURALIST Vol. 46, No. 3 KELLER, B. L. 1978. Dispersal and density of small mam- mals on the Radioactive Waste Management Com- plex, Idaho National Engineering Laboratory Site. Pages 67-73 in O. D. Markham, ed., Ecolog- ical Studies on the Idaho National Engineering Laboratory Site 1978 Progress Report. IDO- 12087. National Technical Information Service, Springfield, Virginia. 371 pp. MarkHaM, O. D. 1978. Activation and fission radionu- clides in the environment near the Idaho National Engineering Laboratory Radioactive Waste Man- agement Complex. IDO-12085. National Techni- cal Information Service, Springfield, Virginia. 19 pp. MARKHAM, O. D., K. W. PUPHAL, AND T. D. FILER. 1978. Plutonium and americum contamination near a transuranic storage area in southeastern Idaho. J. Env. Qual. 7: 422-428. Myers, J. H., AND C. J. Kress. 1971. Genetic, behavioral, and reproductive attributes of dispersing field voles, Microtus pennsylvanicus and Microtus ochrogaster. Ecol. Monogr. 41: 53-78. Nik, N. H., C. H. HULL, J. G. JENKINS, K. STEINBRENNER, AND D. H. BENT. 1975. Statistical packages for the social sciences. 2d ed. McGraw Hill, New York. O'FaArRRELL, M. J. 1974. Seasonal activity patterns of ro- dents in a sagebrush community. J. Mammal. 55: 809-823. RaMSEY, P. R. 1969. Analysis of movement patterns in a population of Dipodomys ordii. Unpublished the- sis. Texas Tech. College, Lubbock. 25 pp. SANDERSON, G. C. 1966. The study of mammal move- ments—a review. J. Wildl. Mgmt. 30: 215-235. STEBBINS, L. L. 1971. Seasonal variations in circadian rhythms of deer mice in northwestern Canada. Arctic 24: 124-131. STICKEL, L. F. 1968. Home range and travels. Pages 373-411 in J. A. King, ed., Biology of Peromyscus (Rodentia). Special Publ. No. 2, Amer. Soc. of Mammal., Lawrence, Kansas. 593 pp. ROLE OF THREE RODENTS IN FOREST NITROGEN FIXATION IN WESTERN OREGON: ANOTHER ASPECT OF MAMMAL-—MYCORRHIZAL FUNGUS—-TREE MUTUALISM -l : 2 ees C. Y. Li’, Chris Maser”, Zane Maser’, and Bruce A. Caldwell! ABSTRACT. —To determine the role of the California red-backed vole (Clethrionomys californicus), the northern flying squirrel (Glaucomys sabrinus ), and the deer mouse (Peromyscus maniculatus) in the nitrogen cycle of forest stands in western Oregon, bacterial colonies were isolated and purified from feces, and their nitrogen-fixing ability measured by acetylene-reduction assay. The ability of the bacterial species Azospirillum sp. to withstand freezing was also tested. Fecal extracts were used to test whether fecal pellets can provide the nutrients necessary for growth of the bacteria. All the feces tested contained viable nitrogen-fixing bacteria, and both species can survive drying and one can survive freezing. Azospirillum colonies grew well on liquid medium but required yeast extract for growth and nitrogenase activity. Fecal extracts from flying squirrels and chickarees (Tamiasciurus douglasi) were as effective as the yeast. The results suggest another link in the chain of mutualism that unites small mammals, mycorrhizal fungi, and forest trees. Some small forest-dwelling rodents help maintain productivity of forested ecosystems by disseminating viable spores of hypogeous, mycorrhizal fungi (Kotter and Farentinos 1984, Maser et al. 1978, McIntire 1984, Roth- well and Holt 1978, Trappe and Maser 1976). Nitrogen-fixing bacteria have recently been found in fungal sporocarps eaten by small mammals (Hunt and Maser 1985, Li and Castellano 1985, Maser et al. 1978). To deter- mine whether small mammals play a role in the nitrogen balance of the forest by eating hypogeous fungal sporocarps and thereby dis- persing the bacteria, we sought answers to these questions: e Do feces of forest-dwelling rodents contain nitrogen- fixing bacteria? e Are these nitrogen-fixing bacteria viable after passage through a rodent’s intestinal tract? e Can these bacteria survive freezing, drying, and high temperature? e Do the fecal pellets provide the nutrients necessary for growth of nitrogen-fixing bacteria? We studied three common and widely dis- tributed forest rodents: the deer mouse (Per- omyscus maniculatus), the California red- backed vole (Clethrionomys californicus ), and the northern flying squirrel (Glaucomys sabrinus ). The deer mouse, ubiquitous throughout most of North America, feeds on fruits, seeds (including conifer seeds), and hypogeous, my- corrhizal fungi (Hunt and Maser 1985, Maser et al. 1978, 1981). The red-backed vole ranges south of the Columbia River, throughout the forested areas of western Oregon into north- western California, and from the Pacific Coast to the crest of the eastern Cascade Range (Maser et al. 1981). It eats mostly hypogeous fungal sporocarps (Maser et al. 1978, Ure and Maser 1982). The flying squirrel, a noctur- nally active inhabitant of most coniferous forests throughout temperate North America, also eats mostly fruiting bodies of hypogeous, mycorrhizal fungi (Maser et al. Food habits, 1985; Maser et al. Northern flying squirrel, 1985). We recognize that more quantitative data may be desired than is found in this paper. Our data are the first reported for the follow- ing interactions, however, and we are only now determining what quantitative questions can and need to be asked. Further, we have not found a way or person to identify the unknown bacteria and yeasts that we encoun- ter. U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station, Forestry Sciences Laboratory, 3200 Jefferson Way, Corvallis, ! Oregon 97331. 2U_S. Department of the Interior, Bureau of Land Management, Forestry Sciences Laboratory, 3200 Jefferson Way, Corvallis, Oregon 97331. 3Department of Forest Science, Oregon State University, Corvallis, Oregon 97331. ‘Department of Microbiology, Oregon State University, Corvallis, Oregon 97331. 411 412 GREAT BASIN NATURALIST Vol. 46, No. 3 TABLE 1. Acetylene reduction by bacteria isolated from small mammal feces. Bacterium isolated Source Azospirillum sp. Azospirillum sp. Clostridium butyricum ‘average of 5 replicates “Average of 3 replicates METHODS To determine if nitrogen-fixing bacteria survive passage through rodent digestive tracts, techniques for fecal collection and lab- oratory techniques were developed to isolate and purify bacterial colonies from rodent feces (Li and Maser, 1986). Acetylene-ethylene as- say for nitrogenase activity as an indication of nitrogen-fixing ability was as described by Hardy et al. (1968). Yeast populations in feces were determined by dilution-plating on sodium albumenate agar (Waksman and Fred 1922), To test the ability of Azospirillum sp. to withstand freezing, flying squirrel feces were held at —17 C (1 F) for three months. The feces were thawed and the bacteria isolated, grown, and tested for nitrogenase activity (Li and Maser, in press; additional details on file, Forestry Sciences Laboratory, Corvallis, Ore- gon 97331). Azospirillum sp. isolated from feces of the flying squirrel were used to test whether fecal pellets provide the nutrients necessary for growth of nitrogen-fixing bacteria. We used fecal extracts from both the flying squirrel and the chickaree (Tamiasciurus douglasi), a diur- nal tree squirrel that shares the flying squir- rel's habitat in the Pacific Northwest. Fecalex- tracts were prepared by homogenizing the fecal pellets in deionized water (2:100 w/v) ina Polytron with a saw-tooth generator, 165 x 12 mm, at maximum speed for | min. Debris was removed by centrifugation for 30 min at 14,500 x g. The supernatant was filter-steri- lized, and 0.2 ml was added to 20 ml of Dobereiner’s nitrogen-free liquid medium (Débereiner and Day 1976). RESULTS AND DISCUSSION Feces of all the small mammals we tested contained viable nitrogen-fixing bacteria. California red-backed vole Northern flying squirrel Deer mouse Acetylene reduction (nmole ethylene/ mg protein per hr) 167.0! 88.67 344.0! Azospirillum sp.—a microaerophilic, nitro- gen-fixing bacterium (Lakshmi et al. 1977)— was isolated from feces of one red-backed vole and five flying squirrels. Clostridium bu- tyricum—an anaerobic, nitrogen-fixing bac- terium (Buchanan and Gibbons 1974)—was isolated from feces of seven deer mice. The bacteria not only survived passage through the digestive tracts but also grew and reduced acetylene in vitro (Table 1). Most bacteria do not survive freezing with- out desiccation because they rupture on thaw- ing; thus, bacteria to be stored are usually freeze-dried (Gherna 1981). Azospirillum sp. can survive freezing in the feces, however, and C. butyricum forms an endospore stage that should also be able to survive freezing. Azospirillum sp. survived at least 15 years in air-dried soil at 28 C + 2 C (82 + 4 F) (Lakshmi et al. 1977). In our study C. bu- tyricum survived 3 months in air-dried feces, presumably in the endospore form, and it re- tained the capacity to reduce acetylene. Many workers have used yeast extract or yeast extract combined with vitamins to pro- mote nitrogenase activity of acetylene-reduc- ing bacteria (Barber and Evans 1976, Haahtela et al. 1981, Murray and Zinder 1984, Rennie 1981, Tyler et al. 1979). The Azospirillum colonies grew well on nutrient agar or trypticase soy agar, and they reduced acetylene when grown under conditions of 99% nitrogen and 1% oxygen. The bacterium required yeast extract for growth and nitroge- nase activity (Table 2). Controls without acetylene were also assayed with negative re- sults. Extracts from the feces of flying squir- rels and chickarees were as effective as yeast extract for inducing nitrogenase activity. Ad- — dition of vitamins into the fecal extract proved unnecessary for growth and nitrogenase activ- ity of Azospirillum sp. (Table 2). Yeast extract, a component of many stan- dard culture media (Tuladhar and Rao 1985), July 1986 TABLE 2. Influence of additives on acetylene reduction by Azospirillum sp. in Débereiner’s nitrogen-free liquid medium (Débereiner and Day 1976). Acetylene reduction” (nmoles ethylene/mg Growth condition protein per hr) Medium with: Yeast extract 71.4 Yeast extract and vitamins 88.6 Vitamins 0 Flying squirrel fecal extract 75.2 Chickaree fecal extract 109.7 Medium without: Yeast extract and vitamins 0 V Average of 3 replicates each is also necessary to the nitrogen-fixing bacteria in vitro (Table 2). We found (in three replica- tions) that fecal pellets of deer mice contained yeast populations that ranged from 33,000 to 40,000 propagules per fecal pellet. A pure cul- ture of yeast and a purified culture of Azospiril- lum sp., both isolated from feces of the flying squirrel, were placed in a nitrogen-free medium and incubated for five days at 30 C (86 F). Results (two replicates each) indicated that the yeast propagules promoted growth and nitrogenase activity of the bacterium. Azospirillum sp. alone or yeast propagules alone in Débereiner’s liquid medium exhibited no nitrogenase activity, but _ Azospirillum sp. and yeast propagules together in Dobereiners liquid medium formed 43 nmoles from acetylene per sample per 17 hr. Viable nitrogen-fixing bacteria, yeast, and spores of hypogeous, mycorrhizal fungi all survived passage through the digestive tracts | of rodents. (Viability of the mycorrhizal fun- | gus spores was tested in studies with seedlings and will be published elsewhere.) The fecal pellets contained the complete nutrients for the nitrogen-fixing bacteria. These findings have several implications for forest habitats. \ Inoculation of soil with organisms carried in _ rodent feces is probably common in forest ecosystems. For example, Azospirillum sp. can penetrate plant roots (Lakshmi et al. 1977, » Patriquin and Débereiner 1978) and is able to _ survive for 15 years in stored, air-dried soil (Lakshmi et al. 1977). The fossorial red- | backed voles and arboreal flying squirrels are _ obligate forest-dwellers. When they dig at the bases of trees, the organisms in their feces can inoculate rootlets with nitrogen-fixing bacteria, yeast, and spores of mycorrhizal fungi. LIET AL.: RODENT-FUNGUS-TREE MUTUALISM 413 The deer mouse is one of the first small mam- mals to occupy clearings after logging or fire, so it could inoculate the soil, even soil that has been severely altered by a hot fire. Although the spores of mycorrhizal fungi may not survive high surface temperatures in openings, they could survive under large woody debris on the soil surface where deer mice are active or below the soil surface in rodent burrows. Unlike the fungal spores, the nitrogen-fixing bacterium C. bu- tyricum has a built-in survival mechanism (the endospore) by which it can withstand tempera- tures up to 80 C (176 F) (Simbert and Krieg 1981). Small rodents have often been seen as detri- mental to timber management (Campbell 1982, Crouch and Radwan 1975, Hooven 1975, Sulli- van 1979, 1980), and poisons and habitat manip- ulation have been used against them. But the more forests are altered by human actions, the more evident becomes the need to understand the interactions of all the organisms in the ecosystem. How each component functions is often far more complex than might be antici- pated, and the role it plays may be essential in maintaining ecosystem health. ACKNOWLEDGMENTS We thank J. W. Witt for helping obtain fresh flying squirrel droppings; D. K. Grayson, R. Molina, R. F. Tarrant, and P. D. Weigl for reviewing the paper; G. Bissell for typing the various drafts. This paper repre- sents a partial contribution (No. 15) of the project entitled “The Fallen Tree—An Exten- sion of the Live Tree.” Cooperating on the project are the U.S. Department of the Inte- rior, Bureau of Land Management; U. S. 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Isolation of Azospirillum from diverse geographic regions. Can. J. Micro- biol. 25: 693-697. Ure, D. C., AND C. Maser. 1982. Mycophagy of red- backed voles in Oregon and Washington. Can. J. Zool. 60: 3307-3315. WAKSMAN, S. A., AND E. B. FRED. 1922. A tentative outline of the plate method for determining the number of | microorganisms in the soil. Soil Sci. 14: 27-28. CANIDS FROM THE LATE PLEISTOCENE OF UTAH Michael E. Nelson’ and James H. Madsen, Jr.” ABSTRACT. —Two canids, Vulpes vulpes and Canis lupus , are recorded from shoreline deposits of Lake Bonneville in north central Utah. Both species are new records in the Pleistocene sediments of Utah and add to our scarce knowledge of the large carnivores that inhabited the shoreline environments of Lake Bonneville. The sand and gravel quarries in the shore- line deposits of Lake Bonneville historically have been the most important sources for late Pleistocene vertebrate fossils in Utah. The Lake Bonneville shoreline mammalian fauna has been partially reviewed by Nelson and Madsen (1978, 1980, 1983), Stokes and Condie (1961), and Stock and Stokes (1969). Miller (1976) described the Silver Creek local fauna from the mountains 20 miles east of Salt Lake City, whereas Heaton (1985) docu- mented the late Pleistocene to Recent Crystal Ball Cave local fauna from westernmost Utah. We report the addition of two canids, the fox, Vulpes vulpes, and a wolf, Canis lupus, to the Bonneville fauna. STRATIGRAPHY AND LOCATION The wolf specimens (UVP 100; UVP 101) and one of the fox dentaries (UVP 082) were collected from what was locally known as the Hardman Gravel Quarries (NW 1/4 and NE 1/4, Sec. 32, TIN, RIE, Salt Lake County) in northern Salt Lake City, an area now largely obscured by large homes and the Ensign Ele- mentary School. Both specimens were col- lected by Golden York, longtime curator of geology in the University of Utah, Depart- ment of Geology Museum. The second fox specimen was collected in 1935 near Bacchus, Utah, southwest of Salt Lake City (locality _ number, 42SL126V in Sec. 8, T25S, R2W), by Mr. A. V. Jenkins. This latter specimen was . found in association with several musk ox _ (Symbos cavifrons) vertebrae. The Hardman Gravel Pit (locality number, 42SLOOIV), at an elevation of 4,800—5, 000 ft, yielded sand and gravel from the shoreline deposits near the maximum level of Lake Bon- neville. Nelson and Madsen (1980), in follow- ing Morrison (1965), thought that the Hard- man Quarry was in the Alpine Formation and deposited during the time interval of 33,000—68,000 years BP. However, Scott et al. (1983) have shown that the Alpine is not a valid formational designation. Currey et al. (1983) have placed these quarries at the Bon- neville level of the Bonneville Lake Cycle, with deposition occurring around 14,500- 18,000 years BP. All specimens have been curated and en- tered into the Paleontology Collections of the Antiquities Section, Utah Division of State History (UVP). The specimens from the re- cent mammal collection of the University of Utah are identified by the initials UM. SYSTEMATIC PALEONTOLOGY Class Mammalia Order Carnivora Family Canidae Canis lupus Linnaeus, 1758 Gray Wolf MATERIAL.—UVP 101, right M'-M” with fragments of palatine and maxilla (Fig. 1); UVP 100, left P* with alveoli for P®, M’ and interior roots of M’ (Fig. 2). These specimens are most likely from the same individual. Discussion.—Nowak (1979) recognized four species of wolves from the late Pleis- tocene of North America: (1) Canis armbrus- teri, an early? Irvingtonian to early Ran- cholabrean form; (2) C. dirus, the Ran- cholabrean to early Recent dire wolf; (3) C. ‘Department of Earth Sciences and Sternberg Memorial Museum, Fort Hays State University, Hays, Kansas 67601-4099. ? Antiquities Section, Utah Division of State History, 300 Rio Grande, Salt Lake City, Utah 84101. 415 416 GREAT BASIN NATURALIST Vol. 46, No. 3 Fig. 1. Canis lupus: UVP 101, occlusal view of right M'—M_°. Solid bar represents | cm. Fig. 2. Canis lupus: UVP 100, lateral view of left P*. Solid bar represents 1 cm. TABLE 1. Tooth dimensions in samples of Canis lupus, C. dirus, and the Bonneville specimen. The measurement is the maximum anteroposterior length of the crown of P* measured on the outer side. Measurements of C. lupus and C. dirus , with the exception of the Utah specimen, are from Nowak (1979). Sample N OR xX C. lupus, male, Recent 233 22.2-30.5 25.92 C. lupus, female, Recent 146 92,.9-98.2- 94.79 C. lupus, Pleistocene, Maricopa 6 23.3-28.5 26.53 C. lupus, Pleistocene, Rancho La Brea 1l z 23.0—29.2 26.19 C. dirus, Pleistocene, Maricopa 4 31.0-32.5 31.90 C. lupus, Recent, Utah 1 — 12.31 UVP 100, Pleistocene, Utah 1 — 29.60 rufus, the early Irvingtonian to Recent, but poorly known, red wolf; and (4) the late Irv- ingtonian to Recent gray wolf, C. lupus. The largest collections of late Pleistocene wolves have come from the southern California tar pits—McKitterick, Maricopa, and especially Rancho La Brea, where a minimum of 1,646 dire wolves, but less than 15 gray wolves, have been documented. Most other collec- tions in the United States represent single, or small, samples from cave deposits. Nowak (1979) believed that C. lupus evolved in Asia, whereas C. dirus arose in North America. However, C. lupus ulti- mately “prevailed over the dire wolf, either through competition or because of external factors, and established itself as the major large predator of North America.” Terminal extinction dates for the dire wolf are around 9500 BP. (Kurten and Anderson 1980), whereas the gray wolf is still extant in North America. However, modern man has extir- pated most of these predators from their origi- nal range in the contiguous United States, and buta single large population remains in north- ern Minnesota (Nowak 1979). Dire wolves were generally larger than gray wolves, having a stockier build and relatively shorter limbs. Most wolves are specifically identified on the basis of their dentition, even though there is some size overlap in these dimensions (Tables 1 and 2). The Bonneville specimen falls into the general size range of a large gray wolf or a small dire wolf. Both C. lupus and C. dirus have similar upper fourth premolars that lack a prominent deuterocone and lingual cingulum (Fig. 2). However, the length of P* on C. dirus is gen- erally much longer than on C. lupus (Table 1). Both species have rather undiagnostic, small, second upper molars. In the first upper molar both species have a large paracone and metacone. However, C. lupus generally has a large hypocone with a complete anterolingual cingulum that joins the hypocone. In C. dirus the hypocone is July 1986 NELSON, MADSEN: PLEISTOCENE CANIDS 417 TABLE 2. Tooth dimensions in samples of Canis lupus, C. dirus, and the Bonneville specimen. The measurement is 9 9 2 < the maximum transverse diameter of M’ from the outermost point to the innermost point of the crown. Measurements of C. lupus and C. dirus, with the exception of the Utah specimen, are from Nowak (1979). Sample . lupus, male, Recent . lupus, female, Recent . lupus, Pleistocene, Maricopa . lupus, Pleistocene, Rancho La Brea . dirus , Pleistocene, Maricopa . dirus, Pleistocene, Rancho La Brea . lupus, Recent, Utah VP 100, Pleistocene, Utah SaQqQqQaaq a? N OR xX 233 11.4-16.7 13.82 146 11.2-16.3 13.44 4 12.2—13.7 12.87 10 12.5-14.3 13.44 4 14.4-16.0 15.02 62 13.1-17.0 15.15 1 — 12.31 13.53 1 aaa Fig. 3. Vulpes vulpes: UVP 82, lateral view of left dentary with P;—M). Solid bar represents | cm. generally reduced, and the incomplete anterolingual cingulum does not reach the hypocone but usually ends somewhere near the protocone. The Bonneville specimen morphologically agrees with the traits as- signed C. lupus (Fig. 1). The large size may simply be an indication of the Pleistocene age of the specimen because many other late Pleistocene carnivores were larger than their recent descendants (Graham 1981). Canis cf. dirus has been reported from the late Pleistocene Silver Creek fauna in the mountains east of Salt Lake City (Miller 1976). The nearest reported occurrences of C. lupus are from the latest Wisconsin to Recent Moonshiner Cave in Brigham County, Idaho _ (White et al. 1984), and Crystal Ball Cave in Millard County, Utah (Heaton 1985). Vulpes vulpes (Linnaeus) Red Fox ) MATERIAL.—UVP 82, left dentary with _ P,—M, and alveoli for P, and canine (Figs. 3, 5); UVP 81, left dentary with P,— M, and alve- oli for I—P; (Fig. 4). Discussion.—Anderson (1984) recognized five species of late Pleistocene foxes from North America, all of which are extant. The gray fox, Urocyon cinereoargenteus, is common in Ran- cholabrean faunas over much of North America (Kurten and Anderson 1980). The arctic fox, Alopex lagopus, is restricted to the arctic regions of North America, Europe, and Asia. It is rarely found in Pleistocene deposits, and in North America has only been reported from the Old Crow River, Yukon Territory (Anderson 1984). Vulpes velox, the swift fox, V. macrotis, the kit fox, and V. vulpes, the red fox are all common in Rancholabrean and Recent faunas of North America. Vulpes velox is the dominant small fox in Pleistocene faunas east of the Rocky Moun- tains, whereas V. macrotis is common in the western United States. Vulpes vulpes is more cosmopolitan in nature and has been identified in numerous Pleistocene sites from Virginia to California. 418 GREAT BASIN NATURALIST Vol. 46, No. 3 Fig. 4. Vulpes vulpes: UVP 81, lateral view of left dentary with P,— Msg. Solid bar represents | cm. Fig. 5. Vulpes vulpes: Lateral view of lower carnassial of UVP 82. Note well-developed cusp at inner junction of talonid and trigonid. Solid bar represents 1 cm. Members of the genus Vulpes differ from Urocyon in their relatively smaller molars and the shape of their mandible (Kurten and An- derson 1980). In Vulpes the“. . . lower border of the mandible forms an even curve without the lobe seen in Urocyon, . . .” whereas in the Arctic Fox “...the premolars are higher crowned, M, has a distinctly shorter talonid, and the tubercular teeth are more reduced than in Vulpes.” The smaller species of Vulpes, V. macrotis and V. velox, may have differences that are only sub- specific; recognition in the fossil record is based mainly on the geographic location of the fauna (Anderson 1984, Kurten and Anderson 1980). Both these forms are much smaller (body weight of 1.4-2.9 kg) than V. vulpes (4.5—-6.7 kg) and are easily recognized in the fossil record. The Bonneville specimens were compared to the large representation of V. vulpes from July 1986 NELSON, MADSEN: PLEISTOCENE CANIDS 419 TaBLE 3. Tooth dimensions in samples of Vulpes vulpes from Moonshiner Cave, Idaho, and the shoreline deposits of Lake Bonneville, Utah. Measurement N OR xX UVPO81 UVP082 Length P, 12 .80—0.90 85 — .88 Length P, 15 .87—1.00 94 — 93 Length P, 14 .82-1.07 .99 1.01 97 Length M, 16 1.44-1.63 1.53 10.62 1.40 Length M, 15 .64—-0.73 .70 .70 — Length P,—M, 16 5.27-6.18 5.75 5.68 — the late Pleistocene to Recent Moonshiner (120 individuals) and Middle Butte (46 indi- viduals) caves in southern Idaho and to Recent specimens (24 individuals) in the University of Utah mammal collections (Table 3). In size and morphology there appears to be little dif- ference in specimens from all localities. UVP 081 lacks the lower third molar and the signifi- cance of this feature is unknown (Fig. 4). In Recent specimens from Utah two individuals had a single dentary lacking the M;; the tooth was present in the other dentaries. In Pleis- tocene specimens from Little Box Elder Cave, Wyoming, 5 of 19 specimens lack an M;, whereas all 11 specimens from Jaguar Cave in Idaho have an M, (Kurten, written communication, 1985). Approximately 6% of the Moonshiner specimens lack the M3. Another variation in the tooth structure can be seen in the lower carnassial (Fig. 3). A small accessory cusp is developed in one of the Bonneville specimens at the internal junction of the trigonid and talonid (Fig. 5). Two Re- cent specimens from Utah lack this cusp, but it is present in 22 individuals (University of Utah mammal collection); however, the size of this cusp shows considerable variation. Kurten (1967) believed that there might be a connection between the cuspless morpho- types and a cold or continental climate. Hager (1972) observed 30 Recent speci- mens of V. vulpes from Colorado and Wyo- ming and reported that all were cusped mor- photypes. The significance of the cuspless or cusped morphotypes as an indicator of pale- oclimate, therefore, is probably very dubious for V. vulpes (Graham 1981). In examining the Recent specimens of V. vulpes, an additional variation in tooth struc- ture was also noted. An adult individual from Kuskokwim Delta, Alaska (UM 18291), lacked a right, lower, first premolar. Therefore, it appears that variation in dental makeup of V. vulpes is quite common. Documented Pleistocene specimens of V. vulpes have not been reported from Utah. The species is known from Crystal Ball Cave in western Utah (Heaton 1985), but the exact age of all elements of this fauna is difficult to ascertain, and faunal mixing may have oc- curred. ACKNOWLEDGMENTS We are grateful to Elaine Anderson and Bjorn Kurten for sharing their ideas on Pleis- tocene canids with us. Norman Negus allowed us access to the University of Utah mammal collections. Fort Hays State University and DINOLAB Inc. provided some financial assis- tance (Nelson). Jack Jackson from Fort Hays State University provided the photographs. LITERATURE CITED ANDERSON, E. 1984. Review of the small carnivores of North America during the last 3.5 million years. Pages 257-266 in H. H. Genoways and M. R. Dawson, eds., Contributions in Quaternary verte- brate paleontology: A volume in memorial to John E. Guilday, Carnegie Mus. of Nat. Hist. Spec. Pub. no. 8. Currey, D. R., G. ATWOOD, AND D. R. MaBEY. 1983. Major levels of Great Salt Lake and Lake Bonneville. Utah Geol. and Min. Surv. Map 73. GraHaM, R. W. 1981. Preliminary report on late Pleis- tocene vertebrates from the Selby and Dutton archeological/paleontological sites, Yuma County, Colorado. Contributions to Geology, 20(1): 33-56. Hacer, M. 1972. Recent vertebrate fauna from Chimney Rock Animal Trap, Larimer County, Colorado. Pages 63-71 in Contrib. to Geol., v. 11, no. 2. Heaton, T. H. 1985. The Quaternary paleontology and paleoecology of Crystal Ball Cave, Millard County, Utah: with emphasis on the mammals and the description of a new species of fossil skunk. Great Basin Nat. 45(3): 337-390. 420 KuRTEN, B. 1967. Dental microevolution. J. Dental. Res. Supplement to no. 5, 46: 824-826. KURTEN, B., AND E. ANDERSON. 1980. Pleistocene mam- mals of North America. Columbia University Press, New York. 442 p. MILLER, W. E. 1976. Late Pleistocene vertebrates of the Silver Creek local fauna from north central Utah. Great Basin Nat. 36(4): 387-424. Morrison, R. B. 1965. Lake Bonneville: Quaternary stratigraphy of eastern Jordan Valley, south of Salt Lake City. U.S. Geol. Sur. Prof. Paper 477. 80 pp. NELSON, M. E., AND J. H. MADSEN, JR. 1978. Late Pleis- tocene musk oxen from Utah. Trans. Kansas Acad. of Sci. 81(4): 277-295. . 1980. A summary of Pleistocene, fossil vertebrate localities in the northern Bonneville Basin of Utah. Pages 97-113 in J. W. Gwynn, ed., Great Salt Lake: a scientific, historical and economic overview, Utah Geol. and Min. Surv. Bull. 16. . 1983. A giant short-faced bear (Arctodus simus) from the Pleistocene of northern Utah. Trans. Kansas Acad. of Sci. 86(1): 1-9. GREAT BASIN NATURALIST Vol. 46, No. 3 Nowak, R. M. 1979. North American Quaternary Canis. Monograph Mus. of Nat. Hist., Univ. of Kansas, no. 6. 153 pp. Scott, W. E., W. D. McCoy, R. R. SHROBA, AND M. RUBIN. 1983. Reinterpretation of the exposed record of the last two cycles of Lake Bonneville, western United States. Quat. Res. 20: 261-285. STOCK, A. D., AND W. L. STOKES. 1969. A reevaluation of Pleistocene bighorn sheep from the Great Basin and their relationship to living members of the genes Ovis. J. Mammology 50(4): 805-807. STOKES, W. L., AND K. C. ConpiE. 1961. Pleistocene bighorn sheep from the Great Basin. J. Paleon. 35(3): 598-609. WHITE, J. A.. H.G. MCDONALD, E. ANDERSON, AND J. M. SoIsET. 1984. Lava blisters as carnivore traps. Pages 241-256 in H. H. Genoways and M. R. Dawson, eds., Contributions in Quaternary verte- brate paleontology: a volume in memorial to John E. Guilday, Carnegie Mus. of Nat. Hist. Spec. Publ., no. 8. NOTE ON FOOD HABITS OF THE SCREECH OWL AND THE BURROWING OWL OF SOUTHEASTERN OREGON Barbara A. Brown’, John O. Whitaker’, Thomas W. French!”, and Chris Maser® ABSTRACT.— Diets of the Common Screech Owl (Otus asio) and Burrowing Owl (Athene cunicularia) from the Great Basin, Malheur County, southeastern Oregon, were studied. Although there was considerable overlap in the diets of these owls, there were differences related to habitat use. Few data are available on the food habits of owls from the Great Basin of southeastern Oregon. The Barn Owl (Tyto alba) is the only one whose food habits have been studied in this part of the state (Maser et al. 1980), al- though some data are available on food habits of owls from the rangelands of central Oregon: Barn Owl (Maser and Hammer 1972); Great Horned Owl (Bubo virginianus) (Brodie and Maser 1967, Maser et al. 1970); Short-eared Owl (Asio flammeus) (Maser et al. 1970; Maser et al. A note on the food habits of the short-eared owl, 1971); Long-eared Owl (A. otus) (Maser et al. 1970), and Burrowing Owl (Athene cunicularia) (Maser et al. Food habits of the burrowing owl, 1971). This paper presents information on food habits of the common Screech Owl (Otus asio ) and the Burrowing Owl in the rangelands of Malheur County, Oregon. STUDY AREA The study area, Malheur County, in ex- treme southeastern Oregon, lies within the Owyhee Upland physiographic province. The major vegetation zone is described as shrub- steppe (characterized by big sagebrush, Artemisia tridentata) (Franklin and Dyrness 1973). Plant communities were defined by Dealy et al. (1981), and the more restrictive habitats were described by Bohn et al. (1980) and Maser et al. Geomorphic and edaphic habitats, 1979; Maser et al. Manmade habi- tats, 1979). METHODS Castings were collected from April 1975 through July 1978. They were placed in plas- tic bags and were soaked in water before dis- section. Prey items were identified to species whenever possible, and individuals were counted. Total counts of leaves and seeds were taken, but other plant parts, fur, and feathers were listed only as the number of pellets in which they occurred. Comparisons between vertebrate and invertebrate foods were based on total percentages. Diversity of prey was calculated for all food items as the number of items per total number of castings. RESULTS AND DISCUSSION Vertebrates formed 20.2% of the prey indi- viduals in screech ow] diets (Table 1); inverte- brates, 79.8% (Table 2). Vertebrates com- prised 14.3% of the prey items in burrowing owl diets (Table 3) and invertebrates 85.7% (Table 4). Vertebrate Prey Both owls fed heavily on the Ord kangaroo rat (Dipodomys ordi), but the kangaroo rat was more important to the Burrowing Owl than to the Screech Owl. The northern pocket gopher (Thomomys talpoides) was important in the diet of the Burrowing Owl but accounted for less than 1% of the Screech Owl diet. The similarity in weight between the Ord kangaroo rat (the average weight of 32 individuals from both Department of Life Sciences, Indiana State University, Terre Haute, Indiana 47809. ?Present address: The Nature Conservancy, Eastern Regional Office, 294 Washington Street, Boston, Massachusetts 02108. USDI Bureau of Land Management, Forestry Sciences Laboratory, Corvallis, Oregon 97331. 421 422 GREAT BASIN NATURALIST Vol. 46, No. 3 TABLE 1. Vertebrate foods of the Screech Owl (Otus asio) from southeastern Oregon, based on analysis of 205 castings. Number of Percentage of Number of Percent Prey item individuals diet castings frequency MAMMALIA Rodentia Cricetidae Peromyscus sp. 16 5.2 10 4.9 Cricetinae unidentified 69 22.6 4] 20.0 Microtinae unidentified 25 8.2 18 8.8 Microtus sp. 2 0.6 1 0.5 Lagurus curtatus 11 3.6 10 4.9 Geomyidae Thomomys talpoides 2 0.6 2 1.0 Heteromyidae Dipodomys ordi 39 12.8 32 15.6 Perognathus parvus 1 0.3 1 0.5 Scuiridae Spermophilus sp. 8 2.6 1 0.5 Sciuridae unidentified 16 ee, 16 7.8 Lagomorpha Leporidae Lepus californicus 1 0.3 1 0.5 Leporidae unidentified 16 5.2 11 5.4 Mammal Unidentified 47 15.4 46 22.4 REPTILIA Squamata Iguanidae Phrynosoma platyrhinos JU 3.6 9 4.4 Cnemidophorus tigris 1 0.3 1 0.5 Lacertilia Unidentified 23 7.5 10 4.9 Reptile Unidentified sil 3.6 11 5.4 AVES < Columbiformes Columbidae Zenaida macroura 1 0.3 1 0.5 Passeriformes Unidentified 2 0.6 2 1.0 Bird Unidentified 3 1.0 1 0.5 *Eggshell Unidentified — — 6 *Feathers Unidentified — — 5 *Claws Unidentified = = 8 305 99.5 *Not included in total count central and southeastern Oregon was 56.8 g; Maser, unpublished data), and the northern pocket gopher in Malheur County (61.3 g) probably allowed the Burrowing Owl to ex- ploit both species. These gophers formed only 0.02% of the Burrowing Owl diet in central Oregon (Maser et al. Food habits of the bur- rowing owl, 1971). Northern pocket gophers of the poorly drained lacustrine soils that oc- cur in Malheur County are small (average weight of 25 individuals was 61.3 g) compared with the same subspecies (quadratus) from the better drained soils of the Steens Moun- tain, Harney County (average weight of eight individuals, 94.6 g), and from the sandy soils of central Oregon, Jefferson and Klamath counties (average weight of 47 individuals 67.4 g; Maser, unpublished data). Possibly, the weight difference (6.1 g) between the go- phers of central and southeastern Oregon al- lowed the Burrowing Owl to exploit this prey in one area but not in the other. Within the family Cricetidae, the Screech Owl had the following taxa available as prey: western harvest mouse (Reithrodontomys megalotis ), deer mouse (Peromyscus manicu- latus), canyon mouse (P. crinitus), northern grasshopper mouse (Onychomys — leuco- gaster), desert woodrat (Neotoma lepida), bushy-tailed woodrat (N. cinerea), montane vole (Microtus montanus), long-tailed vole (M. longicaudus ), and sage vole (Lagurus cur- tatus ). Montane voles were by far more abun- dant than long-tailed voles (Maser, unpub- lished data). The family Cricetidae accounted for 40.2% of the vertebrate prey items. The July 1986 BROWN ET AL.: OREGON OWLS 423 TABLE 2. Invertebrate foods and vegetation of the Screech Owl (Otus asio) from southeastern Oregon, based on analysis of 205 castings. Number of Percentage of Number of Percent Prey items individuals diet castings frequency INSECTA Coleoptera Carabidae Calosoma sp. 90 U5) 34 16.6 Near Anisodactylus sp. 6 0-5 5 2.4 Unidentified 3 0.2 3 1.5 Anisodactylus sp. i 0.6 1 0.5 Curculionidae Unidentified 6 0.5 10 4.8 Alleculidae Unidentified 1 0.1 1 0.5 Tenebrionidae Unidentified 61 Dull 33 16.1 Scarabaeidae Cyclociphala sp. 41 3.4 30 14.6 Rutela sp. 1 0.1 1 0.5 Unidentified 4 0.3 4 2.0 Paracotalpa granicollis 6 0.5 4 2.0 Silphidae Necrophorus sp. ll 0.9 a 3.4 Coleoptera Unidentified 21 Hed 20 9.8 Diptera Unidentified 3 0.2 3 eo) Hemiptera Unidentified 1 0.1 1 0.5 Homoptera Cicadidae Unidentified 81 6.7 21 10.2 Hymenoptera Formicidae Unidentified 20 16.6 12 5.9 Lepidoptera Larvae 3 0.2 1 0.5 Neuroptera Unidentified 1 0.1 i 0.5 Orthoptera Acrididae Unidentified (mandibles) 424 Sy 48 23.4 Gryllidae Gryllus veletis 59 4.9 35 ei Unidentified (mandibles) 27 D9) 8 3.9 Stenopalmatidae Stenopelmatus sp. (mandibles) 117 9.7 19 9.3 Siphonaptera Unidentified 2 0.2 1 0.5 ARACHNIDA Araneida Unidentified 4 0.3 4 2.0 Scorpionida Vejovidae Vejovis boreus 24 2.0 16 7.8 VEGETATION Leaves Erigonum as — 4] *Vegetation — = 1] *Seeds Unidentified 2 — 1 0.5 *Grass seeds 48 = 3 eS : 1,204 99.7 *Not included in total count subfamily Cricetinae accounted for 27.8% and Microtinae 12.4% of the vertebrate diet. The Burrowing Owl had the same taxa available as prey within the family Cricetidae as did the Screech Owl, except for the canyon mouse and occasionally the desert woodrat. Cricetidae formed 29.9% of the Burrowing Owl vertebrate diet. The subfamily Criceti- nae composed 10.3% and the Microtinae 15.7%. As with the Screech Owl, the montane vole was far more abundant in habitat of the Burrowing Owl than was the long-tailed vole (Maser, unpublished data). Invertebrate Prey Although the Screech Owl ate 26 kinds of in- vertebrates (Table 2) and the Burrowing Owl ate 24 kinds (Table 4), there are some major differ- ences. Beetles accounted for 21.4% of the items in the Screech Owl diet, including Scarabaeidae (4.3%) and Carabidae (8.8%). Beetles were slightly more important to the Burrowing Owl (38.6% of the diet). Although the Burrowing Owl used Carabidae about the same as the Screech Owl (7.8%), Scarabaeidae were more important to the Burrowing Owl (21.7%). 424 GREAT BASIN NATURALIST Vol. 46, No. 3 TABLE 3. Vertebrate foods of the Burrowing Owl (Athene cunicularia ) from southeastern Oregon, based on analysis of 150 castings. Prey item MAMMALIA Number of individuals Rodentia Cricetidae Peromyscus maniculatus 11 Reithrodontomys meglotis 7 Microtus sp. 10 Lagurus curtatus 13 Cricetinae Unidentified 3 Microtinae Unidentified 9 Cricetidae Unidentified 8 Geomyidae Thomomys talpoides 21 Heteromyidae Dipodomys ordi 49 Perognathus parvus 11 Unidentified 1 Sciuridae Unidentified 14 Mammal Unidentified 19 AMPHIBIA Unidentified 21 REPTILIA Squamata Iguanidae Phrynosoma platyrhinos Reptile Unidentified AVES Columbiformes Columbidae Zenaida macroura *Feathers Percentage of Number of Percent diet castings frequency 5.4 10 6.7 3.4 7 4.7 4.9 8 5.3 6.4 11 7.3 1.5 3 2.0 4.4 8 5.3 3.9 6 4.0 10.3 21 14.0 24.1 47 31.3 5.4 ll 7.3 0.5 1 0.7 6.9 14 9.3 9.4 18 12.0 10.3 14 9.3 0.5 1 0.7 2.0 4 eri; 0.5 1 0.7 eeesy tat 4 99.8 *Not included in total count The other insect order of major importance to both owls was Orthoptera. This item was more important to the Screech Owl (52.0%) than to the Burrowing Owl (34.3%). Within Orthoptera, grasshoppers (Acrididae) were more important to the Screech Owl (35.2%) than to the Burrowing Owl (21.6%), but the Jerusalem cricket (Stenopelmatus sp.) was eaten more by the Burrowing Owl (12.7%) than by the Screech Owl (9.7%). The Screech Owl also consumed the cricket (Gryllus veletis) (4.9%), but the Burrowing Owl did not. Prey Diversity Total prey diversity per casting for the Screech Owl was 0.3 species and averaged 7.4 individual items. The Burrowing Owl was sur- prisingly close, 0.3 species per casting and averaged 9.5 individuals. Owls The Screech Owl generally inhabited ripar- ian zones, abandoned homesteads, and some cliffs (Bohn et al. 1980; Dealy et al. 1981, Maser et al. Geomorphic and edaphic habi- tats, 1979; Maser et al. Manmade habitats, 1979; Maser, unpublished data). The Burrow- ing Owl, on the other hand, was associated with badger (Taxidea taxus ) burrows, primar- ily in the basin big sagebrush/bunchgrass and black greasewood (Sarcobatus vermiculatus )/ grass communities (Dealy et al. 1981; Maser, unpublished data). Flexibility in selection of habitat by the Screech Owl brought it into contact with a wider prey base than was available to the Bur- rowing Owl with its more rigid selection of habitat. For example, Screech Owls in cliffs had canyon mice and both species of woodrats available; Screech Owls in abandoned home- steads also had both species of woodrats avail- able and were known to take the desert woodrat (Maser, unpublished data). The Bur- rowing Owl, however, occupied habitat that was inhospitable to canyon mice and to bushy- tailed woodrats, and the desert woodrat only occasionally inhabited the black greasewood/ grass community (Maser, unpublished data). Both species of owl are opportunistic and catholic in diet (Gleason and Craig 1979; Maser et al. Food habits of the burrowing July 1986 BROWN ET AL.: OREGON OWLS 425 TABLE 4. Invertebrate foods and vegetation of the Burrowing Owl (Athene cunicularia) from southeastern Oregon, based on analysis of 150 castings. Number of Percentage of Number of Percent Prey item individuals diet castings frequency INSECTA Coleoptera Carabidae Calosoma sp. 51 4.2 36 24.0 Anisodactylus sp. 31 25 13 8.7 Unidentified 14 1.1 8 5.3 Curculionidae Unidentified 19 1.6 11 U8) Scarabaeidae Paracotalpa granicollis 121 9.9 25 16.7 Cyclocephala sp. 1 0.7 I 0.7 Rutela sp. 88 Co? 26 17.3 Unidentified 48 3.9 9 11.3 Tenebrionidae Unidentified 30 2.5 22 14.7 Silphidae Necrophorus sp. 49 4.0 30 20.0 Elateridae Unidentified 11 0.9 2 ES Coleoptera Unidentified 1 0.1 1 0.7 Diptera Unidentified 28 2.3 4 od Homoptera Cicadidae Unidentified 13 1.1 12 8.0 Hymenoptera Formicidae Unidentified 10 0.1 3 2.0 Braconidae Unidentified 98 8.0 8 5.3 Lepidoptera Unidentified (larvae) 7 0.6 3 2.0 Orthoptera Acrididae Unidentified 263 21.6 55 36.7 Stenopelmatidae Stenopelmatus sp. (mandibles) 155 12.7 9 6.0 Insect Unidentified 2 0.2 2) 1.3 Unidentified (mandibles) 48 3.9 5 oe ARACHNIDA Scorpionida Vejovidae Vejovis boreus 126 10.3 56 Boe Araneida Unidentified 3 0.25 3 2.0 Acari Unidentified 2 0.2 2 IES VEGETATION *Leaves Eriogonum sp. 103 18 *Vegetation 17 6 *Seeds Grass 52 2 Feathers Unidentified 1 i 1,219 100.3 *Not included in total count owl, 1971; Smith and Wilson 1971; Zarn 1974), and their diets in southeastern Oregon overlapped considerably. They used totally different habitats, however, which physically isolated the owls and avoided competition. ACKNOWLEDGMENTS Sam Shaver (Vale, Oregon) helped collect _ the castings. D. Grayson (Department of An- _ thropology, University of Washington, Seat- | tle), Z. Maser (Department of Forest Science, Oregon State University, Corvallis), and D. ' Toweill (Department of Fisheries and | Wildlife, Oregon State University, Corvallis) read and improved the manuscript. V. Bissell (USDI Bureau of Land Management, Forestry Sciences Laboratory, Corvallis, Ore- gon) typed the various drafts. We thank them for their help. LITERATURE CITED BOHN, C., C. GALEN, C. MASER, AND J. W. THomas. 1980. Homesteads—manmade avian habitats in the rangelands of southeastern Oregon. Wildl. Soc. Bull. 8: 332-341. BropIE, E. D., JR., AND C. MASER. 1967. Analysis of great horned ow! pellets from Deschutes County, Ore- gon. Murrelet 48: 11—12. 426 DEALY, J. E., D. A. LECKENBY, AND D. M. CONCANNON. 1981. Wildlife habitats in managed rangelands— the Great Basin of southeastern Oregon: plant communities and their importance to wildlife. USDA For. Serv. Gen. Tech. Rep. PNW-120. Pac. Northwest For. and Range Expt. Sta., Port- land, Oregon. 66 pp. FRANKLIN, J. F.. AND C. T. DyrNEss. 1973. Natural vegeta- tion of Oregon and Washington. USDA For. Serv. Gen. Tech. Rep. PNW-8. Pac. Northwest For. and Range Expt. Sta., Portland, Oregon. 417 pp. GLEASON, R. L., AND T. H. Craic. 1979. Food habits of Burrowing Owls in southeastern Idaho. Great Basin Nat. 39: 274-276. MasSER, C., J. M. GEIsT, D. M. CONCANNON, R. ANDERSON, AND B. LOVELL. 1979. Wildlife habitats in man- aged rangelands—the Great Basin of southeastern Oregon: geomorphic and edaphic habitats. USDA For. Serv. Gen. Tech. Rep. PNW-99. Pac. North- west For. and Range Expt. Sta., Portland, Ore- gon. 84 pp. MASER, C., AND E. W. HAMMER. 1972. A note on the food habits of barn owls in Klamath County, Oregon. Murrelet 53: 28. GREAT BASIN NATURALIST Vol. 46, No. 3 Maser, C., E. W. HAMMER, AND S. H. ANDERSON. 1970. Com- parative food habits of three owl species in central Oregon. Murrelet 51: 29-33. ____. 1971. Food habits of the Burrowing Owl in central Oregon. Northw. Sci. 45: 19-26. Maser, C., E. W. HAMMER, AND R. MASER. 1971. A note on the food habits of the Short-eared Owl (Asio flammeus) in Klamath County, Oregon. Murrelet 52: 27. Maser, C., S. SHAVER, C. SHAVER, AND B. PRICE. 1980. A note on the food habits ofthe Barn Owl in Malheur County, Oregon. Murrelet 61: 78-80. Maser, C., J. W. THomas, IL. D. LUMAN, AND R. ANDERSON. 1979. Wildlife habitats in managed rangelands—the Great Basin of southeastern Oregon: manmade habi- tats. USDA For. Serv. Gen. Tech. Rep. PNW-86. Pac. Northwest For. and Range Expt. Sta., Portland, Oregon. 40 pp. Situ, D.G., ANDC. R. WILSON. 1971. Notes on winter food of screech owls in central Utah. Great Basin Nat. 31: 83-64. ZARN, M. 1974. Habitat management series for unique or endangered species: Burrowing Owls, Speotyto cu- nicularia hypugaea. U.S. Dept. Inter., Bur. Land Manage. Tech. Note. 250, Rep. 11, Denver, Colo- rado. 25 pp. Purchased by the USDA Forest Service for official use. : | DISEASES ASSOCIATED WITH JUNIPERUS OSTEOSPERMA AND A MODEL FOR PREDICTING THEIR OCCURRENCE WITH ENVIRONMENTAL SITE FACTORS E. D. Bunderson!, D. J. Weber’, and D. L. Nelson® ABSTRACT.—On 17 Utah juniper (Juniperus osteosperma [Torrey] L.) sites studied in Utah, Gymnosporangium inconspicuum was the most common rust fungus, followed in frequency and severity by G. nelsoni, G. kernianum, and G. speciosum. The incidence of G. kernianum was correlated with moderate temperatures and greater than average precipitation. True mistletoe, Phoradendron juniperinum Engelmm., was present on seven sites. Incidence of foliage diseases of the mold-mildew type was low on sites with low spring and summer temperatures and high on sites with high summer and fall precipitation. Wood rot was common, and incidence seemed to be correlated with low winter temperatures and low soil nitrate but not with annual precipitation. Needle blight, shoot dieback, and needle cast symptoms were common and considered of abiotic origin. Their nonparasitic nature was indicated by lack of association with pathogenic organisms and the positive correlation of their incidence with winter injury and summer drought factors. Needle blight was also positively correlated with high soil salinity but negatively with high soil calcium regardless of salinity. A nonparametric model was developed that accurately predicted the frequency of the mold-mildew type diseases of J. osteosperma based on measured environmental site factors. The pinyon-juniper woodland is a wide- spread vegetation type in the southwestern United States, estimated to cover from 30 to 40 million hectares (Allred 1964). Histori- cally, pinyon-juniper woodland vegetation has provided numerous benefits including fuel, building materials, charcoal, nuts, Christmas trees, medicines, etc. (Tueller et al. 1979, Hurst 1977, Lanner 1975, Cronquist et al. 1972, Gallegos 1977). About 80% of the total acreage is grazed, contributing signifi- cantly to the available forage for livestock and wildlife (Clary 1975), and pinyon-juniper woodlands are becoming increasingly valued for their watershed, aesthetic, and recre- ational values (Gifford and Busby 1975). This ecosystem is a large component of the vegetation of Utah (62,705 km° or 28.6%, Kuchler 1964), and it has the potential to add substantially to the economic and aesthetic activity of the state. Despite this potential, little research has been done to explore the physiological relationships and autecology of this vegetation type. Many complex environmental factors con- tribute to the variety of interactions in a pinyon-juniper ecosystem. Consumable “sup- _ ply factors” of the environment such as light, Utah Technical College, Provo, Utah 84604. Fig. 1. Location of 17 study sites in Utah. water, nutrients, oxygen, and carbon dioxide interact with “site quality” or “condition” fac- tors such as temperature and precipitation (Harper 1977). The effects of species density, Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. SUSDA Forest Service, Intermountain Research Station, Shrub Sciences Laboratory, Provo, Utah 84601. AS @ 428 GREAT BASIN NATURALIST Vol. 46, No. 3 TABLE 1. Description of symptoms and assessment methods for diseases of Juniper osteosperma. Injury Needle blight Needle cast Tip dieback Pathogen Gymnosporangium rusts: G. inconspicuum G. nelsoni G. kernianum G. speciosum Diseases of J. osteosperma symptom or sign Senescent or decadent foliage or twigs. Terminals may show yellow or red-brown coloration. May or may not show evidence of fungal fruiting bodies. Bare twig terminals from which foliage scale leaves have fallen (twig terminals of living branches). Senescent or decadent foliage or twig terminals showing brown to grey coloration. May or may not show evidence of fungal fruiting bodies. Presence of follicolous or caulicolous telia having cartiform pedicels accompanied by little or no fasciation of branches. Presence of galls with irregularly compressed, wedge-shaped telia; on twigs or branches. Presence of well-defined or modified witch's brooms (branch fasciation) with foliicolous, bluntly conical telia. Presence of cristiform or crenate telia on-fusiform Rating scale Percent of the total plant affected Percent of the total plant affected Percent of the total plant affected Rated as light, moderate, or heavy: a) light: 1-3 telial infect- tions in a 2 ft breast high circum- ference section of tree. b) moder- ate: 4-10 telial infections in equivalent area. c) heavy: more than 10 telial infections in equiv- alent area. Actual or estimated number of galls per tree. Actual or estimated number of witch's brooms per tree. Actual or estimated number of swellings. Wood rot fungi (i.e., Fomes juniperinum) branches. Fungal fruiting bodies may or may not be present. Mold and/or mildew Fungal mycelial growth on foliage. (unidentified spp.) Mistletoe (Phoradendron juniperinum) habitable niches, pathogens, seasonality, and roles of predators such as grazing animals in- fluence the diversity and stability of pinyon- juniper woodlands, as do naturally occurring catastrophic events (i.e., fire) or deliberate manipulation or intervention by man. The ecological dynamics of the pinyon- juniper ecosystem have been studied by Pear- son (1920), Woodbury (1947), Daniel et al. (1966), and Vasek (1966) and typically have been oriented toward range and resource management. Synecological studies of pinyon-juniper have included work in latitu- dinal and elevational patterns (Daubenmire 1943), interactions with understory vegeta- tion (West et al. 1975), paleoecological influ- ences (Cottam 1959), and climatic and edaphic relationships (Beeson 1974, Hunt Presence of heart rot or decayed areas in trunk or Presence of viable mistletoe foliage in tree. fusiform swellings per tree. Presence rated as light, moder- ate, or heavy. Presence rated as light, moder- ate, or heavy. Actual or estimated number of growth areas per tree. 1974). Several studies have treated disease and/or insect factors of pinyon pines (McCam- bridge and Pierce 1974, McGregor and San- drin 1968, Hepting 1921), but little research in these particular areas has dealt with ju- niper. The objectives of this study were to survey the diseases of Utah juniper (Juniperus os- teosperma [Torrey] L.), particularly those in- duced by fungi, and to relate their occur- rence, frequency, and severity to selected environmental factors in several pinyon- juniper habitats throughout Utah. MATERIALS AND METHODS Seventeen representative pinyon-juniper sites in Utah were studied from April to Octo- July 1986 BUNDERSON ET AL.: JUNIPER DISEASES 429 TABLE 2. Location of 17 primary juniper study sites in Utah. Site name Site no. Weather station Jackson Springs 1 St. George Tobin Bench 2 Veyo Power House Peters Point 3 Monticello Alkali Ridge 4 Blanding Cyclone Flat 5 Natural Bridges Natl. Mon. Indian Peak 6 Desert Expt. Range Ephraim 7 Ephraim Sorensens Fld. Manti 8 Manti Black Mountain 9 Salina Triangle Mountain 10 Salina Beaver Ridge ll Beaver Gordon Creek 12 Hiawatha Dutch John 13 Flaming Gorge Taylor Flat 14 Allen’s Ranch Rabbit Gulch 15 Hannah Henry Mountains (Stevens Narrows) 16 Capitol Reef Natl. Park Henry Mountains (Airplane Flat) i, Boulder Fig. 2. Observations of four Gymnosporangium spe- cies in selected Utah sites. | ber 1982 (Fig. 1). Sites were selected for cli- max pinyon-juniper vegetation and minimal _ structural disturbance by people, although _ most sites had a long history of grazing by _ domestic animals and wildlife. Soils were pri- | marily derived from marine shales, conglom- | erates, siltstones, and sandstones (Bunderson et al. 1985). Transects at each site consisted of 96 Utah juniper trees randomly selected by the quar- ter method (Phillips 1959). Each tree was measured for height, stem diameter, and age and then assessed for signs and symptoms of disease. Diseases were classed in three causal categories: (a) rust fungi (Gymnosporangium spp.), (b) miscellaneous fungus diseases (e.g., tip burn, die back, stem decay), and (c) para- sitic higher plants. Nonparasitic injury was also assessed. Descriptions of diseases, patho- genic agents, and assessment methods are found in Table 1. The percentage of deca- dence for each tree was estimated and an overall vigor score (1 = good, 2 = moderate, 3 = poor) was assigned to each tree. Since some areas of the state were not rep- resented in our 17 primary sites (Table 3), we added 15 additional sites (Fig. 2) in which just the incidence of Gymnosporangium rusts was studied. We used the standard U.S. Forest Service ratings as listed in the addendum to Table 4. Since each site had its own pattern of infec- tion, we calculated severity indexes to reflect both the total number of trees infected with each causal agent and the severity of that in- fection on each tree (Table 2). We measured concentrations of several soil mineral nutrients (N, P, K, Ca, Mg, Na, Fe, SO,) and other soil properties (pH, EC, sand, silt, clay) by digging five soil pits along each transect and pooling samples from the top 12 inches of soil from each pit. Soils were acid digested and analyzed for mineral content us- ing a Technicon II Auto Analyzer and atomic absorption spectroscopy. Temperature and 430 GREAT BASIN NATURALIST Vol. 46, No. 3 TABLE 3. Observations of three Gymnosporangium species in selected Utah sites. G. inconspicuum G. nelsoni G. kernianum . Immigrant Pass Rosette . Bear Lake . Hwy 39, 6 mi west of Woodruff 5. Chalk Creek 6. Hwy 150—Yellow Pine Campground 7. Hastings Canyon 8. Lookout Pass 9. 12 mi Northeast of Eureka 10. Silver City 11. Scipio Pass 12. Sego 13. 2 misouth of Fremont Junction 14. Escalante Canyon 15. 8 mi east of Parowan (very heavy) Sener oo QaQarea (some fasciation) Oe fre ozZooqg orf (on both foliage and twigs) > Oro C1@ (no live galls) (heavy when present) (very heavy) (most galls dead) (heavy when present) BA OAPAMNOG AO Prere@e aeQ. Of OO@S©O ez A Abundant (€ Common O Occasional R Rare N None observed in area precipitation records were obtained from Na- tional Oceanic and Atmospheric Administra- tion recording stations at or near the research sites. For complete descriptions of the 17 pri- mary site locations and weather recording sta- tions, see Table 3. These values were divided seasonally into four quarters: Decem- ber—February, March—May, June-August, September—November. Location and _ fre- quency of alternate hosts for Gymnospo- rangium rusts were also noted. We used linear correlation coefficients to relate frequencies and severity of the diseases of Utah juniper to the soil factors, tempera- ture, and precipitation. A nonparametric weighted least squares analysis for categorical data tested hypotheses relating interactions between the environmental data and patho- genic factors (Forthofer and Lehnen 1981). The hypotheses were checked for statistical significance and offered as a possible tool to predict the incidence of disease that would occur within a given site if the conditions of the hypotheses were met. RESULTS AND DISCUSSION Gymnosporangium Rusts Frequent occurrences of Gymnospo- rangium spp. on Utah juniper have been recorded in Utah (Arthur 1934, Hodges 1962, Peterson 1967), but these occurrences have not been quantified. Figures 3 and 4 quantify the frequency of the three Gymnosporangium rusts we encountered most often on Utah ju- niper in the 17 transects. Gymnosporangium inconspicuum Kern was present on a substantial proportion of the trees at 15 of the 17 sites. The caulicolous and/or foliicolous telia of G. inconspicuum are inconspicuous and difficult to locate, and probably more trees were infected than recorded. The high frequencies support ob- servations by others that G. inconspicuum has a high endemic population in juniper stands throughout Utah (Peterson 1967). We exam- ined the two sites where G. inconspicuum was not recorded on three separate occasions without finding any indication of this fungus. Since these sites are located in the St. George area in the warm Mohave Desert ecotype rather than in the cooler Great Basin ecotype, environmental factors may influence the oc- currence of this rust in ways other than those we have measured. | We found Gymnosporangium nelsoni Arth. in all research sites. The galls stimulated by this rust on stems and branches of J. os- teosperma varied from 2 to 12 cm in diameter, and both live and dead galls were counted until gall numbers exceeded 300 per tree. Excluding the St. George sites, incidence of "i { July 1986 BUNDERSON ET AL.: JUNIPER DISEASES 431 TABLE 4. Description of rating system for diseases of Juniper osteosperma. Infection level = Number of trees infected with pathogen. Severity a. G. inconspicuum = Average rating of infections per tree 0 = No infection 1 = Light infection 2 = Medium infection 3 = Heavy infection b. G. nelsoni = Average number of galls/tree c. G. kernianum = Average number of witch's broom/tree Severity Index a. G. inconspicuum = Proportion of infected trees (expressed as a %) Site rating — 3 b. G. nelsoni = Severity x 100 Average number galls/meter — proportions of infected trees (%) c. G. kernianum = Severity Average height of trees at site — proportions of infected trees (%) Mold mildew Severity = Percentage of trees infected; Severity index = Proportion (%) < percentage infected (severity) Wood-rotting fungi Severity = Sum of percentage of trees infected with trunk, branch, twig rot; Severity index = Proportion (%) severity Mistletoe Severity = Average number of broom/tree; severity index = Proportion (%) < severity Needle blight Severity = Average percentage of infection/infected tree; Severity index = Proportion (%) x severity Needle cast Severity = Average percentage of mildew on all infected trees (cumulative); Severity index =’Proportion (%) * severity Tip dieback Severity = Average percentage of infection/infected tree; Severity index = Proportion (%) * severity G. nelsoni was much more variable than that of G. inconspicuum. We found Gymnosporangium kernianum Bethel on fewer sites than either G. incon- spicuum or G. nelsoni. This species stimu- lates shoot fasciations (witch's brooms) on which foliicolous telia are formed. Numbers of brooms per infected tree ranged from 0 to 23 and size of brooms from 3 to 130 cm in diame- ter. The incidence of G. kernianum was con- siderably less than that of either G. incon- spicuum or G. nelsoni. As with G. incon- spicuum, only active infections were re- corded. Correlations between temperature and precipitation in our research sites indicate that G. kernianum is favored by moderate temperatures and a greater than average an- nual precipitation, particularly during the summer (Tables 5, 6). We found a fourth species, Gymnospo- rangium speciosum, on only one primary site (Site 12) and one supplementary site (Sego). It forms bright orange cristiform or crenate telia on fusiform stem swellings, with loose fascica- tions usually associated with the infected area. Variation in host susceptibility complicates establishing cause and effect relationships with physical environmental factors for Gym- nosporangium rusts. Differences in resistance to cedar-apple rust have been known for many years (Moore 1940) as well as differences in pathogenicity of the rust (Aldwinkle 1975). It has been suggested that the genetic variability Gymnosporangium inconspicuum Percent of infected trees oO oO 40 30 20 10 0 1 2 3 4° 65 & 7 -& 9 BVO vd da IW 14 WS IG V7 Sites 90 Gymnosporangium nelsoni 80 7) oo 2 mo} 2 Ss) 2 = io) ie o 2 ® a ? 2 4 8 @ “4% 8 8 VON) 12 18 1 18 1) Iza ' Sites 90 BO Gymnosporangium kernianum 7p) ® 2 no} so) [s) 2 iS re) ic ® o. 7) a U #2. 3 OS BB FBO 10 dW 1 A Ae WS GB. U7 Sites Fig. 3. The percent of infected trees (Juniperus os- teosperma) in relation to three rusts. Top chart (Gymnospo- rangium inconspicuum), middle (Gymnosporangium nel- soni), bottom (Gymnosporangium kernianum). of Utah juniper may surface in the average num- ber of stems in any given stand (Kimball Harper, personal communication). After pooling the data from the 17 sites, we found a correlation be- tween environmental parameters such as amount of precipitation and number of juniper stems (a = 0.01 with average annual precipita- tion, a = 0.05 with summer precipitation), indi- cating that as the total precipitation decreases the number of stems. increases. However, there are also positive correlations between increasing numbers of stems and the increased frequencies of the various rusts (a = 0.05 for G. nelsoni, GREAT BASIN NATURALIST Vol. 46, No. 3 80 70 Gymnosporangium inconspicuum 60 50 40 Severity index 30 1 2°95 G2 5G 7 8 © 10 Wi 12 WS 14 WS WB 47 Sites 30 Gymnosporangium nelsoni Severity index TO ad ve ws) Wes aS aS aly 4.0 Sites 3.5 Gymnosporangium kernianum Severity index 10 11 12 13 14 15 16 17 1} 2. 8c OG 7.-8..09 Sites Fig. 4. The severity index of three rusts (top) Gym- nosporangium inconspicuum, (middle) Gymnospo- rangium nelsoni, (bottom) Gymnosporangium kerni- anum on Juniperus osteosperma trees. a = 0.10 for G. kernianum). Gymnosporangium inconspicuum has been described as either folii- colous or caulicolous, and our data indicate that G. inconspicuum on the foliage correlates posi- tively to the number of stems of J. osteosperma (a = 0.10) and the caulicolous form of G. incon- spicuum correlates negatively to the number of stems (a = 0.10). Further studies are needed to understand these correlations. Miscellaneous Fungus Diseases Several types of wood-decay fungi were in- dicated along our transects (Figs. 6, 7) by July 1986 BUNDERSON ETAL.: JUNIPER DISEASES 433 TABLE 5. Correlations between J. osteosperma diseases and insects and temperature in the different time quarters (Q). 5-year average HiQ Low Q Annual 5-year 5-year Pathogen Q, Q, Q; Q, average average average high low Needle blight —.55* —.45 —.40 —.55* —.43 —.63* Soll — 4 —.44 Needle cast —.04 12 .10 13 02 —.09 10 .07 12 Tip dieback —.14 — .06 = IL == Pall = 2 —.08 —.16 .20 = oD) Mold-mildew —.46 —.49* = ond —.4] Saye RAT —.46 = po) Soll) Galls-foliage .07 — 11 .00 14 —.09 18 04 = 587 41 Galls-branch PAU —.32 — .24 —.20 — .33 al = 26 — .64* —.09 Witch’s broom —.43 —.55 —.4] —.45 —.56* —.34 =50* = 65% =pIl6 Lesions-foliage —.30 —.34 —.44 —.33 —.38 iD, = 6X0} — .26 oA) Lesions-branch — .65** SOO —.37 aA ics —.43 as = le = (GBF 929) Rot-twigs = All —.05 —.09 —.11 —.16 —.50* alls} —.13 —.14 Rot-trunk —.48* —.38 = 20 —.39 =< 3 —.39 = 5 6h9) —.06 —.30 Mistletoe 10 04 13 14 .13 ell 13 —.29 15 Insect borers —.47* =. 2) = 2 —.38 —.16 =o — 33 —.09 — .48* Scale insects =) —.29 —.36 —.30 =o —.30 —.29 —.38 —.19 Insect galls 19 20 15 12 34 41 Ig 30 —.09 * Significant at .05 level ** Significant at .01 level TABLE 6. Correlations between J. osteosperma diseases and insects and precipitation (p) and the four different time quarters (Q). Average PQ PQ: PQ; PQ, PQ, PQs PQ; PQs annual Pathogen High High High High Low Low Low Low _ precipitation Needle blight =, (1 —.38 .02 —.09 —.26 10 .68** —.00 —.46 Needle cast —.00 22, —.21 .34 .39 —.19 10 =i = 5941/ Tip dieback —.44 —.13 —.20 —.29 —.34 .O1 .04 9 =) Mold-mildew —.19 —.31 .02 .08* —.05 —.05 BAT — .29 —.09 Galls-foliage .80** .34 31 fOTE* .30 12 —.10 lll soe Galls-branch .40 —.15 .67** 40 —.21 —.13 14 .09 .oo* Witch's broom 5 —.08 1642 OO =.05 .04 25 .05 Eons Lesions-foliage —.03 =, 112) —.02 £32 .49* —.04 18 =o) == 15) Lesions-branch —.09 = .52* 25 RATE —.10 —.44 .34 ll .06 Rot-twigs —.42 —.51* .05 13 —.02 opal 42 0) Oot Rot-trunk = 29 —.38 —.14 43 02 —.20 18 —.14 = All Mistletoe —.03 = 2) .08 .29 roll —.29 .03 —.31 —.07 Insect borers —.89** =o OF —.03 = 5 —.63** —.40 .46 Ol =, Scale insects —.38 —.18 223 —.09 —.18 19 161 eal) Sai Insect galls —.02 oA AT SS —.12 —.04 —.39 16 —.03 * Significant at .05 level ** Significant at .01 level decayed areas and fruiting bodies. Only tenta- tive identifications of the species were made. Fourteen sites contained trees with wood rot (2-64 trees/transect). The incidence of unidentified fungal diseases on the foliage (mold-mildew) was very similar (Figs. 7, 8) to that of the wood-rotting fungi. These mold- mildew fungi may be similar to those recorded by others (Hodges 1962, USDA In- dex of Plant Diseases 1960, Horst 1979), i.e., Cercospora spp., Stigmina spp., Phoma spp., Dimerium spp., etc. We observed the symp- toms of leaf-tip burn, shoot-tip dieback, leaf- needle blight, and needle cast but did not establish specific causal agents for these dis- eases. Distinct negative correlations existed be- tween the amount of mold-mildew occurring in a site and temperatures at the site (a = 0.05 with spring and summer temperatures) and definite positive correlations between the amount of mold-mildew and the amounts of summer (a = 0.01) and fall (a = 0.05) precipi- tation. This corresponds to behavior of foliage pathogens (such as phomopsis, cercospora blight, etc.) in which moderate temperatures 434 Wood Rotting Fungi Percent of infected trees 12 3. 8 5 68.7 8 © Sites Mold Mildew 1) Wade Teh ch IS UG) V7 Percent of infected trees 1 2 6 OG Bo 7 OG O- 10 W-de Aga vS IG We Sites 80 Mistletoe Percent of infected trees Ue Gus ty G7 8 OOS 1248-14 16-78 ar Sites Fig. 5. The percent of trees (Juniperus osteosperma) infected with wood-rotting fungi (top), mold mildew (middle), and mistletoe (bottom). and high amounts of relative humidity and/or free moisture favor infection. Incidence of wood-rotting fungi seems to be correlated with low winter temperatures (a = 0.05), and no correlations exist with amounts of precipi- tation. There is, however, high correlation with the incidence of wood-rotting fungi in the host and variations in amounts of soil nu- trients. If the nitrogen supply to the host is relatively low, the frequency of wood-rotting fungi on the host is greater (a = 0.05). GREAT BASIN NATURALIST Vol. 46, No. 3 Wood Rotting Fungi Severity index ee eh ee O38 77 8) Sites We Mil We We Wes 1S WS 17 Mold Mildew Severity index YW 28s) oh G& 7 Bg Sites VO Wak ve ale) 2b YS WG-7 Mistletoe Severity index 1 2°83) 2 8-6. 7. oO Sites 10 11 12 13 14 15 16 17 Fig. 6. The severity index of wood-rotting fungi (top), mold mildew (middle), and mistletoe (bottom) on Junipe- rus osteosperma trees. Parasitic Higher Plants True mistletoe (Phoradendron juniperium) Englm. has been reported on Utah juniper in Utah by Hawksworth and Wiens (1966) and in Arizona by Hreha (1978). We found Phora- dendron juniperium in seven of our transects (Figs. 5, 6). Injuries of Unknown Origin A third disease category was designated as injuries of unknown origin. Diseases in this July 1986 90 Needle Blight Percent of infected trees on lo} 30 20 10 0 | 12 9 2 5 GG 728-9 JOU) 12 1d WAS AlGaly Sites 90 Tip Dieback Percent of infected trees 10 11 12 13 14 15 16 17 12989 4&4 6 6 7 8-6 Sites Needle Cast Percent of infected trees uo oO 1 2 38 4 6 Bw 2-8-9 Sites VO V1 12 1 V2 1S Wey Uz Fig. 7. The percent of infected trees (Juniperus os- teosperma) with needle blight (top), tip dieback (middle), and needle cast (bottom). category had symptoms similar to the tip- burn, tip-dieback, needle blight, needle cast, and general chlorosis (Figs. 5, 6, 7, 8). Envi- ronmental factors such as desiccation, winter injury, high temperature, and nutrient imbal- ances could cause these injuries. Since some could also be induced by biotic agents, how- ever, we classified them as being of unknown origin. Isolates from these tissues produced a BUNDERSON ET AL.: JUNIPER DISEASES 435 14 Needle Blight Severity index @o o 1 2 8 4 65-6. 7 8 9 Sites VO Vd TW WAS We a7 Tip Dieback Severity index -— fo?) @ ny io 2s) - G-B @ 7s oof Sites 1 V4 We We We 1S. WG: ay Needle Cast Severity index 1 2 Gt SG. 7 BQ Sites NOTA 2S 14s AS) A617, Fig. 8. The severity index of three diseases: needle blight (top), tip dieback (middle), and needle cast (bot- tom) on Juniperus osteosperma trees. variety of fungi, most of which were probably saprophytes. Although a variety of fungi were isolated from these tissues, their pathogenic- ity was not established. Consequently, no specific cause was assigned for these diseases. It was possible, however, to separate in- juries into three groups (Figs. 7, 8) as was done earlier by Horst (1979). Symptoms of needle blight and tip dieback occurred at ev- ery site, and needle-cast was observed on ap- proximately one-third of the sites. 436 GREAT BASIN NATURALIST Vol. 46, No. 3 TABLE 7. Correlations between J. osteosperma diseases and insects versus soil variables. Pathogens pH ECx10° JN ppm P ppm K ppm Ca Needle blight 31 Roce 30 = Il .20 = Needle cast oll =, 119) —.10 —.24 —.01 oO Tip dieback Ol .26 —.08 .66** 43 —.04 Mold-mildew .09 01 Aes =P .28 08 Galls-foliage =.) —.34 07 =5119 —.39 41 Galls-branch = Mil —.13 —.33 = —.34 BOT Witch's broom —.15 —.18 =.43 —.18 —.20 7 Lesions-foliage .49* —.07 ollie =o 117 12 .02 Lesions-branch 15 —.08 Oat = AT* —.03 —.02 Rot-twig .26 ll 52} —.38 07 —.03 Rot-trunk .20 —.08 = =) .48* —.02 Mistletoe = 08} =) 20 —.33 ll —.19 Insect borers .29 .24 62** —.18 .49* = 99) Scale insects 23 .04* 16 = 22 —.03 —.34 Insect galls 30 —.24 18 56* 44 31 * Significant at .05 level ** Significant at .01 level The incidence of needle blight was the most responsive of the three diseases to variations in the environmental factors we measured. Incidence was consistently negatively corre- lated (a = 0.05) with sites that had low tem- perature periods throughout the year, and particularly for the low winter temperatures (a = 0.01). The correlation with precipitation was similar. Needle blight was more intense in those sites with the lowest winter precipita- tion (a = 0.05). Incidence of needle blight was positively correlated with summer drought (a = 0.01). These results suggest that environ- mental stress contributes significantly to this symptom, and it may be the result of desicca- tion or winter injury. The pinyon-juniper woodlands rely heavily on winter precipita- tion to replenish soil moisture since precipita- tion in other periods of the year is sporadic. In areas where soil reserves are marginal, mois- ture deficits during hot summer months may produce the symptoms we observed. The incidence of needle blight was also cor- related with two soil factors (Table 7). Sites with a higher soil salinity as measured by the electrical conductivity (EC) had higher per- centages of trees with needle blight. That this may be a direct response is supported by pre- vious work (Bunderson 1983) in which small elevations in the amount of EC in the soil were correlated with lower concentrations of total chlorophyll in the foliage of Utah ju- niper. The amount of total soluble carbohy- drate in the foliage increased with higher EC in the soil, which may indicate a decrease in the metabolic efficiency of the plant. The cor- relation (a = 0.05) of needle blight with the amount of calcium in the soil indicates that, when calcium is relatively plentiful, the per- cent of needle blight on the site is lower, and we hypothesize that as soil calcium content increases, the detrimental effect of salinity on Utah juniper is ameliorated. Tip dieback, while also having a moderate negative correlation with low winter precipi- tation, has no correlation with temperature. Contrary to expectations, tip dieback was more common on trees with higher nitrogen levels in the foliage (a = 0.05) and higher phosphorus (a = 0.01) and potassium (a = 0.10) levels in the soil. Until fungi can be associated with these symptoms and patho- genicity established, relationships to other factors remain speculative. Gymnosporangium Rusts The most frequent juniper pathogens we encountered on our research sites were the Gymnosporangium rust fungi. These rust fungi are parasitic and most are heteroecious. The alternate hosts of G. inconspicuum and G. kernianum are species of Amelanchier , most commonly A. utahensis Koehne. (A. al- nifolia Nutt. is also a frequent alternate host for Gymnosporangium rusts, although usually for those species that parasitize Rocky Moun- tain juniper (Juniperus scopulorum (Sarg.) rather than J. osteosperma). Gymnospo- rangium nelsoni has been found on Ame- lanchier and, locally, on Peraphyllum ramos- July 1986 BUNDERSON ET AL.: JUNIPER DISEASES 437 ppm Mg ppm Na ppm SO-S ppm Fe % Sand % Silt % Clay —.06 —.25 .24 olka oA —.24 —.20 .23 .24 07 — .28 —.04 —.09 .26 Bi = PA BOK 29 = 5B) = 0 lal 18 —.01 .00 =P) 04 —.06 .O7 09 .45 —.49* — .28 .48* 42 cSill —.02 37 —.24 —.04 39 Ol .20 16 —.09 —.45 —.09 04* oOAU .35 .20 —.16 .24 —:04 —.24 = All 18 13 —.05 —.30 —.48* .46 .05 wll) —.04 .25 45 ll —.34 aoe —.08 31 —.47* = OP — .35 .26 mls a2 —.38 13 —.11 = PAT .23 = 0H) —.48* .O1 —.21 18 —.03 —.03 =O) = Al) —.30 .28 45 .26 —.43 —.40 =, oll .06* —.13 —.30 —.04 .22 30 oo sissimum Nutt. The alternate hosts of G. spe- ciosum have been reported on species of Fendlera and Philadelphus (Kern 1973). In our transects the incidence of rust on juniper was not significantly correlated with the frequency and close proximity of the alter- nate host. This was not unexpected consider- ing the differing ecological habitats of Ame- lanchier and Juniperus species. Since basid- iospores are capable of traveling considerable distances and remaining viable (Parmelee 1968), and because aeciospores may remain viable as long as one year (Miller 1932), alter- nate hosts need not to be in close proximity for infection of either to occur. Temperature and moisture requirements for the release of Gymnosporangium ba- sidiospores have been well documented by Pearson et al. (1980), Pady and Kramer (1971), and Pearson et al. (1977). Although little is known about the specific requirements for infection of juniper, it is obvious that these requirements were met in our research sites (Table 8). Gymnosporangium speciosum and G. kernianum were less widespread than ei- ther G. inconspicuum or G. nelsoni. Peterson (1967) noted that G. kernianum is absent from many stands throughout the Great Basin where both hosts occur together. Our data support this observation. A Predictive Model Disease occurrences are determined by complex environmental interactions and causal agents. In the present study we have used descriptive data to develop hypotheses for correlating the incidence of disease with climate, soil, and other site factors in Utah juniper habitats. Distributions of several of the disease vari- ables were sufficiently peaked or had such high degrees of skewness that the populations did not fit the normal distributions that are implicit in correlational or factor analyses. Even though the correlations in our study are unusually stable because of the size of our data base (96 trees in each of 17 sites), nonparamet- ric methods of analysis were used to generate predictive models from our data. We applied the weighted least squares analysis for categorical data (Forthofer and Lehnen 1981). Using the environmental vari- ables that generally encourage the growth and proliferation of fungi causing mold and/or mildew symptoms, the following hypotheses were generated: 1. High amounts of fall precipitation characterize sites with high amounts of mold-mildew on the foliage. Low summer temperatures characterize sites with high amounts of mold-mildew on the foliage. High nitrogen nutritional status of the plant is condu- cive to low amounts of mold-mildew on the foliage. 2, 3. Five of the 17 sites were selected, each of which contained trees infected with a mold/ mildew pathogen. Each of these sites also dif- fered from the others on the three environ- mental variables chosen to formulate the three hypotheses. The hypotheses can be re- flected as linear weights where: 438 GREAT BASIN NATURALIST TABLE 8. Correlations between J. osteosperma diseases and insects and site characteristics. : —.13 24 pathogen Distance Height Needle blight —.45 HAG) Needle cast .54* — 2] Tip dieback = 8} = a8!) Mold/mildew — 17 —.02 Galls-foliage 43 .66** Galls-branch 16 .48 Witch’s broom 19 .47* Lesions-foliage se —.4] Lesions-branch — 29 4] Rot-twig 13 aT Rot-trunk — 27 .09 Mistletoe —.4] .20 Borers =o =U Scale insects — 95 —.63** Insect galls 02 mill * Significant at .05 level ** Significant at .01 level 1 = The hypothesis is true for this site. —] = The hypothesis is not true for this site. 0 = This site was not considered when making this hypothesis. These weights were entered into the fol- lowing X matrix where row | is equivalent to site 5, row 2 to site 6, row 3 to site 9, row 4 to site 14, and row 5 to site 15. X Matrix: Row 1 1 0 1 1 Row 2 1 0 1 0 Row 3 1 1 IL 0 Row 4 1 0 =I =I Row 5 1 —] =Il 1 Column | represents the presence or absence of mold/mildew at each of the selected sites. Since each site was selected because mold/ mildew was present, column 1 is a constant. Column 2 represents the linear weights of hy- pothesis 1; column 3, hypothesis 2; and column 4, hypothesis 3. From this matrix of linear weights based on our hypotheses, we have pre- dicted the incidence of mold-mildew at sites 5, 6, 9, 14, and 15. The predicted and observed occur- rences of mold/mildew expressed as proportions of infected trees are presented below. From the small residual figures, the success of the three hypotheses at predicting the proportion of infec- tion seems good. Site Observed Predicted Residual 5 .073 .O74 —.001 6 042 .041 .0O1 9 .229 .228 .0O1 14 .375 RON all —.002 15 .625 .624 001 Vol. 46, No. 3 Soil Diameter Age depth —=:38 .20 .08 —.33 = Ils} 12 —.30 21 21 — 34 05 —.26 .24 —.10 5315) —.02 22 15 .07 06 —.05 —.03* .28 .03* —.02 —.14 .10 = 55 BOS .69** —.03 —.16 —.02 .08 01 .0o* —.30 = a7 SRO Oise = ou 11 —.05 19 The overall chi’ (X’) goodness of fit for the observed versus predicted data was 0.004, and the beta weight for the matrix constant (column 1) was 0.226. The beta weights for each of the hypotheses were — 0.181, — 0.184, and 0.033. The very small chi”s and the small residual values indicate an excellent fit for the hypothetical model to the observed incidence of mold-mildew. The hypotheses were tested for significance of the X° values: Hypothesis 1 = 27.7 Hypothesis 2 = 126.3 Hypothesis3 = 1.6 Significance of X° values: .05 level 3.84 .01 level 6.63 .001 level 10.83 Using this procedure we can affirm that high amounts of fall precipitation and low summer temperatures do indeed characterize sites with high amounts of mold-mildew on Utah juniper foliage. The first two hypotheses of our model have reliably predicted the pro- portion of trees in a pinyon-juniper site that could be expected to be infected with mold- mildew. The third hypothesis was not con- firmed. As can be seen from the predictive ability of the generated hypotheses and the power to statistically confirm their validity, such a non- parametric statistical model offers a valuable tool for understanding the interactions be- tween the many variables that exist in any ecosystem. July 1986 BUNDERSON ET AL.: JUNIPER DISEASES 439 Alternate Vigor Decadence Density Slope Elevation Exposure host .38 422) 2 = PA 533 — .20 —.09 —.01 .09 = 08 —.20 —.03 .05 54D) .36 .04 m0) .05 mle == 605) —.05 —.07 .64** .08 .05 nO Dit =o) .34 —.56** 16 —.31 OL 04 = il) Atl —.45 .16 —.16 19 .o4* —.06 ONS —.67** .do* —.19 06 .49* 04 739 .02 .05 —.14 —.14 olka sell .39 —.40 Do 35) 08 20055 olla Boll .29 .03 —.19 00 a5 —.30 sls} —.34 .89** atl .02 34 .06 ROS —.07 10 .08* .05 seid Ol 44 .30 .33 45 18 Roles =, 19 mil .60** —.40 .20 —.14 44 = oll apa 16 =e) —.18 48* —.19 .09 —.08 LITERATURE CITED DAUBENMIRE, R. F. 1943. Vegetational zonation in the Rocky ALDWINKLE, H. S. 1975. Field susceptibility of 41 apple cultivars to cedar apple rust and quince rust. Plant Dis. Rep. 58: 696-699. ALLRED, B. W. 1964. Problems and opportunities on U.S. grasslands. Amer. Hereford J. 54: 70-72., 132 p. ARTHUR, J. C. 1934. Manual of the rusts in the United States and Canada. Purdue Res. Found., Lafayette. 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Observations on witch’s broom formation autoparasitism and new hosts in Phoradendron. Madrono 18: 218-224. Heptinc, G. H. 1971. Diseases of forest and shade trees of the United States. U.S. Dept. Agric. Handbook 386. 658 pp: Hopces, C. §. 1962. Comparison of four similar fungi from Juniperus and related conifers. Mycologia 54: 62-69. Horst, R. K. 1979. Westcott’s plant disease handbook. Van Nostrand Reinhold Co., New York. 803 pp. Hrena, A. M. 1978. A comparative distribution of two mistle- toes: Arceuthobrium divaricatum and Phoranden- dron juniperinum (South Rim, Grand Canyon Na- tional Park, Arizona). Unpublished thesis, Brigham Young University, Provo, Utah. Hunt, C. B. 1974. Natural regions of the United States and Canada. Freeman, San Francisco. Hurst, W. D. 1977. Managing pinyon-juniper for multiple benefits. In Ecology, uses, and management of pinyon-juniper woodlands. USDA For. Serv. Gen. Tech. Rep. RM-39. 48 pp. KERN, 1973. A revised taxonomic account of Gymnospo- rangium. 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Madrono 19: 79-91. PHILLirs, E. A. 1959. Methods of vegetation study. Holt, Reinhart and Winston, Inc., New York. 107 pp. TUELLER, P. T., C. D. BEESON, R. J. Tauscu, N. E. WEST, AND K. H. REA. 1979. Pinyon-juniper woodlands of the Great Basin: distribution, flora, vegetal cover. USDA For. Serv. Res. Pap. INT-229. 22 pp. VASEK, F.C. 1966. The distribution and taxonomy of three western junipers. Brittonia 18: 350-372. WEIMER, J. L. 1917. Three cedar rust fungi, their life histories and the disease they produce. Cornell Univ. Agric. Expt. Stn. Bull. 390: 523-524. West, N. E., K. H. REA, AND R. J. TAuscuH. 1975. Basic synecological relationships in pinyon-juniper. Pages 41-52 in The pinyon-juniper ecosystem: a symposium. Utah Agric. Expt. Sta. Wooppury, A. M. 1947. Distribution of pigmy conifers in Utah and northeastern Arizona. Ecology 28: 113-126. STATUS AND DISTRIBUTION OF THE FISH CREEK SPRINGS TUI CHUB, GILA BICOLOR EUCHILA Thomas M. Baugh’, John W. Pedretti’, and James E. Deacon” ABSTRACT. —The Fish Creek Springs tui chub, Gila bicolor euchila, is present in large numbers throughout its native habitat in spite of extensive man-caused habitat disturbance. This subspecies occurs further downstream in Fish Creek than previously reported. The Fish Creek Springs tui chub Gila bicolor euchila was described by Hubbs and Miller (1972) as an endemic subspecies restricted to the springs and outflows of Fish Creek Springs in southeastern Eureka County, Nevada, in R.53E., T. 16N., S. 8 and 9 (USGS Bellevue Peak Quadrangle 1956). Subsequently, Hardy (1979a, b) found this subspecies in only one spring and failed to find any in the outflows. In 1983 (Federal Register 1984), the fish was re- ported from three springs, but not from the out- flows, and was proposed for listing as “threatened” with critical habitat designated un- der 50 Code of Federal Regulations Part 17. METHODS AND MATERIALS Parts of the present habitat of G. b. euchila were sampled on 9-10 July 1984 and 26-27 June 1985. Unbaited 6-mm wire mesh, Gee- type minnow traps were used. Catch and to- tal-length data are presented in Table 1. Some physical and chemical characteristics of this system are reported in Table 2. Temperature was measured with a standard mercury bulb thermometer, and pH was measured with a Hach High Range pH test Cube No. 12519- | 00. Conductivity was measured with a Lab- | Line Lectro MHO-Meter, Model MC-1, Mark IV and is reported in Micromhos/cm. In | 1985 dissolved oxygen was measured with a ’ Hach Portable Dissolved Oxygen Meter. RESULTS | The Fish Creek Springs system is com- | posed of two isolated northern spring-pools and a number of other springs connected by channelized outflows (Fig. 1, Sites 13-14). In 1938, and at least until 1956, the two northern springs were connected by outflow to the re- mainder of the system (Hubbs et al. 1974). The combined outflows form Fish Creek, which passes just south of Fish Creek Ranch and termi- nates in a reservoir about | km east of Nevada State Route 20. Man-caused modification of this system has occurred frequently in the past. For example, Hubbs et al. (1974) report that in 1938 Fish Creek was dry where it crossed Nevada State Route 20 because of water consumption on Fish Creek Ranch. Prior to our 1984 survey, a dragline had been used to clear Fish Creek throughout sections 9 and 10. In 1984 the westernmost spring (map reference 1) was little more than a muddy swale with a diameter of about 15 m and an area of dense rushes (Juncus sp.) in the middle. Water, however, continued to flow from this spring into an outflow stream about 300 m long (map refer- ence 2) into the main body of the aquatic system. By 1985 the spring-pool had been cleaned of emergent aquatic vegetation and deepened and was, essentially, an open body of water. By 1985 heavy equipment had been used to dredge al- most all the spring-pools and their outflow chan- nels. Emergent aquatic vegetation was reduced throughout the system by about 75% compared to 1984. Dredged material was placed in piles alongside most of the spring-pools and outflows. Gila bicolor euchila presently occurs in large numbers in most components of this system (Table 1). In addition, it was found in 1This work was accomplished when the author was with the Department of Biological Sciences, University of Nevada, Las Vegas. Present address: U.S. Fish } and Wildlife Service, Office of Endangered Species, 1000 N. Glebe Road, Arlington, Virginia 22201. | "Department of Biological Sciences, University of Nevada, 4505 Maryland Parkway, Las Vegas, Nevada 89154. 44] 442 GREAT BASIN NATURALIST Vol. 46, No. 3 TaBLE 1. Catch and total length (TL) data for Gila bicolor euchila from Fish Creek Springs, 1984—1985. Numbers in parentheses represent the number of fish measured. Map # Trap-hours Fish caught 1984 Dy 15 64 3 4 1 5 2 76 6 1 30 9 1 28 10 1 19 13 9 120 14 3 0 15 14 8 16 10 4 1985 2 6 86 5 2D; 126 6 ») 106 13 17.5 eel 1Sites 5 and 6 combined. x Total [(N) = fish measured ] length—mm Catch per trap-hour Total length range—mm 4.3 (64) 50.4 31— 64 0.3 105.0 nae 38 (50) 46.3 32- 78 30 (30) 63.0 44— 78 28 (28) 54.5 38— 68 19 (17) 54.9 ABE iA 13.3 (75) 64.0 38-113 0 0 0 6 (8) 74.3 40-122 4 (4) 61.8 45— 84 14.3 (50) 59.9 AM Ti 63.0 (50) 64.9! 46-122! 53.0 g % 1588 (100) 77.3 33-118 TABLE 2. Physical and chemical characteristics of the Fish Creek Springs system, 1984-1985. Area Temperature Map #_ type C 1984 2 outflow-stream — 3 spring-pool 22 4 spring-pool a 5 spring-pool — 9 spring-pool a 10 outflow-stream _ 13 spring-pool PAIL) 14 spring-pool 26.0 15 canal 11.0 1985 2 outflow-stream 23.8 5 spring-pool Mie 13 spring-pool 16.2 the outflow of Fish Creek Springs in the southern portion of section 10 closely adjacent to the Fish Creek Ranch headquarters. This is beyond the range reported for this subspecies by Hubbs et al. (1974) and in the Federal Register (1984). In 1984 this subspecies was also taken in section 12 at the intersection of the outflow with Nevada State Route 20. Fish were also seen but not captured about 50 m downstream from this intersection in 1985. This area is about 4.8 km east of the previous Conductivity Dissolved oxygen pH micromhos ppm 8.0 505 — 7.5 500 — = 590 — 7.0 — — 7.0 530 — 7.9 540 — 8.0 570 — 8.0 094 —_— 8.5 420 — 8.0 618 15.4 7.0 617 6.3 7.6 557 8.5 easternmost record for this subspecies (Fed- eral Register 1984). Visual inspection of the remainder of Fish Creek down to and includ- ing the reservoir revealed no fish. In 1984 a total of 350 fish were captured from 10 trap sites beginning in section 8 and terminating in section 12. Of the fish cap- tured, 276 were measured (Table 1). In 1985, 595 fish were captured from four sites, and | 200 of these fish were measured (Table 1). Thousands of fish ranging from about 5 mm July 1986 13 14 0 BAUGH ET AL.: TUICHUB 443 10 P] Fig. 1. General configuration of Fish Creek Springs and outflow. Numbers indicate the location of various components of the system within section 8. Sampling site 15 is in section 10, and sampling site 16 is in section 12; neither are shown here. Elevation of the westernmost spring (site 1) is 6,040 ft. It and the northernmost spring (site 13) joined near site 3 on the Bellevue Peak Quadrangle (1956) in 1938. (TL) to more than about 150 mm were seen during visual inspections of the spring system in both 1984 and 1985. No other fish species were seen or trapped in either 1984 or 1985; however, Hubbs et al. (1974) report that rancher Isador Sara planted both rainbow trout (Salmo gairdneri) and brook trout (Salvelinus fontinalis) as early as about 1934, and Pat Coffin (Nevada Department of Wildlife, personal communication) reports that rainbow trout (Salmo gairdneri) are planted in the springs each spring. The Fed- 444 eral Register (1986) states that 7,000 “catch- able-sized” brown trout, Salmo trutta, were planted in Fish Creek Springs in 1973, 1976-1978, and 1981 and notes that the in- crease in tui chub numbers ‘corresponds with the cessation of brown trout stocking in the springheads . Trap results and length measurement data for map reference areas 2, 3, 6, 10, 13, 15, and 16 (1984) and 2, 5, 6, and 13 (1985) are pre- sented in Table 1. Catch per trap-hour ranged from .3 to 38 in 1984 and 14.3 to 63.0 in 1985. The total length (TL) range from all sites in which fish were taken in 1984 was from 31 mm to 122 mm (x = 54.8) and in 1985 from 33 to 122 mm (x = 69.8). Catch per trap-hour was greater at all the sites sampled in 1985 than at the same sites sampled in 1984. Table 2 presents some of the physical and chemical characteristics of the areas referred to in Figure 1. Water temperature ranged from 11 to 26 C, pH from 7.0 to 8.5, conduc- tivity from 420 to 680 micromhos, and dis- solved oxygen from 6.3 to 15.4 ppm. SUMMARY The aquatic system at Fish Creek Springs is composed of a number of springs and their outflows. Human modification of the site in- cludes dredging of the spring-pools and their outflow channels, cattle grazing, and intro- ductions of trout. Even with extensive modifi- cation of the natural system, Gila bicolor eu- chila is present in large numbers in most components of the system. The presence of GREAT BASIN NATURALIST Vol. 46, No. 3 these fish in sections 10 and 12 extends the range of this subspecies about 4.8 km east of localities previously reported. Fish numbers appear to have increased following 1981 when Salmo trutta was last planted in the Fish Creek Springs system. ACKNOWLEDGMENTS We thank the U.S. Fish and Wildlife Ser- vice and the Nevada Department of Wildlife for their financial support of this project. Su- zanne Pedretti assisted in the field during the 1985 survey. LITERATURE CITED FEDERAL REGISTER. 1984. Endangered and threatened wildlife and plants; proposed threatened status and critical habitat for the Fish Creek Springs tui chub (Gila bicolor euchila). Fed. Reg. 49(110): 23409-23412. FEDERAL REGISTER. 1986. Endangered and threatened Wildlife and plants; notice of withdrawal of pro- posed rule to list the Fish Creek Springs tui chub as threatened. Fed. Reg: 51(46): 8215-8216. Harpy, T. 1980a. Interbasin report to the Desert Fishes Council, 1978. Proc. Desert Fish. Council 10: 68-70. —___. 1980b. Interbasin report to the Desert Fishes Council, 1979. Proc. Desert Fish. Council 11: 5-21. Husps, C. L., AND R. R. MILLER. 1972. Diagnoses of new cyprinid fishes of isolated waters in the Great Basin of western North America. San Diego Soc. Nat. Hist. Trans. 17(8): 101-106. Husps, C. L., R. R. MILLER, AND L. C. Huss. 1974. Hy- drographic history and relict fishes of the north- central Great Basin. Memoirs of the California Academy of Sciences 7: 1-259. PONDEROSA PINE CONELET AND CONE MORTALITY IN CENTRAL ARIZONA J. M. Schmid", J. C. Mitchell!, and S. A. Mata! ABSTRACT.—Ponderosa pine conelets in 10 stands on the Coconino and Kaibab national forests were observed periodically from July 1982 until they matured in September 1983. Abortion, ponderosa pine cone beetles (Conophtho- rus ponderosae Hopkins), and ponderosa pine coneworms (Dioryctria sp., probably Auranticella [Grote]) were the significant mortality factors. Cattle, tip moths, and squirrels rarely destroyed conelets or cones. Good cone crops in stands of ponderosa pine, Pinus ponderosa Lawson, in the South- west occur on the average of every three to four years (Schubert 1974), with large crops about every five years (Larson and Schubert 1970). Because good cone crops occur infre- quently and natural regeneration is difficult to obtain, any factor adversely influencing cone production, and therefore seed production, needs to be identified. Cone mortality is caused by insects, small mammals, and weather factors (Schubert 1974). Cone beetles (Conophthorus) have occasionally destroyed 50% of the cone crop in some locales (Pearson 1950). Mice, chipmunks, and ground squir- rels may eat seeds but are not important while the cones are attached to the branches. Abert squirrels, Sciurus aberti Woodhouse, on the other hand, destroyed an average of 20% of the cone crop over a 10-year period (Larson and Schubert 1970). Freezing temperatures in June have also killed conelets (Schubert 1974). This paper reports on the conelet and cone mortality caused by different factors during a two-year period. METHODS Ten cone-producing areas were selected in ponderosa pine stands on the Coconino and Kaibab national forests in north central Ari- zona (Fig. 1). For an area to be selected, it had ‘to have 10 trees bearing conelets. In most areas, each tree usually bore more than 30 -conelets, but only 30 conelets were randomly selected for study. In one area, Dutch Kid, | each tree had less than 15 conelets, so only 10 O\Highway 89 Hart Prairie\ 9 Williams 5Devil Dog ODutch Kid a Mormon Lake Fig. 1. Location of study areas on the Coconino and Kaibab national forests, Arizona. were selected. Two other areas, Devil Dog and Ten-X, had individual trees with fewer than 30 available cones, so the desired num- ber of 300 conelets per area was not attained. Table 1 shows the total number of conelets observed per area. The conelets were generally on the lower crown branches of large-diameter, open- grown trees. The conelets were randomly se- lected from those within 4 m of ground level because no equipment was available to exam- ine cones above that height. Conelets were present individually and in clusters of two to 1USDA Forest Service, Rocky Mountain Forest and Range Experiment Station, Fort Collins, Colorado 80526-2098. | | | | 445 446 TABLE 1. Number of ponderosa pine cones observed in each area, Arizona 1982—1983. National forest/Area Number of cones Elevation m Coconino National Forest Cable 302 2,226 Deadman Flat 301 2,290 Hart Prairie 301 2,561 Highway 89 301 2,195 Rockledge 300 2,195 Schnebly Hill 300 1,980 Kaibab National Forest Devil Dog 281 2,012 Dutch Kid 100 2,012 Grandview 301 2,256 Ten-X 291 2,025 five. Their cone cluster class was recorded to determine whether insect damage varied with the number of cones per cluster. The conelets were selected in July 1982 and were examined about every two weeks there- after until September 1982. Observations were discontinued during the winter of 1982-1983 but were begun again in March or April 1983, depending on accessibility of the site. In August-September 1983 observations were terminated and the cones collected for additional study. During each observation pe- riod, factors causing damage or mortality were identified when possible. Some of the areas were the same as those used by Schmid et al. (1984) in an earlier study, in which 32 cones were collected from the lower and middle crowns of large-diame- ter, open-grown trees in September 1982. However, in that study cones were not ob- served from the time they were conelets. RESULTS AND DISCUSSION Abortion, ponderosa pine cone _ beetles (Conophthorus ponderosae Hopkins), and ponderosa pine coneworms (Dioryctria sp., probably D. auranticella Grote) were the three most important mortality factors, with abortion consistently causing the greatest amount of conelet mortality (Table 2). Abor- tion, as used herein, encompasses several mortality factors. Conelets died and failed to mature, but the exact cause of their death was undetermined. Many of the conelets died the first summer, which suggests some physiolog- GREAT BASIN NATURALIST Vol. 46, No. 3 ical disfunction, i.e., inadequate pollination or water stress. The 1983 mortality was pri- marily observed when observations were re- sumed, which indicates that the mortality oc- curred during the winter months. Schubert (1974) and the USDA Forest Service (1974) note cold temperatures can be lethal, so much of the 1983 mortality may be attributed to freezing temperatures. In December 1982 and January and February 1983, tempera- tures dropped below 0 F in Flagstaff, but whether these low temperatures caused over- winter mortality was not determined. And finally, insects like seedbugs (Leptoglossus oc- cidentalis Heidemann) could cause conelet mortality, although neither the adults nor damage were observed. In the life of the cone, mortality factors during the first year are the least understood. Although Conophthorus-caused mortality was 10% or less in 7 of the 10 areas, Conoph- thorus beetles generally caused the second greatest amount of mortality, causing up to 62% mortality in one area (table 2). Conoph- thorus- caused mortality changed significantly between 1982 and 1983 in two locations and remained the same in two other areas. In 1982, Conophthorus-caused mortality aver- aged 39% in the Highway 89 area and 2% in the Ten-X area (Schmid et al. 1984). In 1983, the second year of this study, mortality at- tributable to Conophthorus was 1% in High- way 89 and 28% in Ten-X. Whereas the two studies have slightly different sampling de- signs, which may have created some of the difference in the incidence of mortality, enough difference exists to suggest that Conophthorus populations, and therefore Conophthorus-caused mortality, exhibit sig- nificant variation from year to year. The frequency of Conophthorus-killed cones mirrored the frequency of the cone cluster class for all levels of Conophthorus- caused mortality. When two-cone clusters were the most abundant cone _ cluster, Conophthorus damage was greatest in that | class. When three-cone clusters predomi- nated, damage was greatest in that class. Thus, even though female beetles can infest more than one cone (Kinzer et al. 1970), mul- tiple cone clusters do not apparently reduce beetle mortality during dispersal and host lo- cation nor do they increase the probability of — ~ a July 1986 SCHMID ET AL.: PONDEROSA PINE CONES 447 TABLE 2. Percent mortality* of developing ponderosa pine cones by location and mortality factors, Arizona 1982-1983. Mortality Coconino National Forest Kaibab National Forest factor Cable Deadman Hart Highway Rockledge Schnebly Devil Dutch Grandview Ten-X Flat Prairie 89 Hill Dog Kid Abortion 1982 13 15 10 6 5 8 13 10 8 2] 1983 15 4 a cae ali 22 15 14 _8 13 Total 28 19 17 13 22, 30 28 24 16 34 Cattle 0 0 0 0 0 1 0 0 1 0 Conophthorus <1 14 62 1 3 1 2 a= a 80 60 40 Surviving (percent) 20 Hart Prairie ) Jul Sep Apr May Jun Jul Aug Sep 15 30 15 15 15 15 15 15 1982 1983 ----------------—-----------------+ > Fig. 2 Ponderosa pine conelet survival for three study areas on the Coconino and Kaibab national forests, Ari- zona, 1982-1983. their relative importance. Losses to Conoph- thorus and Dioryctria can be drastically re- duced using chemical insecticides. If an area GREAT BASIN NATURALIST Vol. 46, No. 3 produces a good conelet crop and seed is needed from it, a preventive spray might be applied in May or early June of the year the cones mature to keep these cone beetles and coneworms from infesting the cones. LITERATURE CITED KInzER, H. G., B. J. RIDGILL, AND J. G. Watts. 1970. Biol ogy and cone attack behavior on Conophthorus ponderosae in southern New Mexico (Coleoptera: Scolytidae). Annals Entom. Soc. Amer. 63: 795-798. Larson, M. M., AND G. H. ScHuBERT. 1970. Cone crops of ponderosa pine in central Arizona, including the in- fluence of Abert squirrels. USDA For. Serv. Res. Pap. RM-58. Rocky Mtn. For. and Range Expt. Sta., Fort Collins, Colorado. 15 pp. PEARSON, G. A. 1950. Management of ponderosa pine in the Southwest. U.S. Dept. of Agric. Monograph 6. Wash- ington, D.C. 218 pp. SCHMID, J. M., J. C. MITCHELL, K. D. CarRLIN, AND M. R. Wac- NER. 1984. Insect damage, cone dimensions, and seed production in crown levels of ponderosa pine. Great Basin Nat. 44: 575-578. SCHUBERT, G. H. 1974. Silviculture of southwestern pon- derosa pine: the status of our knowledge. USDA For. Serv. Res. Pap. RM-123. Rocky Mtn. For. and Range Expt. Station, Fort Collins, Colorado. 71 pp. USDA Forest SERVICE. 1974. Seeds of woody plants in the United States. USDA For. Serv. Agric. Handbook 450. Washington, D.C. 883 pp. NUMBER AND CONDITION OF SEEDS IN PONDEROSA PINE CONES IN CENTRAL ARIZONA J. M. Schmid’, S. A. Mata’, and J. C. Mitchell! ABSTRACT.—Ponderosa pine cones from 10 areas in Arizona were collected prior to natural seed dispersal and dissected to determine the number of sound, hollow, and insect-damaged seeds in each cone. Total and sound seed yields per cone did not vary significantly among areas but did vary significantly among trees within each area. Numbers of hollow and Megastigmus-infested (Hymenoptera: Torymidae) seeds varied significantly among areas and trees within areas. Numbers of sound seed increased significantly with increasing cone length but did not change with increasing numbers of cones per cluster. The percentages of Megastigmus-infested seed did not change significantly with increasing cone length or number of cones per cluster. Good cone crops are the first step in obtaining natural regeneration in stands of ponderosa pine, Pinus ponderosa Douglas ex Lawson. Good cone crops occur every three to four years (Schubert 1974), with exceptional crops every five years (Larson and Schubert 1970). Good to exceptional cone crops, however, do not insure good to exceptional sound seed yield. Cone crops can be substantially destroyed by insects (Schmid et al. 1986), or seed within the cones can be damaged or destroyed so that sound seed yield is substantially lessened. Seed chalcids may destory 77% of the seed in some instances (Schmid et al. 1984), and other insects and physi- ological impairments may further reduce sound seed yield. This note reports on the total, sound, hollow, and insect-damaged seed from 10 areas in Arizona. METHODS Ten areas with first-year conelets were se- lected on the Coconino and Kaibab national forests in Arizona in July 1982 (Schmid et al. 1986). Trees with conelets were generally more than 18 inches dbh and 50 feet tall in 8 of the 10 areas. In the other 2 areas, trees were 8-12 inches dbh and 20—45 ft tall. On all sites the trees were open-grown, and stand density was less than 50 square feet of basal area per acre. Ten trees were selected in each area, and at least 30 conelets were tagged on each tree in 9 of the 10 areas. In the tenth area fewer than 15 conelets were available on each tree, so only 10 per tree were tagged. Conelets were present individually and in clusters of two to five. All tagged conelets were within 12 feet of ground level, so they could be observed from the ground or from a stepladder. The conelets were ob- served periodically during the summers of 1982 and 1983. In September 1983 all tagged live cones on each area were picked (Table 1) and taken to the laboratory. Each cone was mea- sured lengthwise, then dissected to determine the number of sound, hollow, insect-damaged, and total seeds. Sound seeds had white en- dosperm completely filling the seedcoat. Hollow seeds were without contents or had a shriveled remnant inside the seedcoat. Mean numbers of each seed category per cone were calculated for each area on a per tree basis. Numbers of each category per cone were tested by nested analysis of variance for significant dif- ferences among areas and among trees within areas, a = 0.05. Percentages of each seed cate- gory were also tested by analysis of variance for significant variation associated with cone length and the number of cones per cluster, a = 0.05. When significant differences were indicated, Tukey’s multiple range test was used to deter- mine which mean values were significantly dif- ferent from each other. RESULTS AND DISCUSSION Total Seeds Mean number of total seeds per cone per area ranged from 47 to 68 (Table 1) and was not USDA Forest Service, Rocky Mountain Forest and Range Experiment Station. 240 W. Prospect St., Fort Collins, Colorado 80526 449 450 GREAT BASIN NATURALIST Vol. 46, No. 3 TABLE 1. Number of cones dissected and mean number of total seeds per cone per tree within each area, Arizona 1983.1 National forest/Area Number of cones Coconino National Forest Cable 199 Deadman Flat 176 Hart Prairie 53 Highway 89 230 Rockledge 214 Schnebly Hill 140 Kaibab National Forest Devil Dog 173 Dutch Kid 66 Grandview 194 Ten-X 99 ‘Means followed by the same letter are not significantly different, a = 0.05. TABLE 2. Mean number of sound, hollow, and Megastigmus -infested seeds per cone per tree by area.' National forest/Area Sound Coconino National Forest Cable 49 + 23a Deadman Flat 40 + 13a Hart Prairie 58 + 19a Highway 89 50 + 17a Rockledge 46 + 17a Schnebly Hill 35 + lla Kaibab National Forest Devil Dog 42 + 20a Dutch Kid 38 + 15a Grandview 42+ 8a Ten -X 37 + 12a x +$.D Range 67 + 24a 21- 92 56 + 15a 29- 72 61 + 19a 41-100 68 + l6a 46- 89 56 + 16a 33- 83 52 + 14a 30- 72 59 + 22a 25- 94 56 + 15a 24- 74 47+ 8a 31- 56 50 + 14a 30- 68 Hollow Megastigmus x +S.D. 16+ 5ced 0.4 + 0.7a 6+ 2ab 8 +1 b ej} as. BA 0.2 + 0.5a 16 + 14cd Q- sD a 7+ 4abc 5.9 ED a 12 + Tabed 3 =e 8 a 16+ 8bcd 0.6+1 a 19 + lld Oa 4+ la 7 se 6h 12 + 3abcd <— lee alla ‘Means within the same column followed by the same letter are not significantly different, a = 0.05. significantly different among areas. The mean number of seeds per cone per area was gener- ally greater in 1983 than in 1982 for those areas common to an earlier report (Schmid et al. 1984) and this study. The mean number of total seeds per cone per tree varied significantly in all areas and accounts for the large variation in the range of mean number of seeds per area. Why some trees produce 20 to 30 seeds per cone whereas others produce 80 to 90 is unknown. Competi- tion between adjacent trees was not important because the trees were widely spaced and open-grown. Total seeds per cone increased with in- creasing cone length (r = 0.33). Sound Seeds Mean number of sound seeds per cone per tree was not significantly different among ar- eas (Table 2) but varied significantly among trees. The percentages of sound seed in- creased significantly with increasing cone length (r = 0.20) but did not change significantly with increasing numbers of cones per cluster when all areas were pooled. Because total seeds per cone did not vary significantly among areas, the non- significant variation in sound seeds was ex- pected. The increase in sound seed percentages with cone length and the nonsignificant change associated with increasing numbers of cones per cluster indicates that the percentage of sound seed does not decline even though trees produce larger and multiple cones per cluster. Cone length may thus serve as an indicator of sound seed, providing insect damage is negligible. As- suming insect-caused seed loss is negligible, longer cones should be selected over shorter cones because they have a greater probability of containing more sound seed. Hollow Seeds The mean number of hollow seeds per cone varied significantly among areas (Table 2) and July 1986 TABLE 3. Frequency of cones by cone length and by cones per cluster, all areas, Arizona 1982-1983. Cone Number length Frequency cones per Frequency (cm) (%) cluster (%) <6.0 11 1 11 6.0-9.0 80 2 43 >9.0 9 3 32 4 11 5 3 trees within areas. The percentage of hollow seeds decreased significantly with increasing cone length (r = 0.22) but did not change significantly with increasing numbers of cones per cluster when all areas were pooled. The relationships between the number or per- centages of hollow seeds and cone length and cones per cluster were the opposite of those for sound seeds. The greater percentage of hollow seeds as- sociated with shorter cone length suggests that some factor(s) causing poorer cone devel- opment might also be associated with greater hollow seed production. Cones that fail to develop to full size also fail to produce a higher percentage of sound seed. Most “hollow” seeds had a small shriveled remnant inside the seedcoat, although a small percentage were completely hollow. Whether these “hollow” seeds resulted from physiological disfunction or insect damage is not certain. Seedbugs were not observed in the areas, so their influence, if any, was minimal. Megastigmus -infested Seed Megastigmus -infested seed varied signifi- cantly among areas and among trees within areas, which agrees with the results of Schmid et al. (1984). In general, Megastigmus (Hy- menoptera: Torymidae) levels of damage were less in 1983 than in 1982. In four of five SCHMID ET AL.: PONDEROSA PINE SEEDS 451 areas common to both studies, mean numbers of Megastigmus -infested seed declined signfi- cantly in 1983. This indicates the substantial annual fluctuations occurring in the incidence of infestation, caused perhaps by substantial changes in Megastigmus population levels. The percentage of Megastigmus -infested seed remained about the same regardless of cone length or numbers of cones per cluster. Thus, Megastigmus damage did not have an important influence on the trend in the per- centage of sound seed associated with these two variables. Also, the presence of two or more cones in a cluster did not apparently induce females to oviposit more eggs such that the incidence of damage increased with an increasing number of cones per cluster. Cone Characteristics Eighty percent of the cones were 6—9 cm in length, with the remaining 20% of the cones about equally present as shorter and longer lengths (Table 3). Two-cone clusters were the most abundant, whereas five-cone clusters were relatively rare. LITERATURE CITED Larson, M. M., ANDG. H. SCHUBERT. 1970. Cone crops of ponderosa pine in central Arizona, including the influence of Abert squirrels. USDA For. Serv. Res. Pap. RM-58, 15 pp. Rocky Mtn. For. and Range Expt. Sta., Fort Collins, Colorado. SCHMID, J. M., J. C. MITCHELL, K. D. CARLIN, AND M. R. WAGNER. 1984. Insect damage, cone dimensions, and seed production in crown levels of ponderosa pine. Great Basin Nat. 44: 575-578. SCHMID, J. M., J.C. MITCHELL, AND S. A. Mata 1986. Pon- derosa pine conelet and cone mortality in Arizona. Great Basin Nat. 46: 445-448. SCHUBERT, G. L. 1974. Silviculture of southwestern pon- derosa pine: the status of our knowledge. USDA For. Serv. Res. Pap. RM-123, 71 pp. Rocky Mtn. For. and Range Expt. Sta., Fort Collins, Colo- rado. NEW SPECIES OF PROTOCEDROXYLON FROM THE UPPER JURASSIC OF BRITISH COLUMBIA, CANADA David A. Medlyn' and William D. Tidwell* ABSTRACT. —Protocedroxylon macgregorii sp. nov., from Jurassic strata of British Columbia, Canada, is the first reported occurrence of this genus in North America. Protocedroxylon macgregorii combines the tracheal pitting of the araucarians with the crossfield pitting of modern genera of the Abietineae. This species is similar to the type species Protocedroxylon araucarioides. They differ in that P. araucarioides has tangential pitting, tracheid septations, and entirely uniseriate rays. These features are lacking in P. macgregorii with the exception of the rays, which are partially biseriate in the latter species. Protocedroxylon macgregorii has traumatic resin canals or cysts that have not been reported in P. araucarioides. Protocedroxylon was proposed by Gothan (1910) for woods combining characters of abi- etineous and araucarian conifers. The type species Protocedroxylon araucarioides Gothan (1910) was described from Upper Jurassic strata near Esmarks Glacier of Spitzbergen. Tracheal pitting of P. araucarioides is consid- ered araucarioid, whereas its crossfield pitting is typically abietineous. Hence, the generic epithet of the type species refers to its abieti- neous characters, and the specific epithet im- plies araucarian affinities. Specimens of the petrified wood in this report were collected by D. C. McGregor of the Geological Survey of Canada from the northwest shoulder of an unnamed moun- tain situated about 2.5 km east of Elbow Mountain (across Graveyard Creek). The locality is at about 51°9’ N Latitude and 123°5’ W Longitude. The age is considered Lower Portlandian (Upper Jurassic) based upon the ammonite Buchia mosquensis, which occurs at several levels within the unit (Jeletzky and Tipper 1967). The largest specimen of fossil wood measures approxi- mately 15 cm long and 10 cm in diameter and consists of mature secondary xylem only. This specimen falls within the general parameters of Protocedroxylon as defined by Gothan (1910). Since it differs from other species of this genus, it is proposed as a new species. SYSTEMATICS Coniferales Protopinaceae Protocedroxylon Gothan Protocedroxylon macgregorii sp. nov. Figs. 1-12 DiaGnosis.—Growth rings distinct, 19-70 tracheids wide, transition from early to late wood gradual, occasionally abrupt; late wood tracheids radially flattened with narrow ellip- tic lumens and walls 5—7 wm thick, early wood tracheids large, angular, with walls 4-7 pm thick, lumens large, 50-85 wm in radial di- ameter; traumatic resin canals occasionally present having 6-12 thick walled and pitted epithelial cells, horizontal resin canals absent; rays uniseriate, frequently partially biseriate, occasionally entirely biseriate, never multise- riate, 1-40 cells high (commonly 12-25); indi- vidual ray cells round to elliptical, largest cells 17-25 pm wide, 25—30 pm high; tracheal pit- ting variable, 1—4 seriate; early wood tracheal pitting typically multiseriate, alternate and tightly appressed (araucarioid), rarely oppo- site or in stellate pit clusters; late wood pitting mostly uniseriate, separate or contiguous; pit borders 17—25 wm in diameter with rounded apertures; tangential pits and wood _par- enchyma absent; rays homogenous, ray par- enchyma highly resinous, horizontal and tan- gential walls pitted, end walls slightly to ‘Uinta Basin Education Center, Utah State University, Roosevelt, Utah 84066. ?Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 452 July 1986 MEDLYN, TIDWELL: FossiL PLANT 453 iA 2, UA Figs. 1-6. Protocedroxylon macgregorii: 1, Transverse section—note the relatively narrow band of late wood (33X). 2, Transverse section showing large, angular tracheids in the early wood (110X). 3, Tangential row of traumatic resin canals in a transverse section (66X). 4, Tangential section showing the uniseriate nature of the vascular rays (66X). 5, Tangential section showing the uniseriate rays containing occasional paired cells (110X). 6, Tangential section showing epithelial cells of a traumatic resin canal (110X). 454 GREAT BASIN NATURALIST Vol. 46, No. 3 Figs. 7-12. Protocedroxylon macgregorii: 7, Radial view of a traumatic resin canal illustrating the pitted, thick — walled epithelial cells (245X). 8, Radial section showing variation in tracheary pitting, note particularly the stellate pit clusters at the left (160X). 9, Radial section showing the pitted nature of the horizontal and vertical walls of the vascular rays (300X). 10, Radial section showing araucarioid tracheal pitting that typifies this genus (300X). 11, Radial section showing crossfield pitting, note the bordered pit (arrow) (750X). 12, Closeup of radial view of vascular ray (150X). July 1986 acutely oblique, crossfields with 1—3 circular, thinly bordered pits (7-10 um), pit borders 2 wm wide, with rounded to elliptic lumens, the cross-field pits tend to be horizontally aligned. HOoLotyPE: Geological Survey Canada Col- lection No. 6776. PARATYPE: Brigham Young University 5027. EtyMo.ocy: This species is named for its collector D. C. McGregor. DISCUSSION Since Gothan (1910) described the genus Protocedroxylon, the history of this taxon has become rather complex. Metacedroxylon was proposed by Holden (1913) to replace Proto- cedroxylon Gothan on the basis that the latter name implies an abietineous affinity. She fur- ther concluded that because Metacedroxlyon lacked “bars of Sanio” and possessed araucari- oid pitting, it could be none other than an araucarian conifer. In the early 1900s many workers, including Gerry (1910), Holden (1913), and Stopes (1916), debated the diag- nostic value of bars or rims of Sanio (crassu- lae). The presence of crassulae was said to indicate possible abietineous relationships, and the absence of crassulae suggested arau- carian affinities. Holden (1913) maintained that presence or absence of crassulae was the only sure criterion for diagnosing fossil conifers. This theory was later investigated by Bailey (1933), who concluded that, although crassulae do not occur in the wood of extant Araucariaceae, it is fallacious to assume that these structures would be preserved under all conditions of fossilization. Metacedroxylon araucarioides (Gothan) Holden and M. latiporosum Holden were de- scribed by Holden (1913) from the Upper Jurassic of the Yorkshire Coast of England. _ Holden (1915) later reported M. scoticum from the Jurassic Corallian beds of the Suther- land Coast of Scotland. Seward (1919) subse- _ quently pointed out the invalidity of Holden's name and combined Metacedroxylon with _ Protocedroxylon. Stopes (1916) not only questioned the use of Metacedroxylon but also Protocedroxylon and Cedroxylon Kraus (1872), believing them to be “taxonomic misfits.” She further added that it is not justifiable to assume an affinity of fossil genera, whose fructifications are unknown, to MEDLYN, TIDWELL: FossiL PLANT 455 living genera. Therefore, to her, Gothan’s use of the names Protocedroxylon and Cedroxy- lon and Holden’s use of Metacedroxylon all seemed inappropriate. Stopes (1916) then proposed a new genus, Planoxylon, in which the principal characters are similar to Gothan’s genus Protocedroxylon. She as- signed two species to her new genus: Planoxy- lon hectori Stopes from the Cretaceous of New Zealand and Planoxylon lindleii (Whi- tham) Stopes. The latter species was first de- scribed by Whitham (1833) as Peuce lindleii and, since that time, has been shuffled from genus to genus. Stopes (1916) removed P. lindleii from Araucarioxylon Kraus, where it was previously classified by Seward (1904). This species has since been placed in Pro- topiceoxylon Gothan by Eckhold (1922), Pro- tocedroxylon Gothan by Edwards (1925), Pinoxylon Knowlton by Read (1932), Dadoxy- lon Endl. by Shimakura (1936), and Yorkoxy- lon Vogellehner by Vogellehner (1968). Krausel (1949) placed Protocedroxylon, as well as various species of Metacedroxylon, Paracedroxylon Sinnott, Paracupressinoxy- lon Holden, and Thylloxylon Gothan into syn- onymy with Araucariopitys Jeffrey. Vogelleh- ner (1968) subsequently included Araucari- opitys, Metacedroxylon, and Planoxylon in Protocedroxylon. Lemoigne (1970) included Protocedroxylon araucarioides — Gothan, Metacedroxylon scoticum Holden, and Mesembrioxylon libanoticum Edwards (1929) in the new genus Embergerixylon Lemoigne (1968). However, based upon priority, Proto- cedroxylon should have precedent over Em- bergerixylon. Shilkina and Khudayberdyyev (1971) fol- lowed Krausel (1949) and concluded that Planoxylon should be separated from Proto- cedroxylon based upon a difference in their tracheal pitting. Nishida and Nishida (1984) retained Planoxylon for fossil conifer woods having typical araucarian pitting and vertical pairs of pits in the crossfield. COMPARISONS Protocedroxylon has been reported from strata of Middle Jurassic to upper Lower Cret- aceous age (Gothan 1910, Seward 1919, Negri 1914, Vogellehner 1968). Protocedroxylon araucarioides has been reported from Meso- 456 zoic strata of Svalbard, West Spitzbergen, and Manchuria (Gothan 1910, Walton 1927, Shi- makura 1940). Among the characteristics of Protocedroxylon araucarioides, Gothan made a special point of noting the absence of normally formed resin canals in this species. Axillary or wood parenchyma is sparse to ab- sent in P. araucarioides, and its vascular rays are always uniseriate. The horizontal and tan- gential walls of the ray parenchyma of P. arau- carioides are heavily pitted and in each cross- field are 1-3 circular pits. The tracheal pitting in P. araucarioides is araucarian. It is charac- terized by uniseriate to triseriate, large, bor- dered pits (20-24 um in diameter) with flat- tened and contiguous borders. Gothan (1910) did not mention the occurrence of crassulae between the bordered pits in this species, and their presence could not be observed in any of his figures. Numerous tangential pits are present in the late wood tracheids. Protocedroxylon transiens (Gothan) Shilk- ina and Khudayberdyyev (= Cedroxylon transiens Gothan) is similar to P. araucario- ides. Terminal parenchyma and stellate pit clusters are the only specific differences be- tween P. transiens and P. araucarioides. Gothan (1907, 1910) cited two occurrences of the former species, one from the Lower Cre- taceous of King Charles Land and the other from the Upper Jurassic or Lower Cretaceous of Spitzbergen. Protocedroxylon macgregorii sp. nov. from the Upper Jurassic of British Columbia is the first report of this genus from North America. Protocedroxylon macgregorii is placed in Pro- tocedroxylon Gothan on the basis of a pre- dominance of araucarioid tracheal pitting cou- pled with pitted horizontal and tangential walls of the rays. A combination of the forego- ing characters is unique to this genus and the araucarioid type tracheal pitting excludes P. macgregorii from either Araucariopitys Jef- frey (1907) or Cedroxylon Kraus (1872). The absence of normally formed resin canals pre- cludes the possibility of close affinities to Pro- topiceoxylon Gothan. Protocedroxylon macgregorii is remarkably similar to P. araucarioides but differs from it primarily in the absence of tangential pitting, the presence of traumatic resin canals or cysts, the marked absence of tracheid septations (a feature notably present in Gothan’s figured GREAT BASIN NATURALIST Vol. 46, No. 3 specimen, P1.5, Fig. 4 [1910]), and occur- rence of partially biseriate vascular rays in P. macgregorii. The rays of P. araucarioides are always uniseriate. The tracheal pitting of both P. transiens and P. araucarioides is similar to that of P. macgregorii. However, the pres- ence of stellate pit clusters in P. macgregorii makes it more closely allied with P. transiens, although differing from the latter by not hav- ing terminal parenchyma. Protocedroxylon macgregorii differs additionally from both P. transiens and P. araucarioides in having occa- sional quadraseriate rows of appressed pits. Yorkoxylon lindleianum (Whitham) Vol- gellehner is very similar to P. macgregorii, but the height of the vascular rays (1-12 cells), the presence of wood parenchyma (Holden 1914), and the absence of stellate pit clusters in the former are notable differences. Protoce- droxylon hectori (Stopes) is similar to our spe- cies; however, the presence of terminal par- enchyma in P. hectori and the height of the vascular rays separate them. Protocedroxylon scoticum (Holden) Seward (1919), with its predominance of uniseriate pitting, is distinct from P. macgregorii. Protocedroxylon paro- nai Negri, from the Cretaceous of North Africa, is not well preserved and therefore not adequately described. A comparison of P. macgregorii with P. paronai would be incon- clusive. Protocedroxylon macgregorii is similar to several Protocedroxylon species from Japan. This species differs from P. japonicum Nishida (1967) from the Cretaceous of Choshi by the latter species having septate tracheids and lower rays (1-4, rarely 6 cells high). Protoce- droxylon okafujii Nishida and Oishi (1982) from the Triassic of Yamaguchi perfecture can be distinguished from P. macgregorii by hav- ing abundant wood parenchyma in incre- ments and septate tracheids. Also from the same strata as P. okafujii are specimens of P. mineense (Ogura) Nishida and Oishi. Origi- nally reported as Araucarioxylon mineense Ogura (1960), this species was subsequently placed in Protocedroxylon by Nishida and Oishi (1982). They further included P. trias- sicum Yamazaki and Tsunada (1981, Yamazaki et al. 1980) in this species. Protocedroxylon macgregorii is separated from P. mineense by the tangential pitting and septate tracheids in the latter and the presence of triseriate pitting July 1986 and stellate pit clusters in the former. Proto- cedroxylon yezoense Nishida and Nishida from the Cretaceous Upper Yezo Group, Hokkaido differs from P. macgregorii in that P. yezoense has septate tracheids, typically uniseriate pitting, the presence of wood par- enchyma, and lower rays (1—28 cells high) that do not occur in P. macgregorii. Protocedroxylon macgregorii and Protopi- ceoxylon canadense Medlyn and _ Tidwell (1979), are presently the only species of petri- fied conifer wood of Jurassic age reported from British Columbia. Additional studies of fossil woods, as well as compression materials from this area, will be necessary before the nature and composition of the Jurassic forest of this region can be fully understood. ACKNOWLEDGMENTS We express appreciation to Dr. S. R. Ash of the Department of Geology, Weber State College, Ogden, Utah, for reviewing this pa- per. We also appreciate the critical comments of Dr. E. M. V. Nambudiri, who examined the specimens. Thanks are also due to Dr. Brian Norford of the ISPG, Calgary, Alberta, for sending us the specimens from which Pro- tocedroxylon macgregorii was described. Par- tial financial support for this study is also ac- knowledged from the Society of Sigma Xi and Brigham Young University Research Grant 115-22-053. LITERATURE CITED BAILEY, I. W. 1933. The cambium and its derivative tis- sue, VII. Problems in identifying the wood of Mesozoic Coniferae. Ann. Bot. 47(185): 145-157. ECKHOLD, W. N. 1922. Die Hoftupfel bei rezenten und fossilen Coniferen. Jb. Preuss. Landesanst. 42: 472-505. Epwarps, W. M. 1925. On Protopiceoxylon johnseni (Schroeter), a Mesozoic coniferous wood. Ann. Bot. 39: 1-7. ———. 1929. Plants from Syria and Transjordania. Ann. Mag. Nat. Hist., Ser. 10°, 4: 401—402. Gerry, E. 1910. “Bars of Sanio” in Coniferales. Ann. Bot. 24: 119-123. GoTHAN, W. 1907. Die fossilen Holzer von Konig Karls Land. Kung]. Svenska Vetensk. Handl. 42(1): 1-41. ———. 1910. Die fossilen Holzreste von Spitzbergen. Kungl. Svenska Vetensk. Hand]. 45(8): 13-56. HOLDEN, R. 1913. Contributions to the anatomy of Meso- zoic conifers. No. 1. Jurassic coniferous woods from Yorkshire. Ann. Bot. 27: 533-545. MEDLYN, TIDWELL: FossiIL PLANT 457 ——. 1914. Contributions to the anatomy of Mesozoic conifers. No. 2. Cretaceous lignites from Cliff- wood, New Jersey. Bot. Gaz. 58: 168-177. ———. 1915. A Jurassic wood from Scotland. New Phyt. 14: 205-209. JEFFREY, E. C. 1907. Araucariopitys, a new genus of araucarians. Bot. Gaz. 44: 435-444. JELETZKyY, J. A, AND H. W. Tipper. 1967. Upper Jurassic and Cretaceous rocks of Taseko Lakes map area and their bearing on the geological history of southwestern British Columbia. Geol. Surv. Canada Paper. 67—54: 1-218. Kraus, G. M. 1872. Bois fossiles de Coniferes. Pages 363-385 in W. P., Schimper, Traite de Paleon- tologie Vegetale ou la Flore du Monde Primitive (1870-1872). J. B. Bailliere et Fils, Paris. KRAUSEL, R. 1949. Die fossilen Koniferenholzer (unter Asschluss von Araucarioxylon Kraus) II. Tiel. Kritsche untersuchungen zur diagnostik lebender und fossiler Koniferenholzer. Palaeontographica 89B: 83-203. LEMOIGNE, Y. 1968. Un nouveau genre de structure ligneuse de type gymnospermien: Embergerixy- lon nov. g. Ann. Soc. Geol. du Nord. 88: 155-157. 1970. Les especes Protocedroxylon araucarioides Gothan, Metacedroxylon scoticum R. Holden et Mesembrioxylon libanoticum W. N. Edwards doivent etre referees au genre Embergerixylon Y. Lemoigne. Bull. Soc. Geol. France, Ser. 7°, 12: 398-402. MEDLYN, D. A., AND W. D. TIDWELL. 1979. A review of the genus Protopiceoxylon with emphasis on North American species. Canadian J. Bot. 57: 1451- 1463. NEGRI, G. 1914. Spora alenni legni fossili del Gebel Tripoli- tano. Bull. Soc. Geol. Italiana 33: 321-344. NisHipa, M. 1967. On some petrified plants from the Cretaceous of Choshi, Chiba Perfecture V. Bot. Mag., Tokyo 80(954): 487-497. NisHipA, M., AND H. Nisuipa. 1984. Structure and affini- ties of the petrified plants from the Cretaceous of northern Japan and Saghalien. I. Petrified plants from the Upper Cretaceous of Hokkaido. J. Jap. Bot. 59(2): 48-57. NIsHIDA, M., ANDT. OtsuI. 1982. The identity of Araucar- ioxylon mineense and a new species of Protoce- droxylon from the Triassic of Miné, Yamaguchi Perfecture. J. Jap. Bot. 57(4): 97-104. OcurRa, Y. 1960. Tyloses in tracheids in Araucarioxylon. J. Fac. Sci., Univ. Tokyo, Sect. III. (Bot.) 7: 501-509. READ, C. B. 1932. Pinoxylon dakotense Knowlton from the Cretaceous of the Black Hills. Bot. Gaz. 93: 173-187. SEWARD, A. C. 1904. Catalogue of the Mesozoic plants in the British Museum (Natural History). The Juras- sic flora. II. Liassic and Oolitic floras of England, London. _____. 1919. Fossil plants. Vol. IV. Cambridge Univer- sity Press, London. SHILKINA, I. A., AND R. KHUDAYBERDYYEV. 1971. New lo- calities and a review of the genera Protocedroxy- lon and Xenoxylon (in Russian). In Palaeobotany in Uzbekistan 2: 117-133. Bot. Inst., Acad. Sci. Uzbeck S.S.R. 458 SHIMAKURA, M. 1936. Notes on fossil woods from Japan and adjacent lands. I, Some Jurassic woods from Japan and Manchoukuo. Sci. Rep. Tohok Imp. Univ. Sendai, Ser. 2, Geol. 18: 267-298. . 1940. The presence of Protocedroxylon araucari- oides Gothan in Manchuokuo and its geologic meaning. J. Shanghai Sci. Inst. sect. II, 2(8): 283-289. Stopes, M. C. 1916. An early type of the Abietineae (?) from the Cretaceous of New Zealand. Ann. Bot. 30(117): 111-125. VOGELLEHNER, D. 1968. Zur Anatomie und Phylogenie Mesozischer Gymnospermenholzer 7: Prodromus zu einer Monographie der Protopinaceae. I. Die protopinoiden Holzer des Jura. Palaeontographica 124B: 125-162. GREAT BASIN NATURALIST Vol. 46, No. 3 WALTON, J. 1927. On some fossil woods of Mesozoic and Tertiary age from the Arctic zone. Ann. Bot. 41(142): 239-254. WuitHaM, H. 1833. The internal structure of fossil veg- etables, accompanied by representations of their internal structure, as seen through the micro- scope. Blackwood, Edinburgh. YAMAZAKI, S., AND K. TSUNADA. 1981. Fossil coniferous woods belonging to Protocedroxylon Gothan and Xenoxylon Gothan, obtained from the Upper Tri- assic Miné Group, southwest Japan. Bull. Sci. and Eng. Res. Lab., Waseda Univ. 97: 1-18. YAMAZAKI, S., K. TSUNADA, AND N. KOIKE. 1980. Some fossil woods from the Upper Triassic Nariwa Group, southwest Japan. Mem. Sch. Sci. and Eng., Waseda Univ. 44: 91-131. i NEW TAXA AND NOMENCLATURAL CHANGES IN UTAH PENSTEMON (SCROPHULARIACEAE) Elizabeth C. Neese! ABSTRACT. —New taxa include Penstemon angustifolius Pursh var. dulcis Neese, P. leonardii Rydb. var. higginsii Neese, P. scariosus Pennel var. cyanomontanus Neese, and P. thompsoniae (Gray) Rydb. var. desperatus Neese. New nomenclatural combinations include: Penstemon acaulis L. O. Williams var. yampaensis (Penl.) Neese, P. duchesnen- sis (N. Holmgren) Neese, P. leonardii Rydb. var. patricus (N. Holmgren) Neese, and P. pachyphyllus Gray ex Rydb. var. mucronatus (N. Holmgren) Neese. Studies leading to a revision of Penstemon for the Utah flora project dictated the following nomenclatural changes and demonstrated the presence of some taxa that require names and descriptions. The justification for these deci- sions will appear in the version of the Utah flora to be published subsequently. Penstemon acaulis L. O. Williams var. yam- paensis (Penl.) Neese stat. nov. [based on Pen- stemon yampaensis Penl. Madrono 14: 156. 1958]. Penstemon angustifolius Pursh var. dulcis Neese var. nov. Similis P. angustifolius var. venosus sed in foliis gracilioribus et colore floribus omnibus roseolioribus differt. TyPE: USA Utah. Millard Co.; T17S, R5W, $35, SE1/ 4, Canyon Mts.—Sevier Desert, 10 km 200 dgrs from Oak City, 1,586 m, juniper community on aeolian sand, 19 May 1981, S. Goodrich 15403 (Holotype BRY; 7 isotypes distributed previ- ously as P. angustifolius Nutt. ex Pursh). ADDI- TIONAL SPECIMENS: Utah. Millard Co.; T17S, R5W, S36, NW1/4, Canyon Mts., 9 km 195 dgrs from Oak City, S. Goodrich 15402; ibid., T17S, R5W, S36 NW1/4, Canyon Mts., Clay Springs Wash, 9 km 195 ders SW of Delta, S. Goodrich 18313; ibid., Millard Sand Dunes, Stanton 5281. Juab Co.; T14S, R5W, Delta, E. Rose 1521; ibid., Little Sahara Dunes, K. H. Thorne 78 (all BRY). Four-wing saltbush, sagebrush-eriogo- num, and juniper communities at 1,400 to 1,650 m in Millard and Juab counties; endemic; 9 (0). Penstemon duchesnensis (N. Holmgren) Neese stat. nov. [based on Penstemon dolius var. | duchesnensis N. Holmgren Brittonia 31: 219. 1979]. Penstemon leonardii Rydb. var. higginsii Neese var. nov. Plantis similis P. leonardii var. leonardii sed in corollis lavandulis (nec caeruleis) et plus suffruticosis differt. TYPE: USA Utah. Washington Co., T39S, R13W, $20, along the Browse road to Guard station, E side of Pine Valley Mts., at 2,000 m, sandy soil, pinyon-juniper-mountain brush commu- nity, 7 June 1983, Larry C. Higgins 13578 (Holotype BRY, 4 isotypes distributed previ- ously as Penstemon). ADDITIONAL SPECIMENS: Utah. Washington Co., Pine Valley, 26 June 1941, W. P. Cottam 8863; ibid, Kolob, 30 July 1965, A. H. Barnum 1407; ibid., Beaverdam Mts., 18 June 1966, Larry C. Higgins 692; ibid., summit of the Beaverdam Mts., 14 June 1970, Larry C. Higgins 3393; ibid., 21 km S of Enterprise, 18 June 1970, Larry C. Higgins 3483; ibid., 4 km N of Cougar Pass, 17 June 1976, N. D. Atwood 6824; ibid., Pine Valley Mts., 8 June 1981, N. D. Atwood 7886; ibid., Kolob Terrace, 7 July 1981, S. L. Welsh 20696; ibid., 10 km S of Enterprise, 10 June 1983, N. D. Atwood 9355; ibid., Cole Flat on road to Slaughter Creek, 10 June 1983, Larry C. Higgins & N. D. Atwood 13711, ibid., Kolob Plateau, 14 July 1983, B. Albee 5615; ibid., Pine Valley Mts., 18 June 1984, Larry C. Higgins 14392; ibid., Kolob Terrace, 7 June 1984, S. L. Welsh, Larry C. Higgins, and K. Thorne 22926: ibid., Pine Valley mts., 8 July 1984, B. & J. Franklin 843; ibid., 6 km E of Enterprise, 13 June 1985, Larry C. Hig- gins & E. Higgins 15710; ibid., Pine Valley Mts., 6 July 1985, G. Baird 1705; ibid., N slope of Pine Valley Mts., 26 June 1985, E. 1 ife Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 459 460 Neese 17092 (all BRY). Pinyon-juniper, mountain brush, ponderosa pine-manzanita, and aspen conifer communities at 2,000 to 2,750 m in the Valley and Bull Valley mts. and Kolob Plateau, Washington County; en- demic; 18 (i). Earlier reports of P. thurberi Torr. from Washington County belong here. Penstemon leonardii Rydb. var. patricus (N. Holmgren) Neese [based on: Penstemon patricus N. Holmgren Brittonia 31: 238. 1979]. Penstemon scariosus Pennell var. cyano- montanus Neese var. nov. Similis P. scariosus var. garrettii (Pennell) N. Holmgren sed in corollis glandularibus extus caeruleo-pur- pureis plerumque et caulibus plus decumben- tibus differt. TypE: USA Utah. Uintah Co., T4S, R25E, S25 S/E1/4, near Reservoir, Blue Mountain Plateau, 2,349 m, in dry creek bed, with Ivesia, Carex, and Rumex, 20 June 1982, E. Neese et al. 11808 (Holotype BRY; five isotypes distributed previously as Penste- mon). ADDITIONAL SPECIMENS: Utah. Uintah Co., Doug Chew's cabin, Blue Mt., 2 June 1956, S. L. Welsh 470; ibid., Warren Draw, 15 June 1978, E. Neese 5693; ibid., Blue Mt. Plateau, 12 June 1982, E. Neese & B. Neely 11752; ibid., 6 July 1982, F. Smith & J. Trent 1742; ibid., Yampa Plateau, 20 June 1982, F. Smith, E. Neese, & J. Trent 1703; ibid., Blue Mt., 13 June 1982, K. Thorne & B. Neely 2003; ibid., 6 July 1982, E. Neese & K. Snyder 11908; ibid., 9 July 1983, R. Dorn 3881 (all BRY). Colorado. Moffat Co., top of Douglas Mt., 1938, Brown 194 (BRY). Sand- stone crevices in sagebrush-grass communi- ties on the Blue Mountain Plateau summit GREAT BASIN NATURALIST Vol. 46, No. 3 near the Colorado/Utah border, Uintah Co.; Colorado; a Uinta Basin endemic; 13 (vi). The taxon was previously reported from the region as P. cyanocaulis Payson (Welsh 470 BRY). Penstemon thompsoniae (Gray) Rydb. var. desperatus Neese var. nov. Ab P. thompsonae var. thompsonae in caulibus et inflorescentiis longioribus et foliis minoribus differt. TyPE: USA. Utah. Beaver Co., ca 8 mi SW of Saw- tooth Peak, Indian Peak Range, T29S, R19W, $3, at 2,013 m, 26 May 1976, S. Welsh, K. Taylor, & F. Peabody 13287 (Holotype BRY; 2 isotypes distributed previously as P. thomp- sonae ). ADDITIONAL SPECIMENS: Utah. Beaver Co., Hamblin Valley, 31 May 1978, K. Ostler & D. Anderson 1197; ibid., Wah Wah Mts., 22 June 1978, K. Ostler 1465 (all BRY). Iron Co., Hamblin Valley, 16 June 1978, K. Ostler & D. Anderson 1407; ibid., ca 21 km W of Enterprise, 13 June 1985, L. C. & E. Higgins 15725; ibid., Enterprise-Panaca road, 6 km E of the Nevada border, 12 June 1984, N. H. & P. K. Holmgren 10445 (all BRY). Sagebrush and pinyon-juniper communities at 1,800 to 2,075 m in Beaver and Iron counties; Nevada; 6 (0). One specimen tentatively assigned here (Atwood 5151 BRY) has stems unusually long, and is probably more closely allied to materi- als from the Mohave Strip region of Arizona. Possibly it represents still another taxon wor- thy of naming. Other, as yet unrecognized, infraspecific taxa may occur within the species beyond Utah. Penstemon pachyphyllus Gray ex Rydb. var. mucronatus (N. Holmgren) Neese stat. nov. [based on: Penstemon mucronatus N. Holmgren Brittonia 31: 234. 1979]. RELICT OCCURRENCE OF THREE “AMERICAN” SCOLYTIDAE (COLEOPTERA) IN ASIA Stephen L. Wood! and Hui-fen Yin? ABSTRACT. —The first non-American representatives of the bark beetle genus Pseudopityophthorus , of the subtribe Corthylina (Corthylini), and of the Ips concinnus species group are named. All are apparently relicts of an early faunal exchange with North America. The new taxa include: Gnatharus, new genus, and its type-species Gnatharus tibetensis , new species (Tibet); Xenophthorus, new subgenus, and its type-species Pseudopityophthorus (Xenophtho- rus) peregrinus , new species (Tibet), and Ips orientalis , new species (Tibet, China). The intercontinental exchange of other taxa in Scolytidae are also mentioned. Faunal interchange between North Amer- ica and Asia has been widely recognized ever since the discovery of America. Among the Scolytidae this has included the invasion of the Eurasian genus Polygraphus (about 45 species) into North America (3 species), Phloeosinus (40 Asian to 29 American spp.), Cryphalus (over 200 Asian to 3 American spp.), Dryocoetes (over 80 Asian to 7 Ameri- can spp.) Orthotomicus (12 Asian to 1 Ameri- can spp.), Trypodendron (12 Eurasian to 5 American spp.), etc. Migration in the oppo- site direction has included Carphoborus (21 American to 14 Eurasian spp.), Scolytus (over 60 American to 47 Eurasian spp.), Pityophtho- rus (over 300 American to about 35 Eurasian spp.), Ips (83 American to 18 Eurasian spp.), Pityokteines (6 American to 3 Eurasian spp.), etc. In each case anatomical and phylogenetic diversity in the invaded area is conspicuously less than in the area of origin such that no species group occurred in the invaded area that was not also present in the area of origin. More species groups in each genus always occurred in the area of origin than occurred in the invaded area. In view of the above, it should be expected that additional examples of faunal exchange will be found. The Eurasian occurrence of Dendroctonus micans Kugelann, a geographi- cal replacement species of the subpolar D. punctatus LeConte, in a genus that otherwise was exclusively American, has been known for almost a century. However, the discovery of D. armandi Tsai & Li in China, with no allied sister species in America was surprising. Equally unexpected was the discovery that the species named from China as Gretshckinia mongolica Sokanovskii actually belongs to the American genus Pseudothysanoes, and, in fact, is the only represenative of its tribe in Asia. In a review of the Scolytidae of China, repre- sentatives of three additional groups thought to be exclusively American were found. The first is a previously unknown species of Ips that is allied to I. concinnus (Mannerheim). The second is an aberrant species of Pseudopityophthorus. The third is a previously unknown genus in the sub- tribe Corthylina, the first native species found outside of America. It is anticipated that addi- tional relicts of an early faunal exchange will be found in both Asia and North America as the faunas become more completely known. These new taxa are described as follows. Ips orientalis, n. sp. This species is distinguished from the closely allied concinnus (Mannerheim) of western North America by the stouter body form, by the absence of a frontal fovea and tubercle, by the less deeply excavated elytral declivity, with the lateral margins less strongly elevated and the denticles smaller, and by the much more pronounced sexual dimorphism of the declivity. MALE.—Length 4.4 mm (paratypes 3.5— 4.4 mm), 2.3 times as long as wide; color dark reddish brown. lLife Science Museum and Department of Zoology, Brigham Young University, Provo, Utah 84602. "Institute of Zoology, Academia Sinica, 7 Zhongguancun Lu, Haidian, Beijing, China. 461 462 Frons resembling concinnus except tubercles distinctly smaller, particulary those on epis- tomal margin; central fovea entirely absent, me- dian tubercle on epistomal margin and not larger than other tubercles. Antennal club with base significantly thicker and more corneous, sutures equally procurved. Pronotum as long as wide, essentially as in concinnus except discal area much shorter. Elytra 1.3 times as long as wide, 1.4 times as long as pronotum; basic outline and structure as in concinnus except stouter, discal punctures less numerous, larger, deeper, confused; decliv- ity of same basic structure as concinnus except much less strongly excavated, rather weakly ex- planate, lateral margins on upper half rounded, all denticles smaller and more broadly conical, 2 without a ventral carinate ridge extending to- ward 3. Vestiture about as in concinnus. FEMALE.—Similar to male except frons with a poorly formed central fovea about as in female concinnus; elytral declivity with apical margin subacute, but not explanate, and not extending as far laterally; declivital excavation narrower and lateral margins not as high as in male. TYPE LOCALITY. —Baxoi, Xizang (Tibet). TYPE MATERIAL.—The male holotype, female allotype, and one male and two female paratypes are from Baxoi, Xizang (Tibet), 18-VIII-1973, Picea, Huang Fu-sheng. Two female paratypes are from Riowge, Xizang (Tibet), 19-VIII-1976, same collector and host. One male and two fe- male paratypes are from Barkam, Sichuan, 24- VII-1955, Picea, Qui De-xun. One male paratype appears to be from the type locality, taken 18-VI-1974. The holotype, allotype, and four paratypes are in the Institute of Zoology, Academia Sinica, five paratypes are in the Wood collection. Notes.—The discovery of this species is sig- nificant because it represents the first occur- rence of this distinctive species group in an area outside of North America. Additional signifi- cance is found in the fact that it is by far the most primitive member of its group, and of Ips, and adds to the evidence that both Ips and Orthoto- micus were derived from Acanthotomicus , and that the obliquely truncate antennal club of this tribe is not primitive in this tribe. Pseudopityophthorus Swaine The genus Pseudopityophthorus has con- sisted of 23 species from North and Central GREAT BASIN NATURALIST Vol. 46, No. 3 America and one species from northern South America (Colombia). All bore in the phloem tissues of Quercus where they form character- istic transverse biramous parental tunnels. It was most surprising, therefore, to find a spe- cies in this uniquely American genus in Asia, also in Quercus. While this species is clearly allied to the first six species in my key (Wood, 1982, Great Basin Nat. Mem. 6: 966), it is thought appropriate to place it in a separate subgenus, based primarily on characters of the antennal club. Xenophthorus, new subgenus.—Antennal club broadly oval, almost subcircular, sutures profoundly procurved, suture 1 attaining mid- dle of club; anterior margin of pronotum not regularly serrate. Other characters as in prim- itive members of genus as described for the type-species. Type-species. —Pseudopityophthorus (Xe- nophthorus) peregrinus , described below. Pseudopityophthorus peregrinus, n. sp. This species is distinguished from all other members of the genus by the very strongly procurved sutures of the antennal club, by the presence of strial punctures on both disc and declivity, by the long vestiture, and by other characters described below. FEMALE.—Length 1.9 mm (paratypes 1.7— 2.0 mm), 3.0 times as long as wide; color very dark brown. Frons convex, a moderate transverse im- pression on lower third; surface reticulate- granulate, upper half and sides with sparse, rather coarse granules; vestiture sparse, rather long, hairlike. Antennal scape shorter than club, slender; funicle 5-segmented; club 1.17 times as long as scape, 1.17 times as long as wide, sutures broadly, very strongly pro- curved, marked by rows of setae, 1 extending slightly beyond middle of club. Pronotum resembling granulatus Blackman except posterior areas more coarsely punc- tured and anterior margin unarmed except for about four irregularly placed serrations. Elytra 1.8 times as long as wide, 1.5 times as long as pronotum; striae not impressed, punctures small; moderately deep, in rows, spaced by about diameter of a puncture; inter- striae almost smooth and shining, three times as wide as striae, punctures feebly granulate, half as large as those of striae, uniseriate, as July 1986 closely spaced as strial punctures. Declivity steep, convex; striae continued to apex, punc- tures much smaller than on disc; interstriae very slightly shagreened, 1 weakly elevated, all with punctures replaced by small uniseri- ate granules. Vestiture consisting of short strial hair and long, coarse, pointed interstrial hair; each interstrial seta longer than distance between rows, up to twice as long on decliv- ity. TYPE LOCALITY.—Zayti, Xizang (Tibet). TYPE-MATERIAL.—The female holotype and four female paratypes were taken at the type locality on 2-VIII-1973, from Quercus, by Huang Fu-sheng. The holotype and two paratypes are in the Zoological Insititute, Academia Sinica, two paratypes are in the Wood collection. Gnatharus, n. g. This genus is the only native member of the subtribe Corthylina to occur outside of Amer- ica. Although aberrant in all characters, it ap- pears most nearly allied to Gnathotrupes Sched to which it is remotely related at best. The prothoracic precoxal flange resembles that of Tricolus Blandford. DESCRIPTION.—Frons convex in both sexes, without special sculpture or ornamen- tation. Eye oval, finely faceted, more than one-third divided by an emargination. Anten- nal scape club-shaped, slightly shorter than club; funicle 4-segmented; club subcircular, with two rather strongly procurved sutures marked by both grooves and rows of setae. Pronotum sexually dimorphic, female much as in small Gnathotrupes, summit at middle, anterior slope strongly declivous and armed by numerous small asperities, very finely sculptured on posterior half, male weakly de- clivous in front, asperities largely obsolete, carinate anterior margin extended cephalad. Elytra simple, striae obsolete, declivity con- vex, rather steep, but suture divaricate from middle of declivity, elytra truncated before apex. Protibia in female inflated on posterior face and armed by numerous, confused tuber- cles; in male flattened, apparently unarmed (concealed from view); lateral margin armed almost to base. TYPE-SPECIES: Gnatharus tibetensis Wood & Yin, described below. WOOD, YIN: RELICT BARK BEETLES 463 Gnatharus tibetensis, n. sp. This unique species has no near relatives. It is characterzied as follows. FEMALE.—Length 2.0 mm (allotype 1.7 mm, paratype 2.0 mm), 2.8 times as long as wide; color yellowish brown, with anterior half of pronotum and posterior half of elytra reddish brown. Frons broadly convex on upper two-thirds, shallowly, transversely impressed, median line from impression to epistomal margin with a weak double-crested carina; surface strongly reticulate, punctures rather coarse, shallow, not close; vestiture of sparse, fine, short, in- conspicuous hair. Eye and antenna as de- scribed for genus. Pronotum 1.2 times as long as wide; sides straight and parallel on posterior two-thirds, broadly rounded in front; anterior margin broadly, almost weakly subcostate, serrations ranging from definite to indefinite, numerous; summit anterior to middle, anterior slope strongly declivous, armed by numerous rather small asperities; posterior areas strongly reticulate, punctures very fine, shal- low. Vestiture of fine, short, rather sparse hair; anterolateral angles with a small tuft of longer hair as in some female Gnathotrupes. Procoxae contiguous, precoxal piece a simple, transverse partition bent (flanged) cephaled as in most Tricolus. Elytra 1.7 times as long as wide, 1.3 times as long as pronotum; scutellum large, flat; sides almost straight and parallel on more than anterior two-thirds, rather broadly rounded behind except divaricate at suture and with a sublateral denticle; disc smooth shining, striae obsolete, punctures strongly confused, small, distinct, rather close. Declivity rather steep, convex, apex complex; sculpture as on disc; suture beginning to divaricate at middle of declivity, separation gradual and modest to lower fourth, then abruptly (almsot 90 de- grees) diverging and curving to meet costal margin in a subspinose point, distance be- tween points equal to almost half of total ely- tral width. Vestiture of fine, short, rather abundant recumbent hair. Tibiae wider than normal for tribe, meso- and metatibiae each armed by three socketed teeth, protibiae as described for genus. MALE.—Similar to female except frontal carina with single summit; pronotum with 464 strongly formed, slightly produced anterior costa (serrations almost obsolete), anterior slope much more gradual, asperities greatly reduced in size, without special tufts of hair on anterolateral angles; posterior face of protibia flat and unarmed. TYPE-LOCALITY.—Médog, Xizang. (Tibet). GREAT BASIN NATURALIST Vol. 46, No. 3 TYPE-MATERIAL.—The female holotype, male allotype, and one female paratype were taken at the type locality on 8-IX-1974, 1400 m, from a Castenopsis by Huang Fu-sheng. The holotype and allotype are in the Zool- gocial Institute, Academia Sinica, the paratype is in the Wood collection. —————————— NEW GENUS OF SCOLYTIDAE (COLEOPTERA) FROM ASIA Stephen L. Wood! and Fu-sheng Huang” ABSTRACT.—Pseudoxylechinus, new genus, and uniformis (type-species) (Yunnan), variegatus (Sanxi), sinensis (Yunnan), rugatus (Yunnan), and tibetensis (Tibet), new species, are described. Kissophagus tiliae Niisima, 1910, is also transferred to this genus. In a review of the Scolytidae of China, a genus new to science was found that appears to be a geographical replacement of the closely related North American Pseudohylesinus Swaine. Of the seven species examined, all are substantially smaller than are the species of Pseudohylesinus and all breed in angiosperm hosts. The species Kissophagus tiliae Niisima (1910, Sapporo Nat. Hist. Soc. 3:2), from Japan, is here transferred to Pseudoxylechinus . Pseudoxylechinus, n. g. D1AGNosis.—This genus is distinguished from Pseudohylesinus Swaine by the smaller, closer strial punctures, by the unarmed discal inters- triae (except in rugatus), by the closer, coarser erect discal interstrial setae, by the more slen- der, apically pointed interstrial ground setae, by the more strongly flattened antennal club with its apex more broadly rounded, and by their occurrence in angiosperm hosts. DESCRIPTION. —Allied to Pseudohylesinus ex- cept species mostly smaller, variegated scale patterns less pronounced to absent. Frons sexu- ally dimorphic, more nearly flattened, often with a median carina or groove. Antennal funicle 7-segmented; club flattened, its apex rather _ broadly rounded. Elytral striae rather narrow, punctures close; interstriae twice as wide as striae usually unarmed on disc. Interstrial ground setae rather slender, each with its apex pointed. TYPE-SPECIES: Pseudoxylechinus uniformis Wood & Huang. Pseudoxylechinus uniformis, n. sp. This species is distinguished from other members of the genus by the large size and by the presence of a median carina on the frons in both sexes. MaALeE.—Length 2.6 mm (paratypes 2.5— 2.8 mm), 2.2 times as long as wide; color dark brown, vestiture uniformly pale. Frons convex except moderately impressed on lower two-thirds on median third, im- pressed area with a fine, long, median carina; surface shining, rather coarsely rugose-punc- tate laterally and above, smooth, shining, and impunctate near carina; vestiture of rather short, coarse, moderately abundant hair. Eye oval, entire, finely granulate, 3.0 times as long as wide. Pronotum 0.81 times as long as wide; widest at base, strongly, arcuately converging on basal three-fourths to moderate constric- tion just behind rather broadly rounded ante- rior margin; surface smooth, shining, with abundant, minute punctures interspersed with less abundant larger punctures (twice diameter of smaller ones), larger punctures spaced by one to two diameters of a large puncture; small punctures bearing scalelike ground setae, each four times as long as wide, strongly tapered from base to acute point; large punctures each bearing a coarse, slender bristle, each about twice as long as scales. Elytra 1.5 times as long as wide, 2.1 times as long as pronotum; sides almost straight and parallel on basal two-thirds, rather narrowly, unevenly rounded behind; striae weakly im- pressed, punctures small, deep, close; inter- striae twice as wide as striae, small punctures minute, shallow, obscure, central row of bris- tle-bearing punctures modestly granulate. Declivity moderately steep, convex; inter- striae 1 and 3 moderately convex, 2 impressed 1Life Science Museum and Department of Zoology, Brigham Young University, Provo, Utah 84602. Institute of Zoology, Academia Sinica, 7 Zhongguancun Lu, Haidian, Beijing, China. 165 466 and narrower than | or 3 and without bristles or granules, otherwise sculpture as on disc. Vestiture of rather numerous ground scales, each slightly longer than wide and tapered from its base to acute point; each interstriae with a row of uniseriate, stout bristles, each about three-fourths as long as distance be- tween rows. FEMALE.—Similar to male except frons more uniformly convex. TYPE-LOCALITY.—Nanjiang, Sichuan, Chi- na. TYPE-MATERIAL.—The male holotype, fe- male allotype, and a male and three female paratypes were taken at the type locality on 15-VIII-1958, from a broad leaf tree by Song Shi-mei. One female paratype is from Taibai, Shaanxi, China, V-1981, Pinus tabulaeformis (accidental?), by Yang Xu-hui. The holotype, allotype, and two paratypes are in the Institute of Zoology, Academia Sinica, three paratypes are in the Wood col- lection. Pseudoxylechinus variegatus , n. sp. This species is distinguished from wuni- formis Wood & Huang by the smaller size, by the more broadly impressed, grooved, male frons, by the more uniformly punctured pronotum, and by the very different elytra as described below. MALE.—Frons 2.0 mm (paratypes 1.8—2.1 mm), 2.1 times as long as wide; color dark brown, vestiture 50 to 70 percent pale, re- mainder darker. Frons almost flat on slightly more than lower half, upper half of flattened area with a median impression (groove); surface almost smooth, shining on lower half, with fine, shal- low punctures; upper areas rugose-reticulate, with obscure, rugose punctures; vestiture of short, rather stout, moderately abundant se- tae. Pronotum resembling wuniformis except sides on basal half subparallel, more strongly arcuate; surface rather coarsely, deeply, uni- formly punctured; vestiture of short, stout, moderately abundant bristles, obscurely bi- colored. Elytra about as in uniformis except inter- strial tubercles higher, sharper; declivity steeper, uniformly convex, interstriae 2 equal in height and convexity to 1 and 3; interstrial GREAT BASIN NATURALIST Vol. 46, No. 3 ground setae longer, each about four times as long as wide, each tapered from its base, sharply pointed, bicolored, forming an irregular pattern of pale and darker scales, pale predominate; erect bristles about 1.5 times as long as ground setae, of equal width. FEMALE.—Similar to male except lower frons more convex, median groove present. TYPE LOCALITY. —Jiangxian, Shanxi, China. TYPE MATERIAL.—The male holotype, female allotype, and three male and three female paratypes were taken at the type locality on 5- VIII-1972, from Elaeagnus sp., by Huang Fu- sheng. The holotype, allotype, and two paratypes are in the Zoological Institute, Academia Sinica, and four paratypes are in the Wood collection. Pseudoxylechinus tibetensis, n. sp. This species is distinguished from variega- tus Wood & Huang by the stouter body form, by the less extensively flattened lower frons and shorter groove, and by the larger inter- strial tubercles. - FEMALE.—Length 2.2 mm (allotype 2.1 mm), 2.1 times as long as wide; color very dark brown, vestiture forming a variegated pattern of half pale and half dark scales. Frons irregularly convex, median groove very short; surface rugose-reticulate, ob- scurely, rather coarsely punctured; vestiture rather short, coarse, moderately abundant. Pronotum about as in uniformis, surface sculpturing similar to variegatus except punc- tures slightly smaller, setae shorter; slender and stout setae clearly discernable. Elytra about as in variegatus except inter- strial tubercles on disc distinctly larger, de- clivital interstriae | distinctly higher than 2 or 3, interstrial ground scales shorter, stouter, each less than twice as long as wide, erect interstrial setae more slender. Vestiture about equally divided between pale and dark. MALE.—Similar to female except frons less extensively flattened on lower half, median groove much shorter than male variegatus. Declivity of allotype destroyed (apparently by a predator). | TYPE LOCALITY.—Zayii, Xizang (Tibet). TYPE MATERIAL. —The female holotype and damaged male allotype were taken at the type | locality on 18-IV-1973, 2500 m, from an | unidentified host, by Huang Fu-sheng. July 1986 The holotype and allotype are in the Zo- ological Institute, Academia Sinica. Pseudoxylechinus sinensis, n. sp. This species is distinguished from tibetensis Wood & Huang by the absence of a frontal groove, by the more slender pronotal and ely- tral ground setae, and by the (apparently) uni- formly pale elytral vestiture. FEMALE.—Length 1.8 mm (paratype 1.9 mm), 2.0 times as long as wide; color dark brown, vestiture apparently uniformly pale (covered by an incrustation). Frons convex, distinctly inflated just above epistoma, groove absent; surface obscurely subreticulate and rugose-punctate above, more finely sculptured on lower third; vesti- ture rather fine, short, inconspicuous. Pronotum about as in tibetensis except scale-like setae much more slender. Elytra as in tibetensis except interstrial tubercles smaller, declivity more broadly convex with interstriae 1-3 more equally, uniformly con- vex, scales in ground vestiture much more slender (each about four times as long as wide) but not longer, apparently of uniform color. TYPE LOCALITY. —Lijiang, Yunnan, China. TYPE MATERIAL.—The female holotype and one female paratype were taken at the type locality on 2-IX-1962, from an unidentified host by Song Shi-mei. The holotype is in the Zoological Institute, Academia Sinica, the paratype is in the Wood collection. Pseudoxylechinus rugatus, n. sp. This species is distinguished from other members of the genus by the strongly im- pressed striae, by the large strial punctures, by the coarse interstrial tubercles, and by the very slender interstrial ground setae. WOoOD, HUANG: ASIAN BARK BEETLES 467 MALE.—Length 2.2 mm (allotype 2.1 mm), 2.0 times as long as wide; color dark brown, vestiture pale. Frons shallowly concave from epistoma to upper level of eyes from eye to eye; surface largely obscured by an incrustation but appar- ently with a short median groove, rugose- reticulate, obscure punctures finely rugose. Pronotum outline about as in variegatus , surface shining, punctures coarse, deep, very close, of mixed sizes; vestiture of recumbent, slender, almost hairlike ground setae and erect, rather stout bristles of almost equal length. Elytra 1.4 times as long as wide, 1.1 times as long as pronotum; sides almost straight and parallel on basal two-thirds, broadly rounded behind; striae rather strongly impressed, punctures coarse, deep, close; interstriae as wide as striae, each armed by a uniseriate row of closely set, coarse, almost subvulcanate, setiferous tubercles, each tubercle almost as high as wide and almost as wide as interstriae. Declivity steep, convex; punctures and tuber- cles distinctly smaller than on disc. Vestiture limited to a row of ground setae on each side of each interstriae, each seta slender (about eight times as long as wide), and erect bristles, one arising from each tubercle, each bristle only slightly longer and stouter than ground setae. FEMALE.—Similar to male except frons convex, modestly inflated just above flattened epistomal area. TYPE LOCALITY.—Xishuangbanna, Yunnan, China. TYPE MATERIAL.—The male holotype and female allotype were taken 7-V-1962, 750 m, from Cassia sp., by Song Shi-mei. The holotype and allotype are in the Zo- ological Institute, Academia Sinica. NEW PSEUDOXYLECHINUS (COLEOPTERA: SCOLYTIDAE) FROM INDIA Stephen L. Wood’ ABSTRACT. —Pseudoxylechinus indicus is decribed as new to science from India. In my review of the Scolytidae of India and China, the genus Pseudoxylechinus was dis- covered and named. A species new to India is here added to the Chinese and Japanese spe- cies now in this genus. Pseudoxylechinus indicus, n. sp. This species is distinguished from other members of the genus by the strongly, broadly excavated male frons, by the shal- lowly concave female frons, and by the almost hairlike ground setae on the pronotum and elytra. MALE.—Length 2.7 mm (allotype 2.4, paratype 2.8 mm), 2.1 times as long as wide; color very dark brown, elytra lighter brown, vestiture pale. Frons very deeply, broadly concave from epistoma to well above upper level of eyes, from eye to eye, margin from epistoma to eye strongly, acutely carinate; surface rugose- reticulate, punctures sparse, minutely granu- late; vestiture sparse, hairlike, rather short except much longer at inner margin of eye. Pronotum 0.94 times as long as wide, out- line as in variegatus Wood & Huang; surface shining, coarsely, rather shallowly, rugosely punctured, a few small tubercles in median area near anterior margin; vestiture of moder- ately abundant, rather long, coarse hair. Elytra 1.6 times as long as wide, 2.2 times as long as pronotum; outline as in wniformis Wood & Huang; striae slightly impressed, puctures small, deep, close; interstriae twice as wide as striae, smooth, shining, punctures sparse, minute, each with a uniseriate row of small tu- bercles. Declivity rather steep, convex, sculp- ture as on disc. Vestiture of moderately abun- dant, short, slender, almost hairlike ground setae, and interstrial rows of erect, equally slen- der bristles, each bristle slightly more than twice as long as ground setae. FEMALE.—Similar to male except frons shallowly concave on lower third, modestly convex above, lateral margins on lower third acutely carinate; vertex with a short, median, subcarinate callus. TYPE LOCALITY.—Rangirum, Darjeeling, Bengal, India. TYPE MATERAL.—The male holotype, fe- male allotype, and one male paratype were taken at the type-locality on 6-IX-1929, from Quercus lamellosa, by J.C.M. Gardner. The holotype and allotype are in the Forest Research Institute, Dehra Dun, the paratype is in my collection. 1 Life Science Museum and Department of Zoology, Brigham Young University, Provo, Utah 84602. 468 WILDLIFE DISTRIBUTION AND ABUNDANCE ON THE UTAH OIL SHALE TRACTS 1975-1984! C. Val Grant” ABSTRACT. — Distribution and abundance of 215 amphibians, reptiles, birds, and mammals were monitored for 10 years on Utah’s Oil Shale Tracts using line transects, mist netting, and live trapping. Wildlife monitoring was conducted in four major vegetation types and during all seasons to establish a quantitative baseline for use in impact identification during oil shale mining. Habitat preferences were established for many species in cold desert vegetation of two types of desert shrub, and juniper and riparian woodlands. Seasonal, annual, and habitat distribution of each class demonstrated a variety of adaptive responses to environmental variables. The most important environmental variables, that is, those factors resulting in a predictable change in wildlife populations, were, in descending order: weather, food resource, shelter, and competition. Distributional accounts of wildlife are usu- ally limited to a particular class, for instance, birds (Behle and Perry 1975, Hayward et al. 1976, Behle 1981, Cook 1984) and mammals (Durrant 1952, Ranck 1961). Western am- phibians and reptiles are treated on a regional basis (Stebbins 1966). Nevertheless, regional accounts of wildlife are available for the Utah portion of the Upper Colorado River Basin (Hayward et al. 1958) or for the Uinta Basin Oil Shale Area (Olsen 1973). These distributional accounts of wildlife relegate abundance to a subjective reference, such as common, uncommon, rare, occa- sional, etc. Although Behle and Perry (1975) attempt to quantify abundance, there is a ba- sic flaw in an estimator that rates abundance of the golden eagle (Aquila chrysaetos) and the horned lark (Eremophilia alpestris) as com- mon (Twomey 1942, Olsen 1973, Behle and Perry 1975, Hayward et al. 1976, Behle 1981). Counting two eagles and 2,000 larks on the same day leaves some question as to what common really means. Temporal accounts of wildlife, that is, resi- dency status, characterize a state (Durrant 1952, Behle and Perry 1975, Hayward et al. 1976), aregion (Twomey 1942, Behle 1981), a county (Cook 1984), or an area within a county (Ranck 1961, Olsen 1973). Due in part to the geology of Utah, wildlife that are permanent residents in the southern deserts and high plateaus are transients in the northern basins and mountains. Wildlife that summer in the mountains usually winter in the basins. In other words, as scope changes, residency status changes; yet most accounts defer one to another, regardless of reality. Spatial accounts of wildlife, that is, distribu- tions by habitat or vegetation type, are essen- tially absent for most reptiles and broadly stated for most birds (Behle 1981, Walters and Sorenson 1983) and some mammals (Durrant 1952, Hasenyager 1980). Specificity is usually limited to game birds and mammals, some rodents, raptors, and threatened and endan- gered species. In an effort to fill some voids, a detailed monitoring program for amphibians, reptiles, birds, and mammals began in 1975 and ended, in part, in 1984 at the Oil Shale Tracts Ua-Ub in the Uinta Basin of northeastern Utah. A species-by-species account is re- ported herein as well as population dynamics of each wildlife class, except that the mam- mals are divided into bats, rodents, rabbits, carnivores, and ungulates. A detailed account of birds by feeding guild is presented else- where (Steele et al. 1987), as is waterfowl (Steele and Vander Wall 1985) and sampling effort (Steele et al. 1984). STUDY AREA Ua-Ub includes 42 km? (16 mi’) of cold desert. Elevation ranged from 1,463 m in the 1This research was funded by White River Shale Oil Corporation, Salt Lake City and Vernal, Utah. ?Bio-Resources, Inc., 135 East Center Street, Logan, UT 84321. 469 1334 00L IWAY3LNI YNOLNOD S31IW NI31V9S ii =i aa = t A SH3L3SWOTIN NI3TVOS t i) S t 46, No. 3 Vol ainovalvis () G3AOUdWINN -=----- 3DV4UNS G3AONdWI. —-— savou GREAT BASIN NATURALIST ("14 0029-0009) SH313W 0681-6281 (-14 0009-0095) SH313W 6281 -ZOZL ("14 0095-0025) SH3LAW Z0Z1-SBSL ("14 0025-008) SH3.L3W S8SI -ESbL NOILWAJ14 0€.Z1 470 July 1986 45 40 4 --~ ANNUAL PPT SPRING PPT (CM) PRECIPITATION 0 IDES 1976 1977 1978 GRANT: OIL SHALE TRACT WILDLIFE AS) 47] 1980 1981 1982 1983 1984 Fig. 2. Annual precipitation (October-September) and spring precipitation (April-June) at Oil Shale Tracts Ua-Ub, Uintah County, Utah (after VIN, Inc.). [ll] 1978-1982 I \983 7 cm Prectpitatton Z / Fig. 3. Seasonal precipitation for 1978-1982 and 1983 at Oil Shale Tracts Ua-Ub, Uintah County, Utah (after VIN, Inc.). White River fluvium to 1,890 m atop steep walls of Uinta sandstone near the tract’s south- ern perimeter (Fig. 1). Precipitation changed dramatically over the 10 years (Fig. 2); how- ever, seasonal distribution remained the same (Fig. 3). Ambient temperature averaged 8 C and was coldest during the winter along the White River (Fig. 4). Wind speed peaked dur- ing the spring, and airflow during the daytime was generally westerly; during the night cold air drained into the White River canyon. Four vegetation types covered Ua-Ub (Fig. 5). Dominant species in Greasewood vegeta- tion were big sagebrush (Artemisia triden- tata), greasewood (Sarcobatus vermiculatus), rabbitbrush (Chrysothamnus nauseosus ), and cheat grass (Bromus tectorum). Dominant species in Shadscale vegetation were shad- scale (Atriplex confertifolia), big sagebrush, snakeweed (Xanthocephalum sarothrae), and cheat grass. Juniper vegetation was domi- nated by Utah juniper (Juniperus os- teosperma), black sagebrush (Artemisia nova ) in the draws, and cheat grass. Riparian vege- tation was dominated by Fremont cottonwood (Populus fremontii), tamarisk (Tamarix te- trandra), rabbitbrush, greasewood, and cheat grass. Stem leader growth of big sage- bursh and biomass of annual and perennial grasses and forbs were used to estimate floral productivity over 10 years (Fig. 6). MATERIALS AND METHODS Sampling locations for terrestrial wildlife were within and near Ua-Ub’s boundary (Fig. 472 30 8 oo \ 7 AN Joe YY a 18 5- AN iF a“ \ ‘ Us ET PORN 4 / wk Y NN i: ? Fo ty OW ws ¢ a SN YP Hs , \ \f i g \ . ifs : ‘ \ = Hf i ‘ \ 6 Mm / : \ \ . SP ae ry \ 43 y 6 \ \ © Yj U / vi \ = He / a ! cy i \ ho tee EN SON p : yo . o 4 : z \ c /} i a v y Ay . = dé i . ‘ y i= -6 / s © / si [= a . ul sy a a ¢ ° 1s @--—- Juniper - Uplands *@--- Shadscale - Uplands *---- Evacuation Creek Canyon ares White River Canyon -30 Dec Feb Apr Jun Oct Fig. 4. Seasonal temperature patterns from upland and canyon locations at Oil Shale Tracts Ua-Ub, Uintah County, Utah (after Aerovironment, Inc.). Aug 7), including a mist-netting site for bats at a gas well in the southwest corner of the area mapped. Each vegetation type was sampled for amphibians, reptiles, birds, and terrestrial mammals (including rodents, rabbits, carni- vores, and ungulates) by 1 km (0.6 mi) line transects (Emlen 1971). Transects were walked in the early morning for birds, from midmorning to midday for reptiles, and late afternoon to early evening for mammals. All animals observed were recorded by species and number of individuals seen. Perpendicu- lar distance from the transect was estimated for all visual sightings. Auditory identifica- tions were used for species count only, as were tracks, scat, or sign encountered on tran- sect. Transects were surveyed for five consec- utive days each month during February, April, June, August, and October. From 1982 to 1984 transects for birds and reptiles were surveyed only during June and for mammals only during August. GREAT BASIN NATURALIST Vol. 46, No. 3 For small nocturnal rodents, four large grids (12 x 12 array of Serman live traps) covering 2.73 ha (6.6 acres) were trapped every August in each vegetation type. All traps were baited with oats or barley. All rodents were identified to species, weighed to the nearest gram by 100 g and 500 g capacity Pesola scales, aged (juvenile or adult), sexed, marked for individual recognition, trap site recorded, and released. Seasonal trapping was conducted from 1975 to 1981, with live traps set along transects during February, April, June, August, and October, using the same pro- cedures but fewer traps in different arrays (Steele et al. 1984). Bats were mist-netted at a small pond formed by low-quality water from an aban- doned, artesian natural gas well. Cattails (Ty- pha sp.) surrounded the pond in a grease- wood-—cheat grass—covered section of Asphalt Wash. Nets were checked at 15 minute inter- vals maximum from sunset to 0400 hr in June and August 1977-1980. Besides recording time of capture, all bats were identified to species; weighed to the nearest gram; fore- arm, ear, body (nose to base of tail), wing length, and wing width measured to the nearest millimeter; sexed; marked for recap- ture recognition; and released. Abundance was expressed as number per kilometer (no/km) for animals sampled on transects. Abundance of rodents captured in the 12 x 12 grids was calculated by dividing the actual area covered by the grid into the number of individuals captured in five nights of trapping and expressed as individuals per hectare (ind/ha). Rodent abundance from sea- sonal trapping was expressed as individuals per 100 trap nights (ind/100 TN). Bat abun- dance was expressed as individuals per trap night (ind/TN). Species richness was the num- ber of species that occurred on a transect, in a trap grid, or were mist-netted during a sam- pling period. Species diversity expressed as | H’ after Shannon and Weaver (1964) was cal- culated from data for a sampling period. Habi- tat preference and annual changes in abun- dance, species richness, and species diversity from 1975 to 1984 were tested by two-way analysis of variance (ANOVA) with means ranked by least significant differences (LSD) | test (Steel and Torrie 1960). Seasonal prefer- | ence from 1975 to 1981 was tested by one-way ANOVA and LSD. i i ! | ] July 1986 GRANT: OIL SHALE TRACT WILDLIFE 473 Ss © OO C ©& & ht ee ee eg cz O& WM OO YD © | ——— r (G.R.) ol He | h | ioe) ui I rll Hell Wee Hi Wit i! HH { nding area, Uintah County, Utah. Green Rive 4 i hy yi eg i Ht mt il Falla { E i < (ae > EE 4q ai eed ‘ip iY i ee ANA / es ill oT iy tr i oie it A he gg a vil wll a rs ul alr i a Ee mi wa wl aa ‘ ! Hae : i mS ee 474 = 1 Ses C CSS Ee S00 ° s) o° oo oC 0 224.3 590.0 oO N oO oO N oO Annual Plant Biomass, g/m SES) 1976 1977 1978 County, Utah (after NPI, Inc.). Species identification of amphibians and reptiles as well as nomenclature was done according to Stebbins (1966), also using Smith (1946) for the latter group of vertebrates. Avian identification was based on Peterson (1961), Robbins et al. (1966), and Behle and Perry (1975) and nomenclature followed AOU (1983). Mammalian identification was based on Ranck (1961), Barbour and Davis (1969), and, primarily, Armstrong (1972), whose nomenclature was also followed. An explana- tion of abbreviations used in tables and graphs are presented in Table 1. RESULTS AND DISCUSSION The number of species of amphibians and reptiles found at Ua-Ub did not differ markedly from the species previously re- corded in the Uinta Basin (Table 2). The num- ber of avian and mammalian species, how- ever, was considerably lower at Ua-Ub than in the Basin. This was due to the lack of aquatic as well as higher elevation habitats that sup- port a diversity of birds and mammals absent in the desert portion of the Basin. GREAT BASIN NATURALIST 1979 Fig. 6. Sagebrush stem leader growth and annual plant biomass at Oil Shale Tracts Ua-Ub, Uintah Vol. 46, No. 3 ee ee @-—- Greasewood #— Shadscale ~+---- Juniper Rtpartan 1980 1981 1982 1983 1984 Amphibia SALIENTIA.—Five amphibians occurred at Ua-Ub (Table 3), including a new record, a red-spotted toad (Bufo punctatus), found at a small pond in Asphalt Wash, June 1980. The other two toads were the most consistent am- phibians, occurring at stock ponds more fre- quently than along the White River. The two frogs, confined to the White River drainage, were absent more often than not. The tiger salamander (Ambystoma tigrinum) included in the Oil Shale Area by Olsen (1973) was not found at Ua-Ub. Reptilia SQUAMATA.—FEleven reptiles occurred at Ua-Ub (Table 4). The tree lizard (Urosaurus ornatus ) was a new record for the Basin. Two other species, the milk snake (Lampropeltis triangulum) and night snake (Hypsiglena tor- | guata), did not occur at Ua-Ub; however, a milk snake was found and photographed north of Bonanza, Utah, at a small stock pond east of Utah Highway 44 by T. Schultz (WRSP 1977, Final Environmental Baseline Report, SLC, | Utah), augmenting Tanner (1957). | July 1986 TABLE 1. Definition of abbreviations used in text, ta- bles, and graphs for residency status, month, vegetation (habitat) types, and presence/absence. RESIDENCY STATUS P = permanent resident: breeding documented, pre- sent year-round p = permanent resident: breeding not documented, present year-round S = summer resident: breeding documented, present during breeding season, that is, April—-October s = summer resident: breeding not documented w = winter resident: present October—April m = migrant: present during spring (April) and/or fall (October) t = transient: present for short duration and permanent or summer resident at other locales in the Uinta Basin a = accidental: present infrequently, outside of normal range ? = unknown: present frequently or infrequently for long or short durations at specific or random inter- vals VEGETATION TYPES G = Sagebrush-greasewood S = Shadscale-sagebrush J = Juniper woodland R = Riparian woodland Subscript p = pond/marsh PRESENCE/ABSENCE * Present: Species observed, captured, heard, or iden- tified by sign at one or more times during one year Species not observed, captured, heard, or identified by sign at one or more times dur- ing one year — Absent: Except for the short-horned lizard (Phryno- soma douglassi), lizards were abundant and ubiquitous. The eastern fence (Sceloporus un- dulatus) and tree lizards preferred the wooded habitats but occurred in a shallow- soiled shadscale vegetation abutting juniper woodlands on rugged topography present in western Ub and northern Ua. Where exposed sandstone was scattered, soils deeper, and juniper absent (in the shadscale vegetation in the eastern portion of Ub), both lizard species were absent. Rocks and trees served as hunt- ing sites for these sit-and-wait predators. The sagebrush lizard (Sceloporus graciosus) pre- ferred the three upland habitats over riparian vegetation. It foraged in open ground beneath the shrub canopy and in the bare interspace around shrubs. It was not found in dense grass cover and decreased in abundance from 1981 through 1983 (p = 0.05) as greater precipita- tion caused grass cover to increase. The side- GRANT: OIL SHALE TRACT WILDLIFE 475 TABLE 2. Wildlife found in the State of Utah, the Uinta Basin, and Oil Shale Tracts Ua-Ub. Number of species Utah in Uinta Basin Ua-Ub AMPHIBIA Ambystomatidae 1 1 0 Pelobatidae 2 1 1 Bufonidae 5 1 Q* Hylidae 2 1 1 Ranidae 3 1 1 13' 5a 5 REPTILIA Iguanidae 14 4 5* Teiidae 3 1 ] Colubridae 19 6 4 Viperidae 5 1 1 4]' 1 ll AVES Gaviiformes 3 ] 0 Podicipediformes 8 6 0 Ciconiiformes 15 8 3 Anseriformes 37 25 14 Falconiformes 22 17 15 Galliformes 11 9 3 Gruiformes wh 6 3 Charadriiformes 53 36 6 Columbiformes 6 3 1 Cuculiformes 3 0 1* Strigiformes 12 12 4 Caprimulgiformes 4 2 2 Apodiformes 11 5 4 Coraciiformes 1 1 1 Piciformes 11 fe) 4 Passeriformes 186 125 94* 390° 265° 155 MAMMALIA Insectivora 4 2 0 Chiroptera 17 11 Oe Lagomorpha ii 6 2 Rodentia 61 31 20* Carnivora 23 11 10 Artiodactyla 6 5 3 118° 66"° 44 *New record for the Uinta Basin ‘Stebbins 1966 ?Hayward et al. 1958, Olsen 1973 3Behle and Perry 1975, Hayward et al. 1976 ‘Behle 1981 °Durrant 1952 ®Ranck 1961, Olsen 1973 blotched lizard (Uta stansburiana) foraged in the extensive bare ground and _ horizontal sandstone surfaces found in shadscale and ju- niper vegetation and was not found in the grassy areas. The western whiptail (Cnemi- dophorous tigris) demonstrated no habitat preference but was most often associated with greasewood, rabbitbrush, and sagebrush, 476 GREAT BASIN NATURALIST Vol. 46, No. 3 TABLE 3. Amphibian residency status, consistency, spatial distribution, and abundance in four vegetation types at Oil Shale Tracts Ua-Ub. Abbreviations are explained in Table 1. Spatial distribution and abundance ORDER 1975-1984 Family Common name Residency Years Month Vegetation Number/ Genus species status present type Kilometer + SD SALIENTIA Pelobatidae Great Basin Spadefoot 2 June Gp * Scaphiopus intermontanus Sp = R * Bufonidae Woodhouse’s Toad R June Gp Bufo woodhousei Sp R Red-spotted Toad ? June Gp * Bufo punctatus Hylidae Chorus Frog P June R os Pseudacris triseriata Ranidae Leopard Frog 2 June R * Rana pipiens where it foraged actively in the litter at a shrub’s base. The whiptail lizard also avoided grassy areas. During periods of high grass and forb cover, both the side-blotched and whiptail lizards were low (1975 and 1983) compared to peak abundance during the 1977 drought (p = 0.05). The eastern fence and tree lizards, both at low abundance during the 1977 drought, increased in 1983 (p S$ 0.05). Aves CICONIIFORMES AND ANSERIFORMES. —Sev- enteen species of herons and waterfowl oc- curred along the White River (Table 5). Only the great blue heron (Ardea herodias) and Canada goose (Branta canadensis) spent much time along the river. The herons for- aged in the area surveyed in this study, but no nests were found. Abundance was lowest in April (Table 6). The geese nested on sandbars along the river banks and in sandstone cliffs separating riparian vegetation from shadscale vegetation. Adult geese were at peak abun- dance during April (Table 6). There were un- published reports (J. Grandison, UDWR, Vernal, Utah, verbal communication) that these geese flew into southern Wyoming by August, then returned by October. These data support Grandison in that the only time geese stayed along the White was during the 1977 drought. FALCONIFORMES.—Fifteen species of raptors occurred at Ua-Ub (Table 5). Most were ob- served during the nesting season in February and April (Table 6). Three raptors were perma- nent residents; of these, the golden eagle was the most abundant. During 1983 seven pairs of golden eagles successfully nested in the sand- stone cliffs within 2 km (1.2 mi) perimeter of © Ua-Ub. Two additional nests outside the perimeter were also successful in 1983. Five rap- tors were summer residents, and the red-tailed hawk (Buteo jamaicensis ) and American kestrel (Falco sparvarius) were most abundant. Also during 1983 seven pairs of red-tailed hawks suc- cessfully nested in the same area as the eagles and two other successful red-tailed hawk nests occurred outside the perimeter. Most of the nests were in sandstone cliffs, except one, which occurred in a cottonwood tree along the White River. Kestrels were also abundant during 1976, 1982, and 1983, years of high rodent abundance. Red-tailed hawks and eagles both increased as rabbits increased in 1983; both eagles and rab- bits decreased in 1984. The hawks also de- creased in 1984, but not as drastically because they used other prey resources. Among the three winter residents, the goshawk (Accipiter gentilis) arrived at Ua-Ub July 1986 GRANT: OIL SHALE TRACT WILDLIFE 477 TABLE 4. Reptilian residency status, consistency, spatial distribution, and abundance in four vegetation types at Oil Shale Tracts Ua-Ub. Abbreviations are explained in Table 1. Letters after abundance indicate differences at p = 0.05. ORDER Family Common name Genus species SQUAMATA Iguanidae Eastern Fence Lizard Sceloporus undulatus Sagebrush Lizard Sceloporus graciosus Side-blotched Lizard Uta stansburiana Tree Lizard Urosaurus ornata Short-horned Lizard Phrynosoma douglassi Teiidae Western Whiptail Cnemidophorus tigris Colubridae Racer Coluber constrictor Striped Whipsnake Masticophis taeniatus Gopher Snake Pituophis melanoleucus Western Terrestrial Garter Snake . Thamnophis elegans Viperidae Western Rattlesnake Crotalus viridis lEvacuation Creek exclusively Residency status Spatial distribution and abundance 1975-1984 Years Month Vegetation Number/ present type Kilometer + SD 10 June 10 June =) Ne) I+ i=) D ° 10 June — io) I+ re eee. poop 10 June S e I+ SoS — oo of 9 June 10 June MD Dan AnD DPHnO DHHO DPHAHO D-4O ) oo I+ S on 9 June 9 June 10 June A= NO Aan * bo — Cc =) (a>) ee) i=) S — 10 June G OsIR==s05l D4 WD oS (=) — in October (Table 6) and roosted mainly in _ lands north and west of Ua-Ub. The ferrugi- cottonwoods along the White River (Table 5). | nous hawk (Buteo regalis), infrequently found The rough-legged hawk (Buteo lagopus), at Ua-Ub, nested at low abundance and was a though few in number at Ua-Ub, was numer- permanent resident during 1983 in the area ous and consistently found in the open shrub- north and west of Ua-Ub. The bald eagle 478 GREAT BASIN NATURALIST Vol. 46, No. 3 TABLE 5. Avian residency status, consistency, spatial distribution, and abundance in four vegetation types at Oil Shale Tracts Ua-Ub. Abbreviations are explained in Table 1. Letters after abundance indicate differences at p < 0.05. ORDER Family Common name Genus species CICONIIFORMES Ardeidae Great Blue Heron Ardea herodias Snowy Egret Egretta thula Black-crowned Night Heron Nycticorax nycticorax ANSERIFORMES Anatidae Canada Goose Branta canadensis Green-winged Teal Anas crecca Mallard Anas platyrhynchos Pintail Anas acuta Blue-winged Teal Anas discors Cinnamon Teal Anas cyanoptera Northern Shoveler Anas clypeata Gadwall Anas strepera American Wigeon Anas americana Ring-necked Duck Aythya collaris Lesser Scaup Aythya affinis Bufflehead Bucephala albeola Common Merganser Mergus merganser Red-breasted Merganser Mergus serrator FALCONIFORMES Cathartidae Turkey Vulture Cathartes aura Accipitridae Bald Eagle Haliaeetus leucocephalus Northern Harrier Circus cyaneus Residency status m Spatial distribution and abundance Years present 10 10 10 10 10 10 1975-1984 Month Vegetation type June R April R April R April R April Gp R April Gp R April R April R ‘April R April R April R April R April R April R April R April R April R April G S J R February S R April G S R Number/ Kilometer + SD 0.01 2.0 + 1.0 I+ * 0.2 + 0.3 * 0.1 + 0.2 0.4 + 0.7 0.2 + 0.2 4 | Te ee July 1986 Table 5 continued. ORDER Family Common name Genus species Sharp-shinned Hawk Accipiter striatus Cooper's Hawk Accipiter cooperii Northern Goshawk Accipiter gentilis Swainson’s Hawk Buteo swainsoni Red-tailed Hawk Buteo jamaicensis Ferruginous Hawk Buteo regalis Rough-legged Hawk Buteo lagopus Golden Eagle Aquila chrysaetos Falconidae American Kestrel Falco sparverius Merlin Falco columbarius Peregrine Falcon Falco peregrinus Prairie Falcon Falco mexicanus GALLIFORMES Phasianidae Chukar Alectoris chukar Sage Grouse Centrocercus urophasianus Ring-necked Pheasant Phasianus colchicus | GRUIFORMES Rallidae Virginia Rail Rallus limicola Gruidae Sandhill Crane Grus canadensis GRANT: OIL SHALE TRACT WILDLIFE Residency status ie S,P 5,P Spatial distribution and abundance Years present fl 10 10 10 10 10 1975-1984 Month Vegetation type June J R April G J R February J R August G S June G S J R October J February G S J R February G S J R June G S J R August R April G R June G S J R June G R April S June G R June Gp April R 479 Number/ Kilometer + SD * fo) S oS of fp 0.02 480 GREAT BASIN NATURALIST Vol. 46, No. 3 Table 5 continued. Spatial distribution and abundance ORDER 1975-1984 Family Common name Residency Years Month Vegetation Number/ Genus species status present type Kilometer + SD Whooping Crane m 1 October Flying = Grus americana over Ua-Ub CHARADRIIFORMES Charadriidae Killdeer S 10 June Charadrius vociferus mo, * Recurvirostridae American Avocet m 3 April R ss Recurvirostra americana Scolopacidae Greater Yellowlegs m 4 April R 0.01 Tringa melanoleuca Solitary Sandpiper s 2 June R * Tringa solitaria Spotted Sandpiper S 10 June R 0.9 + 0.8 Actitis macularia Common Snipe m 1 April R - Gallinago gallinago COLUMBIFORMES Columbidae Mourning Dove S 10 June Zenaida macroura mn) OOS OR It It I+ I+ aS DN QDwA ~ CUCULIFORMES Cuculidae Yellow-billed Cuckoo s 2 June R Coccyzus americanus STRIGIFORMES Strigidae Western Screech-Owl t 2 April Otus kennicottii J R Great Horned Owl P 10 June G Bubo virginianus S J R R S S xa) I+} * * * 0.2 + 0.3 Long-eared Owl P 6 April Asio otus Short-eared Owl t 4 April S Asio flammeus J R a CAPRIMULGIFORMES Caprimulgidae Common Nighthawk S 10 June Chordeiles minor It it I+ I+ ooso bo Ol bo ooos bo Now * Common Poorwill S 10 June Phalaenoptilus nuttallii APODIFORMES Apodidae White-throated Swift S 10 June G Aeronautes saxatalis S 0.2 + 0.4 J R July 1986 GRANT: OIL SHALE TRACT WILDLIFE 481 Table 5 continued. Spatial distribution and abundance ORDER 1975-1984 Family Common name Residency Years Month Vegetation Number/ Genus species status present type Kilometer + SD Trochilidae Black-chinned Hummingbird S 6 June G 0.01 Archilochus alexandri ij 0.2 + 0.5 R 0.01 Broad-tailed Hummingbird S 8 June G 0.01 Selasphorus platycercus S 0.01 J 0.01 R OF EOk3 Rufous Hummingbird m 1 August R o Selasphorus rufus CORACIIFORMES Alcedinidae Belted Kingfisher t 8 April R 0.05 Ceryle alcyon PICIFORMES Picidae Yellow-bellied Sapsucker t 3 October R 0.3 + 0.4 Sphyrapicus varius Downy Woodpecker P 10 June R 0.3 + 0.5 Picoides pubescens Hairy Woodpecker p 10 June R 0.06 Picoides villosus Northern Flicker 2 10 June G 0.04 Colaptes auratus S 0.01 J 0.03 R 0.9 + 0.6 PASSERIFORMES Tyrannidae Olive-sided Flycatcher m 1 August R 0.02 Contopus borealis Western Wood Peewee S 10 June G Contopus sordidulus J 0.07 R 0.4 + 0.5 Willow Flycatcher S 10 June R 0.5+ 0.5 : _ Empidonax traillii Gray Flycatcher S 10 June (0), 0} ce (OLR Al Empidonax wrightii 13+0.5 b : 0.02 a Eastern Phoebe m 3 August ee . Sayornis phoebe - { * Say’s Phoebe S 10 June 0.3 + 0.2 Sayornis saya Ash-throated Flycatcher S 10 June Myiarchus cinerascens S =) oo) of Pp Pp | Western Kingbird S 9 June Tyrannus verticalis Dn Dong Don BPO B-O —) bo I+ —) bo Fatt 482 GREAT BASIN NATURALIST Vol. 46, No. 3 Table 5 continued. Spatial distribution and abundance ORDER 1975-1984 Family Common name Residency Years Month Vegetation Number/ Genus species status present type Kilometer + SD Eastern Kingbird s 3 June G 0.01 Tyrannus tyrannus R 0.03 Alaudidae Horned Lark P 10 June G = Eremophila alpestris S 0.01 Hirundinidae Tree Swallow t 3 June Gp s Tachycineta bicolor J : R * Violet-green Swallow S 10 June G O Jb = 0.2 Tachycineta thalassina S 0.04 J 0.2+0.4 R 0.04 Northern Rough-winged s 6 June S * Swallow J 0.02 Stelgidopteryx serripennis R 0.04 Cliff Swallow S 10 June G 0.02 Hirundo pyrrhonota S O37 25 1.83 J 0.07 R 30.4 + 33.0 Barn Swallow m,s 10 June Stas z Hirundo rustica J : R 0.01 Corvidae Scrub Jay P 8 June G < Aphelocoma coerulescens J 0.01 R * Pinyon Jay 2 10 April G DO ae BS 2 Gymnorhinus cyanocephalus S 0.09 a ] 7.2=56 b Clark’s Nutcracker t 3 April J = Nucifraga columbiana R = Black-billed Magpie P 10 June G 0.4+0.6 a Pica pica S 0.01 a ] 02+03 a R 14 eSaleoieb Common Crow t 4 April G 0.01 Corvus brachyrhynchos R 0.01 Common Raven P 10 February G 0.6 + 0.6 Corvus corax S 0.04 J 0.2 + 0.2 R 0.3 + 0.2 Paridae Black-capped Chickadee P 10 June G * Parus atricapillus J * R 0.7 + 0.5 Mountain Chickadee w 6 October G < Parus gambeli J 0.08 R 0.2 + 0.5 Plain Titmouse P 10 June G 0.1 + 0.2 Parus inornatus J 0.4 + 0.5 Aegithalidae Bushtit Ww 6 February J ° Psaltriparus minimus R a July 1986 Table 5 continued. ORDER Family Common name Genus species Sittidae Red-breasted Nuthatch Sitta canadensis White-breasted Nuthatch Sitta carolinensis Certhiidae Brown Creeper Certhia americana Troglodytidae Rock Wren Salpinctes obsoletus Canyon Wren Catherpes mexicanus Bewick’s Wren Thryomanes bewickii House Wren Troglodytes aedon Marsh Wren Cistothorus palustris Muscicapidae Ruby-crowned Kinglet Regulus calendula Blue-gray Gnatcatcher Polioptila caerulea Black-tailed Gnatcatcher Polioptila melanura Western Bluebird Sialia mexicana Mountain Bluebird Sialia currucoides Townsend's Solitaire Myadestes townsendi Swainson’s Thrush Catharus ustulatus Hermit Thrush Catharus guttatus American Robin Turdus migratorius GRANT: OIL SHALE TRACT WILDLIFE Residency status t,w S,P Spatial distribution and abundance Years present 4 10 10 10 10 10 1975-1984 Month Vegetation type August J R August R October R June G S J R June G S J R June G J R June R June Gp October G J R June G S J R June G J R April R June G S J R April G J R October R October R June G J Number/ 483 Kilometer + SD 0.01 0.04 0.1 + 0.2 ) bo I+ S bo I+ + i+ I+ i Go Ul Ol Oe S XS a) — S a) S 0.2 *|+ * S ioe) 0.8 I+ 0.8 og 25 OLY) oon an) 484 Table 5 continued. ORDER Family Common name Genus species Mimidae Gray Catbird Dumetella carolinensis Northern Mockingbird Mimus polyglottos Sage Thrasher Oreoscoptes montanus Bendire’s Thrasher Toxostoma bendirei Motacillidae Water Pipit Anthus spinoletta Bombycillidae Cedar Waxwing Bombycilla cedrorum Laniidae Northern Shrike Lanius excubitor Loggerhead Shrike Lanius ludovicianus Sturnidae European Starling Sturnus vulgaris Vireonidae Gray Vireo Vireo vicinior Solitary Vireo Vireo solitarius Warbling Vireo Vireo gilvus Red-eyed Vireo Vireo olivaceus Emberizidae Orange-crowned Warbler Vermivora celata Virginia's Warbler Vermivora virginiae Yellow Warbler Dendroica petechia Yellow-rumped Warbler Dendroica coronata Black-throated Gray Warbler Dendroica nigrescens GREAT BASIN NATURALIST Spatial distribution and abundance 1975-1984 Residency Years Month Vegetation status present type ? 1 June R S 3) June G S R S 10 June G S J ip 1 June S m 3 April S R s iit June R w 3 February G S R - PS 10 June G S J S 10 June G S J R S 1 June J S 8 June J R S 9 June ? 2 June R s 6 June R s 2 June R S 10 June R m,s 10 October G J R S 10 June G J R Vol. 46, No. 3 Number/ Kilometer + SD 0.1 + 0.2 0.5 £ 0.7 O0.1+0.1 a 183+ sk2b 0.04 a July 1986 GRANT: OIL SHALE TRACT WILDLIFE 485 Table 5 continued. Spatial distribution and abundance ORDER 1975-1984 Family Common name Residency Years Month Vegetation Number/ Genus species status present type Kilometer + SD Townsend's Warbler m 2 August R 0.2 + 0.4 Dendroica townsendi MacGillivray s Warbler s 3 June R 0.03 Oporornis tolmiei Common Yellowthroat s 5 June R 0.04 Geothlypis trichas Wilson’s Warbler m 5 October R 0.04 Wilsonia pusilla Yellow-breasted Chat S 10 June R il = O63 Icteria virens Western Tanager S 9 June G 0.01 Piranga ludoviciana J 0.05 R ee} 22 1a Rose-breasted Grosbeak P 2 June J * Pheucticus ludovicianus R 0.01 Black-headed Grosbeak NS) 0 June G 0.01 Pheucticus melanocephalus R 0.4 + 0.4 Blue Grosbeak S 8 June R OFS 015 Guiraca caerulea Lazuli Bunting S 8 June G 0.01 Passerina amoena R 1B 28 IES} Green-tailed Towhee m 4 April G 0.03 Pipilo chlorurus R 0.09 Rufous-sided Towhee S 10 June G 0.02 Pipilo erythrophthalmus J = R 3.1 + 1.4 American Tree Sparrow Ww 2 February R' O57 a=: dats} Spizella arborea Chipping Sparrow S 10 June G OVAEEON/ meal Spizella passerina J 10+0.8 b R 0.09 a Brewer s Sparrow S 10 June G DO) se Oak Bh Spizella breweri S O22 110 |o J 0.1+0.2 b R 0.2+0.5 b Vesper Sparrow m 9 April G 0.4 + 0.9 Pooecetes gramineus S 0.2 + 0.3 Lark Sparrow S 10 June G OF5EELOloerac Chondestes grammacus S ile O68) 10 J 0.2403 a R 0.9+0.7 be Black-throated Sparrow S 10 June G 19+0.6 a Amphispiza bilineata S IL 2 (0.7 |p J 0.7+0.6 b R 0.04 c Sage Sparrow S 10 June € 0.3+0.2 a Amphispiza belli S 1.8 Bs 12 b J 486 GREAT BASIN NATURALIST Vol. 46, No. 3 Table 5 continued. Spatial distribution and abundance ORDER 1975-1984 Family Common name Residency Years Month Vegetation Number/ Genus species status present type Kilometer + SD Savannah Sparrow m 1 June S s Passerculus sandwichensis Song Sparrow p ) June Gp = Melospiza melodia R 0.01 White-crowned Sparrow m,s 10 April G 0.2 + 0.4 Zonotrichia leucophrys S * J * R 0.6 + 1.0 Dark-eyed Junco Ww 10 April G 44+6.4 Junco hyemalis S OIL se O,2) ] 4.2 +59 R 3.4 + 5.8 Red-winged Blackbird S 10 June G 0.04 Agelaius phoeniceus R 0.06 Western Meadowlark S 10 June G 0.9+0.9 a Sturnella neglecta S Meeya= ILO 10) J 0.02 c R - 0.3+0.3 ¢ Brewer s Blackbird s 10 June G 0.2 + 0.8 Euphagus cyanocephalus S 0.02 J 0.1 + 0.2 R 0.1 + 0.3 Brown-headed Cowbird S 10 June G 0.01 a Molothrus ater J 0.6+0.8 a R 44+1.7 b Northern Oriole S 10 June R PAN) se 11.83 Icterus galbula Scott's Oriole S 8 June G 0.08 Icterus parisorum J 0.02 Fringillidae Rosy Finch Ww 9 February G 1.5 + 4.0 Leucosticte arctoa S 0.1 + 0.4 J 1.7+4.5 R 4.8 + 12.6 House Finch S,P 10 June G 0.5+0.4 a Carpodacus mexicanus S 0.4+0.7 a J 25+1.6 a R Gls BO lo Pine Siskin w 4 April G 0.04 Carduelis pinus S ee J * R * Lesser Goldfinch P 1 R * Carduelis psaltria American Goldfinch S 8 June J 0.01 Carduelis tristis R 0.3 + 0.5 Evacuation Creek exclusively July 1986 GRANT: OIL SHALE TRACT WILDLIFE 487 TABLE 6. Avian seasonal distribution and abundance in four vegetation types at Oil Shale Tracts Ua-Ub. Abbrevia- tions are explained in Table 1. Seasonal distribution and abundance 1975-1981 Number/Kilometer + SD Wee teat OT = fa eee ne rete ew a Re Great Blue Heron R — * < (Nil <0.1 <0.1 Canada Goose R 0.3 + 0.8 Mell se MO 0.2 + 0.4 0.2 + 0.5 O67 se ilk Green-winged Teal R a 0.2+ 0.3 <0.1 << 081 — Mallard R OFIE=TOF2 zs <0.1 ms Cinnamon Teal R — <0.1 <0.1 — — Common Merganser R << (MII 0.1 + 0.3 — — * Turkey Vulture G — * * ee, S — <) I+ S ~] rot) — oo I+ — of 42+26 a 16.9 + 10.9b 0.06 0.1+0.3 0.2 + 0.2 oP * 14+ 1.9 * i) x] I+ to fon) pop July 1986 GRANT: OIL SHALE TRACT WILDLIFE 497 Table 7 continued. Spatial distribution and abundance ORDER 1975-1984 Family Common name Residency Years Month Vegetation Individuals/ Genus species status present type trap night + SD Bushy-tailed Woodrat 2 10 August G * Neotoma cinerea S ct i O37 s= 11) R 1.4 + 2.0 Montane Vole P 4 August G * Microtus montanus S OME==053 R Shi) as! 1/609) Muskrat P 10 August R ** Ondatra zibethicus Erethizontidae Porcupine IP 10 August G a Erethizon dorsatum J * R (0.02) Number/ Kilometer + SD | CARNIVORA Canidae Coyote P 10 August G * | Canis latrans S * j J | R ‘ i Red Fox ip 1 August S = Vulpes vulpes ; Gray Fox P 1 October IR o Urocyon cinereoargenteus ( Procyonidae Ringtail ? 1 April J _ } Bassariscus astutus | Raccoon P 7 August R re \ Procyon lotor { Mustelidae Long-tailed Weasel t 2 June J - Mustela frenata Badger 2 9 August G cs Taxidea taxus S * J * R * Striped Skunk 1p 9 August R 0.01 Mephistis mephistis Felidae Mountain Lion t 1 June S = Felis concolor Bobcat P 4 August G “ Lynx rufus S : J * R * )ARTIODACTYLA Cervidae Mule Deer P 10 August G 0.05 Odocoileus hemionus S 0.1 + 0.2 J 0.1 + 0.2 R OFA E1033 498 GREAT BASIN NATURALIST Vol. 46, No. 3 Table 7 continued. Spatial distribution and abundance ORDER 1975-1984 Family Common name Residency Years Month Vegetation Individuals/ Genus species status present type trap night + SD Antilocapridae Pronghorn t 2 August S * Antilocapra americana R Bovidae Bighorn Sheep t 1 April S * Ovis canadensis Domestic Sheep Ww 10 February G 13.5 + 22.9 Ovis aries S a ] 15.8 + 40.7 R 15.0 + 24.7 Domestic Cattle S 10 August G * Bos taurus S 0.06 R 4.5 + 3.9 not cite the record. The fringed bat (M. thysanodes) was also recorded for the Basin with no source in Hasenyager (1980), and the only other record was Krutszch and Heppen- stall (1955) from the Book Cliffs at the south- ern boundary of the Basin. One of the most abundant bats was the western pipistrelle (Pipistrellus hesperus), the smallest bat at Ua-Ub, weighing 4 + 0.4 g. It was a permanent resident that was en- countered in all seasons and was the only spe- cies to increase in abundance during August. During the 1977 drought the western pip- istrelle and the silver-haired bat (Lasionyc- teris noctivagans) were the only species to occur in high abundance. Silver-haired bats, weighing 10 + 1 g, were migrants. Usually the June population was predominantly male; however, females ac- companied the males during 1978 and 1980. During August, only three males were cap- tured in four years. This bat, like the hoary bat, migrates northward in segregated groups, pregnant females preceding the males (Barbour and Davis 1969). The largest and most abundant species was the hoary bat, femaies being larger than males, 26 + 3 gand 24 + 3g, respectively (p =< 0.05), and pregnant females the largest at 34 + 3g(p. = 0.05). During June the popula- tion was usually dominated by males (97%) from 1977 to 1979. During August 1977-1979 the sex ratio was even (49% male). During June 1980 the ratio changed to 65% male. All 13 females trapped during June 1980 were pregnant. During August 1980 the ratio in- creased to 86% male, 50% of these subadults. Only three females were observed. None dis- played the usual signs of copulation. Pregnant females were thought to occur in | the plains for parturition, join the males in the Intermountain West in late summer, and breed at some unknown time in the fall prior to migrating into the southwestern states and northern Mexico for the winter (Findley and | Jones 1964). At Ua-Ub breeding occurred in | August as noted by Merriam (1884) and Tenaza (1966); males and females were not segregated during parturition, which in 1980 | occurred in the Intermountain West. | LAGOMORPHA.—Only two species of lago- | morph occurred at Ua-Ub, the desert cotton- tail (Sylvilagus audubonii) and black-tailed jackrabbit (Lepus californicus ) (Table 7). The white-tailed jackrabbit (L. townsendii) oc- | curred near Bonanza, Utah, but sightings | were rare over a 10-year period. Cottontails were the most numerous mammal encoun- | tered on transects. Although no habitat pref- | erence was evident, annual changes reflected the cyclic nature of lagomorph populations (Fig. 8). Abundance during 1976 and 1983 exceeded all other years (p S 0.01), and 1977-1978 and 1982 exceeded abundance during 1979-1981 and 1984 (p S 0.05). The rapid increase in 1976 following a 1975 desert bloom kept the cottontail population at a high level through the 1977 drought. The severe 499 GRANT: OIL SHALE TRACT WILDLIFE July 1986 ‘says uvLIedu—-y AA ‘sos 1odrun{—[AQ ‘Sozs ofeospeys—S AA ‘SoS poomosevais 1334 001 1WAY3SLNI HNOLNOD S31IW NI31V9OS ~0€, 200601 ai) ANGE yea, ‘Ayunog yew “qQ- BY SORA, B[PYS [IO Ye suoneoo] Surpdwres ofp PAA “2 “BL 20E.ZL SL Se . t t) S¥313W01I> NI31V9S aLnouaivis (_) aaAOudWINn 30V4UNS G3AOEdWI. —— savou pug doy, joasuns, A Wy ee “JONNY OHIUM uo Aaains ajb3 pjog PUD JO}dDyY ‘|MOJJaLDM ~0E,LG ———_— -— | % t ONDNIDAT w —— u Je 12M SED 9/ t \ \ cS pie ua Canyon| a> 0€, LOc6OL 498 Table 7 continued. GREAT BASIN NATURALIST Vol. 46, No. 3 Spatial distribution and abundance ORDER 1975-1984 Family Common name Residency Years Month Vegetation Individuals/ Genus species status present type trap night + SD Antilocapridae Pronghorn t 2 August S * Antilocapra americana R * Bovidae Bighorn Sheep t 1 April S 3 Ovis canadensis Domestic Sheep Ww 10 February G 13.5 + 22.9 Ovis aries S ~ J 15.8 + 40.7 R 15.0 + 24.7 Domestic Cattle S 10 August G 3 Bos taurus ) 0.06 R 45+ 3.9 not cite the record. The fringed bat (M. thysanodes) was also recorded for the Basin with no source in Hasenyager (1980), and the only other record was Krutszch and Heppen- stall (1955) from the Book Cliffs at the south- ern boundary of the Basin. One of the most abundant bats was the western pipistrelle (Pipistrellus hesperus), the smallest bat at Ua-Ub, weighing 4 + 0.4 g. It was a permanent resident that was en- countered in all seasons and was the only spe- cies to increase in abundance during August. During the 1977 drought the western pip- istrelle and the silver-haired bat (Lasionyc- teris noctivagans) were the only species to occur in high abundance. Silver-haired bats, weighing 10 + 1 g, were migrants. Usually the June population was predominantly male; however, females ac- companied the males during 1978 and 1980. During August, only three males were cap- tured in four years. This bat, like the hoary bat, migrates northward in segregated groups, pregnant females preceding the males (Barbour and Davis 1969). The largest and most abundant species was the hoary bat, femaies being larger than males, 26 + 3 gand 24 + 3g, respectively (p = 0.05), and pregnant females the largest at 34 + 3g(p. = 0.05). During June the popula- tion was usually dominated by males (97%) from 1977 to 1979. During August 1977-1979 the sex ratio was even (49% male). During June 1980 the ratio changed to 65% male. All 13 females trapped during June 1980 were pregnant. During August 1980 the ratio in- creased to 86% male, 50% of these subadults. Only three females were observed. None dis- played the usual signs of copulation. Pregnant females were thought to occur in | the plains for parturition, join the males in the Intermountain West in late summer, and breed at some unknown time in the fall prior to migrating into the southwestern states and northern Mexico for the winter (Findley and Jones 1964). At Ua-Ub breeding occurred in August as noted by Merriam (1884) and Tenaza (1966); males and females were not segregated during parturition, which in 1980 occurred in the Intermountain West. LAGOMORPHA.—Only two species of lago- morph occurred at Ua-Ub, the desert cotton- tail (Sylvilagus audubonii) and black-tailed jackrabbit (Lepus californicus ) (Table 7). The white-tailed jackrabbit (L. townsendii) oc- | curred near Bonanza, Utah, but sightings were rare over a 10-year period. Cottontails were the most numerous mammal encoun- tered on transects. Although no habitat pref- erence was evident, annual changes reflected the cyclic nature of lagomorph populations (Fig. 8). Abundance during 1976 and 1983 exceeded all other years (p S 0.01), and 1977-1978 and 1982 exceeded abundance during 1979-1981 and 1984 (p S 0.05). The rapid increase in 1976 following a 1975 desert bloom kept the cottontail population at a high level through the 1977 drought. The severe 499 GRANT: OIL SHALE TRACT WILDLIFE Ne) ee) op) ce iy =} — ‘sous uetedu—-y (A ‘sozs 1odrun{—[ Aq ‘sojIs apRospeys—G AA ‘SoUSs poomasvais—J) MQ “Yey~Q ‘AyUNOD YRIUIY “qn- 1334 00 IWAYSLNI HNOLNOD S3TIW NI3TVOS SS SS SS = —| t i) SH313WOTIM NI31V9OS aLnowaivis () G3AOUdWIINN ----—- 30V4YNS GIAOKAWI. —-— savou pig dos, O joasuns, VA Wy Eee “AOAIY OLIUM uo Aaasns 9j69 pjog puD JO}dDyY ‘|MOJIa}OM t & 5 ~0€,L00601 ~ 20,71 St BY) SpORAT, [CYS [IO 3 SuOHKoo] Surjdwes opi “1, “Bly 7 1A 112M S°D 9/ \ uollpnsDaz 8 yaaa Si is Canyon| ~~, N, Pr a a eS) 2— Greasewood *®--— Shadscale ~-—- Juniper Ee N sro [NY I & 10 i pearien we Wy (E o -a Ee 325 Ze, SAK O ~ IS7o 1973 I9SO 1982 1984 Fig. 8. Desert cottontail (Sylvilagus audubonii) abun- dance in four vegetation types at Oil Shale Tracts Ua-Ub, Uintah County, Utah. winter of 1978-1979 resulted in small pockets of surviving cottontails in juniper and riparian woodlands and extinction in shrub habitats from 1979 through 1980. The crash in 1984 also fol- lowed a severe winter. Seasonally, cottontails were at peak abundance during August and Oc- tober (p S 0.05) (Table 8). RODENTIA.—Twenty species of rodents oc- curred at Ua-Ub (Table 7). In the squirrel family, the least chipmunk (Eutamias minimus ) favored juniper habitat. It was seldom seen or captured in February and was most active during June and August (Table 8). It avoided shadscale habitats that lacked scattered juniper trees and plentiful sandstone outcrops. The Colorado chipmunk (E. quadrivittatus) was a rare capture in juniper (Table 8), though Ranck (1961) found them rela- tively common in similar habitat. The yellow-bellied marmot (Marmota flaven- tris) was rare until 1981, when encounters be- came frequent in the rip-rap used to construct a bridge and pipeline across the White River. The white-tailed antelope squirrel (Am- mospermophilus leucurus) densities in grease- wood and shadscale were five and four per hectare, respectively, and one per hectare in juniper from 1975 through 1976, exceeding the next eight years (p = 0.05). During the 1977 drought through 1980 no squirrels were cap- tured or seen in shadscale and juniper habitats. Visual sighting during the day occurred in February in greasewood, then from April through October after sunset or prior to dawn. The squirrel population appeared to be recover- ing in 1982; however, density declined to 0.1 per hectare in 1984. The white-tailed prairie dog (Cynomys leu- curus ) that resided in a town on the northeast- GREAT BASIN NATURALIST Vol. 46, No. 3 ern perimeter of Ub followed the same pattern as the antelope squirrel. The dog town was aban- doned from 1977 through 1980, was recolonized by individuals from larger towns to the north in 1981, then “boomed” in 1984. Two heteromyid rodents, the Apache pocket mouse (Perognathus apache) and Ord’s kanga- roo rat (Dipodomys ordii), favored greasewood and shadscale habitats (Table 7). Pocket mice were especially susceptible to cold weather, with no captures occurring in February, whereas kangaroos rats remained active through mild winters (Table 8). Annually pocket mice were little affected by drought or the 1975 bloom; however, the severe winter of 1979 may have influenced the population to a small de- gree. Ord’s kangaroo rats were not affected by the drought, but following 1979 the population be- gan to increase, reaching a peak in 1980 and 1981 (p = 0.05) (Fig. 9). Kangaroo rats invaded ripar- ian and juniper habitats, occupying sandy fluvial deposits and sandy alluvial deposits from upland flashfloods in riparian habitat and sandy alluvial | and colluvial deposits in the juniper. By 1984 | these populations were extinct and the normally stable populations in shrub habitats were also declining. The beaver (Castor canadensis), absent from | the White River in previous surveys (Ranck | 1961, Olsen 1973), resided in the river banks and was active year-round, even during winters when the river was frozen over. Young beaver was encountered in shadscale and in ponds in Asphalt Wash during August. The cricetid rodents were species of the wood- lands rather than the shrubs. The western har- | vest mouse (Reithrodontomys megalotis) and | deer mouse (Peromyscus maniculatus) pre- | ferred riparian woodlands, and the canyon mouse (P. crinitus), pinyon mouse (P. truei), and desert woodrat (Neotoma lepida) preferred juniper (Table 7). The bushytailed woodrat (N. cinerea) preferred both woodlands, and it, like the desert woodrat, was subject to periodic ex- tinction in greasewood and shadscale. All these rodents were active year-round (Table 8). Annually two basic patterns in popula- tion were demonstrated by the harvest mice (Fig. 10) and by the deer mice (Fig. 11). The harvest mice were present some years and ab- sent others, as were the canyon mice, the pinyon mice, the least chipmunks, and the montane July 1986 GRANT: OIL SHALE TRACT WILDLIFE 501 TABLE 8. Mammalian seasonal distribution and abundance in four vegetation types at Oil Shale Tracts Ua-Ub. Abbreviations are explained in Table 1. Species Desert Cottontail Least Chipmunk Colorado Chipmunk White-tailed Antelope Squirrel Golden-mantled Ground Squirrel Apache Pocket Mouse Ord’s Kangaroo Rat Beaver Western Harvest Mouse Canyon Mouse Deer Mouse Pinyon Mouse Desert Woodrat Bushy-tailed Woodrat Vegetation type BH nA DHnn BHO BHHO BPHH BHHO DF PHHO FPAHYO 27% DAS NO om DHnD DanNOH February Seasonal distribution and abundance 1975-1981 Individuals/100 trap nights + SD (Number/Kilometer + SD) 9.4 + 6.3 12.0 + 12.4 6.1 + 6.0 7.8 + 4.1 0.4+1.1 = N I+ — oo Go} a8 Shy) August October 0.9 + 1.2 4b 2s JE) 12.1 + 10.3 13.1 + 8.4 9.2 + 6.3 142 eE7e9 <0.1 Do ae 0.1 + 0.3 <0.1 502 Table 8 continued. Vegetation Species type February Porcupine J — R (0.8 + 0.6) Mule Deer G (< 0.1) S (< 0.1) i (0.1 + 0.2) R ae Domestic Sheep G (13.5 + 22.9) S * J (15.8 + 40.7) R (15.0 + 24.7) Domestic Cattle R e Evacuation Creek exclusively #— Greasewood e--- Shadscale -—- Juniper fs) = 15 * +--- Riparian = =) 2 10 > me) as) FESSOR IS Ze 84 1978 Fig. 9. Ord’s kangaroo rat (Dipodomys ordii) density in four vegetation types at Oil Shale Tracts Ua-Ub, Uintah County, Utah. SIGS voles (Microtus montanus). The pinyon mice seemed more like an invader from southerly pinyon-juniper woodlands during years of ovér- all high rodent productivity. Both types of woodrats followed the same pat- tern as the deer mice: high density in 1976, a crash in 1977, and a slow recovery and median densities in upland habitats. Deer mouse den- sity in riparian habitat was possibly skewed up- ward by movement from shadscale into riparian habitats..The reasons for this movement were possibly wet soils in riparian, plus a lack of aboveground nest sites. In northwestern Colo- rado the deer mice moved more than 1 km in one week between dry soils covered by grasses and willow-dominated riparian vegetation (C. V. Grant, unpublished data). GREAT BASIN NATURALIST Vol. 46, No. 3 Seasonal distribution and abundance 1975-1981 Individuals/100 trap nights + SD (Number/Kilometer + SD) April June August October rie oe * * (0.4 + 0.4) (< 0.1) (< 0.1) (< 0.1) — — (< 0.1) — (0.2 + 0.3) (<0.1) (<0.1) def (< 0.1) (< 0.1) (< 0.1) — On20e) TOL200) O5209 7207) * coe! — — (54.3 + 76.6) ES — — * ewe a — * * * * zt ze (6.1 + 3.6) (8.0 + 8.8) Three other rodents of note were the montane vole, the muskrat (Ondatra zibethicus ), and the porcupine (Erethizon dorsatum). The vole was new for this portion of the Uinta Basin (Durrant ~ 1952). In 1981 the first vole was trapped along ~ the White River. By 1982 vole density was 22 individuals per hectare on one alluvial fan, but no sign was present elsewhere along the river. By 1983 voles were present or trapped on every alluvial fan we visited, plus individuals were seen in greasewood habe 4 km south of the river in the Southam Canyon. By 1984 there were no signs nor captures of voles along the White River, but some were captured and more observed in shadscale habitat, 3 kin south of the White River and 2 km east of Evacuation Creek at sampling site S1 (See Fig. 7). Muskrats, previously unrecordéd in these reaches of the White River, were not numerous but consistent. Porcupines were also few in ~° number during summer (Table 7); peak abun- dance occurred during winter (Table 8) when they fed on the upper cottonwood canopy. CARNIVORA.—Ten species of carnivore oc- curred at Ua-Ub (Table 7). The striped skunk, (Mephistis mephistis), coyote (Canis latrans), raccoon (Procyon lotor), and badger (Taxidea taxus) were the only consistent carnivores at Ua-Ub. None were abundant; only one striped skunk was encountered on transect. In areas where the prey base was more stable, carnivores — were not only encountered on transects, but weasels (Mustela spp.) were captured in live traps (C. V. Grant, unpublished data). - 7 } July 1986 10 #— Greasewood e-—- Shadscale “-—- Juniper +--- Riparian Individuals/ha oO -_ 1984 fp? SIGS 1980 Fig. 10. Western harvest mouse (Reithrodontomys megalotis) density in four vegetation types at Oil Shale Tracts Ua-Ub, Uintah County, Utah. 1978 1982 ARTIODACTYLA.—Mule deer (Odocoileus hemionus) occurred at low abundance at Ua- Ub (Table 7). They used both the juniper and shadscale during winter and early spring, then most deer moved into riparian for the | remainder of the year. Pronghorn (Antilo- capra americana) entered the boundary of Ua-Ub north of the White River on only two | occasions, once during the 1977 drought and again in 1983. Bighorn sheep (Ovis canaden- sis) were encountered in high sandstone cliffs separating Asphalt Wash and Southam ‘Canyon in April 1983. In June 1983 a group of | three ewes was seen in high sandstone cliffs north of the White River at Ignatio Stage Stop. These sightings are the only living | sheep reported in the Basin. Two other mammals that are seldom con- ' sidered in wildlife accounts yet play a major role in forage quantity and quality were do- ) mestic livestock. Domestic sheep (Ovis aries ) / were winter residents of Ua-Ub from Decem- ) ber through April (Table 7). One flock of about / 2,000 wintered in and around greasewood, juniper, and riparian habitats of Ua. This flock moved north of the White River in March, but at least two other flocks moved into Ub shad- scale habitat from March through shearing in _ late April (Table 8). Approximately 400 cattle were released in riparian habitat in late June to forage along the river bottom lands through October (Table 7, 8). During 1977 and again in 1983, cattle left |the river bottom to graze in the uplands. In 1977 little forage remained in riparian habitat, ‘and in 1983 grasses were plentiful in the up- lands. GRANT: OIL SHALE TRACT WILDLIFE 503 40 ®— Greasewood ®-— Shadscale =—- Juniper 30 +--- Riparian 20 (2) Individuals/ha 1974S 193 1980 19832 1984 Fig. 11. Deer mouse (Peromyscus maniculatus) den- sity in four vegetation types at Oil Shale Tracts Ua-Ub, Uintah County, Utah. #— Greasewood e-- Shadscale “-—- Juniper +--- Riparian Number/km 1984 1982 197s 1978 19Se Fig. 12. Reptilian abundance in four vegetation types at Oil Tracts Ua-Ub, Uintah County, Utah. POPULATION DYNAMICS The few amphibian species did not reflect any general trends, except that Ua-Ub habitat does not favor an abundant or diverse amphibian fauna. Among the reptiles the snake component was also limited in number but, for the most part, consistently encountered from year to year. Lizards were more abundant, perhaps the most abundant vertebrates at Ua-Ub. Abun- dance measured for all vertebrates included those observations made at zero point on the transect line. For birds and mammals, distance to the farthest observation was unlimited, but usually few individuals were seen beyond 100 m. For lizards maximum observable distance from the transect was 4 m. Annual lizard population dynamics were mixed (Fig. 12). Although there were seemingly large changes, neither abundance, richness, nor diversity changed significantly from year to year. 904 GREAT BASIN NATURALIST Vol. 46, No. 3 TABLE 9. Spatial distribution of reptiles, birds, and mammals according to abundance, species richness, and species diversity at Oil Shale Tracts Ua-Ub. Letters signify differences at p < 0.05. Vegetation Abundance Richness Diversity type +SE +SE +SE REPTILES June 1975-1983 Number/Kilometer Number of species H’ Greasewood 8 ae O08 a 4+ 0.4 1.02 + 0.09 Shadscale 6+ 1.0 b 4+ 0.6 1.14 + 0.18 Juniper 7+0.9 b 5+ 0.2 1.33 + 0.03 Riparian 4+0.6a 5 + 0.6 1.31 + 0.12 BirDs June 1975-1984 Number/kilometer Number of species H’ Greasewood 16+ 20a 21+1.0a 2.41 + 0.08 Shadscale 13+ 10a 16+1.0b 2.22 + 0.08 a Juniper SR =e 28Olra 23 =) 1.0¥a 2.60 + 0.06 b Riparian 82 + 15.0 b 44+2.0c 2.59 + 0.14 b MAMMALS (Lagomorpha, Rodentia, Carnivora, Artiodactyla) August 1975-1984 Number/Kilometer Number of species H’ Greasewood 3+ 1.0 2+0.3a 0.18 + 0.07 a Shadscale 2+ 0.6 2+ 0.3 0.33 + 0.10 Juniper 3 + 0.7 3+ 0.5 0.45 + 0.11 Riparian 4+ 2.0 3+ 0.5 b 0.60 + 0.14 b RODENTS August 1975-1984 Individuals/hectare Number of species H’ Greasewood 18 + 3.0 5+0.5a & 1.15+0.10 a Shadscale 15+3.0a 5 + 0.5 ac 1.20 + 0.07 a Juniper Ig} a2 8.0) @ = 0b Nestle O18) |p Riparian 25 + 6.0 b 4+0.5c 0.69 + 0.ll c Overall, reptiles were the most consistent and stable vertebrates at Ua-Ub despite desert blooms in 1975 and 1983, a drought in 1977, and the severe winters of 1979 and 1984. Lizard abundance was concentrated in two dissimilar vegetation types, juniper and shad- scale (Table 9). Juniper provided lizard popula- tions with the following features: a high percent- age of bare ground and rock (75%); shallow soils; low annual plant productivity; juniper trees, both live and dead, to serve as perches and nest sites; and additional vertical relief due to sand- stone escarpments and randomly strewn boul- ders. Shadscale vegetation type east of Evacua- tion Creek did not meet any of the above criteria; however, anew site selected in 1977 did. Specif- ically it had 80% bare ground and rock, shallow soils, low annual plant productivity, and addi- tional vertical relief from sandstone escarp- ments. Different were the absence of large boul- ders and only a few scattered juniper occurring at the new shadscale site, S 4 (see Fig. 7). Similarity indices derived from Motyka et al. (1950) for reptilian species composition and abundance were used to compare within and between juniper and shadscale vegetation types. When all reptile populations in shad- scale were sampled east of Evacuation Creek, species composition and abundance similarity averaged 88 + 18% and 63 + 12%, respec- tively. A new shadscale site west of Evacua- tion Creek compared to site S 1 at 34 + 21% similarity in species composition and 13 + 5% similarity in abundance. Comparing reptiles between the new shadscale site and juniper resulted in similarity of species composition and abundance at 82 + 1% and 66 + 3%, respectively. These values were not much dif- ferent from similarities within juniper: 69 + | 7% for composition and 72 + 4% for abun- | dance. Using the flora as a sole criterion for delineating a vegetation type was disputed by the reptiles. Classifying a vegetation type based on physical and faunal as well as floral characteristics would be more appropriate for environmental assessment. Reptiles were inactive from November through March, then dependent on daily weather conditions in April through October. Peak lizard activity occurred in June and de- ( ‘ ( ( July 1986 GRANT: OIL SHALE TRACT WILDLIFE 505 TABLE 10. Seasonal distribution of reptiles, birds, and mammals according to abundance, species richness, and species diversity at Oil Shale Tracts Ua-Ub. Letters signify differences at p < 0.05. a a ee eee eee Abundance Richness Diversity Month + SE +SE +SE REPTILES 1975-1981 Number/Kilometer Number of species H’ June 5) as OLS) A 5) ae (083 1.16 + 0.07 August 4+0.4b dyes (0,3 1.09 + 0.07 Birps (Greasewood, Shadscale, Juniper Only) 1975-1981 Number/Kilometer Number of species H’ Sa la MI OC ge a NIRS CAN cae ice ee February ose ORal 5+0.4 a 0.44 + 0.08 a April 12+1.0b 18} S29.) |o) 1.46 + 0.10 b June 144+1.0b OH 250Re 1.96 + 0.06 c August 10 + 2.0 b 14+20b 1.53 + 0.07 b October 11+2.0 b 8+1.0d 0.94+ 0.09 d BirDs (Riparian Only) 1975-1981 Number/Kilometer Number of species H’ February 20+ 12.0 a 1l+10a 1.18 + 0.17 a April 28+ 3.0 a 27 + 3.0 b 2.41 + 0.08 b June 63 + 9.0 b 40+4.0c¢ 2.42 + 0.10 b August i ae, ZL) 8) 30 + 3.0 b 2:26) =) 0512" be October 9) ae 60) A 18+2.0a 1.91 +0.17 c¢ Bats 1977-1980 Individuals/Trap night Number of species H’ June 18+2.0a 5+0.4a 1.19 + 0.09 a August 11+2.0b 6) 2) (058): Lo) 0.91 + 0.10 b MAMMALS (Lagomorpha, Rodentia, Carnivora, Artiodactyla) 1975-1981 Number/Kilometer Number of species H’ February 1+04a 9) se (0, 0.23 + 0.06 April 1+0.3a 9 ae (N),2 0.26 + 0.06 June 2+ 0.4 b QOS 0.36 + 0.08 August 3+0.7 ¢ Ph as (09 0.31 + 0.07 October 3+0.9 ¢ 9) ae (0,8) 0.22 + 0.06 RODENTS 1975-1981 Individuals/100 Trap Night Number of Species H’ February 8+20a Sas 4) a Of51 0:14 a April 14+2.0b 3+ 0.3 b 0.75 + 0.10 b June 144+10b 4+0.4¢ 0.97 + 0.10 c August 15 + 2.0 b 5+0.3¢ 1.01 + 0.09 c October Igss 20) |) 4+0.4b 0.76 + 0.09 b clined by August (Table 10). About 30% of active lizards during August were juveniles. Bird populations at Ua-Ub, especially those in riparian vegetation, were at high levels of abun- dance during 1975, 1982, and 1983 (Fig. 13); yet, when abundance in all vegetation types was ana- lyzed, no significant annual differences were de- tectable. Nor were there any annual differences in species richness or species diversity; how- ever, density, richness, and diversity did show annual differences during 1975-1981 (Steele et al. 1987). Nevertheless, the fact that peak avian abundance occurred in riparian vegetation dur- ring the years mentioned was evident. It was also evident that avian abundance and richness in riparian vegetation far exceeded the values found in upland vegetation, although juniper species diversity ranked it as a vegetation type equivalent to riparian (Table 9). A masking effect of riparian bird populations led to an analysis of annual avian abundance in greasewood, shadscale, and juniper vegetation only. The results showed that bird abundance in upland vegetation during 1975 (23 birds/km) ex- ceeded all years (p < 0.05) except 1983 (21 birds/ km) and 1984 (18 birds/km). The abundance in 506 200 #— Greasewood e--— Shadscale 150 “-—- Juniper +--- Riparian 100 Number/km 90 SAG 1980 1982 1984 Fig. 13. Avian abundance in four vegetation types at Oil Shale Tracts Ua-Ub, Uintah County, Utah. 1978 1977 (9 birds/km) was lower than all other years (p = 0.05) except 1978 (10 birds/km), 1979 (12 birds/km), and 1984 (14 birds/km). The effect of the drought was reflected in bird abundance; however, change in species richness from year to year was imperceptible. Seasonal changes in bird abundance and rich- ness were also separated to eliminate the mask- ing effect of riparian population. Riparian birds reached peak abundance in June, whereas up- land bird populations reached a plateau in April and did not change appreciably through October (Table 10). Both riparian and upland populations were at peak richness in June, also when diver- sity was highest for upland birds; however, di- versity of riparian birds did not differ from spring through late summer. The large mammal community was so limited in species that the desert cottontail population dynamics (see Fig. 8) were essentially the com- munity dynamics (Table 9 and 10). The bat com- munity, on the other hand, was at highest abun- dance, richness, and diversity in June (Table 10). Annually, bat abundance was low during the 1977 drought (8 bats/trap night), then increased dramatically in 1978 (25 bats/trap night), and declined to abundance equivalent to 1977 in 1979 and 1980 (p = 0.05). This pattern mimicked that of aerial-feeding insectivores, i.e., night- hawks, swifts, and swallows (p S$ 0.01; r = 0.976). The rodent community in all vegetation types responded in concert to changes in weather and subsequent changes in floral productivity (Fig. 14). Peak density in 1976 exceeded most years, especially 1977 through 1979 (p = 0.05). By 1980 densities returned to what now appears a me- dian level. However, from 1981 through 1983 only riparian rodents, deer mice and voles, ex- GREAT BASIN NATURALIST Vol. 46, No. 3 60 #— Greasewood *e-- Shadscale “-—- Juniper bh U1 +--- Riparian O1 Individuals/ha W o) ISAS, 1980) NI82ass4 Fig. 14. Rodent density in four vegetation types at Oil Shale Tracts Ua-Ub, Uintah County, Utah. 1978 ceeded the median. Species richness also peaked in 1976 (eight species) compared to all years except 1975, 1981 and 1982 and was lowest during 1977 (three species) and 1979 (four spe- cies) (p S 0.05). Riparian vegetation that supported the highest rodent densities also supported lowest richness and diversity at Ua-Ub; juniper vegeta- tion that supported lowest densities supported the highest richness and diversity (Table 9). Al- though juniper was consistently noted for its low densities, it held the record for rodent biomass prior to the invasion and short stay by montane voles in riparian. Rodent abundance increased from February to April, then maintained from spring through the fall (Table 10). Species activ- ity was highest during the summer and lowest in winter. During the seven-year span of seasonal trap- ping, the vegetation type that supported the highest rodent abundance was greasewood (p S 0.05). The shift to high density in riparian coin- _ cided with flash floods depositing upland sandy alluvium (up to 18 in deep) over the moist silty loams. Although seasonal rodent sampling and annual density sampling agreed on annual and spatial distribution, seasonal sampling during August did not adequately predict density (Steele et al. 1984). Measurable changes in weather, soil depth and type, foliage height profiles, and floral pro- ductivity resulted in significant changes in wildlife. Spatially amphibians lacked the appro- priate conditions for abundant and diverse popu- lations. Reptiles, however, though not repre- _ sented by numerous species, did prefer juniper and certain attributes of shrub vegetation. Bird | were attracted to the most structufally diverse July 1986 and productive vegetation. Larger mammals tended to be best represented in riparian habi- tat; however, small rodent species avoided it. Reptiles, most birds, and bats were most abundant and diverse when weather was moder- ate or tending toward hot and annual grasses, forbs, and insects were most abundant and di- verse. Larger birds, i.e., raptors and waterfowl were active during early spring, whereas larger mammals and rodents were most abundant and diverse from late summer into fall. Shelter for raptors was not important, since usable nest sites exceeded raptor density. Waterfowl nesting ac- tivity preceded spring runoff, and by midsum- mer waterfowl were virtually absent. Cottontails and the other large mammals were most suscep- tible to winter conditions and also responded in kind to floral production, especially perennial shrub production that peaked in fall. Rodents followed the same pattern and were also subject to soil distribution. Desert wildlife reached different levels of abundance, richness, and diversity at different seasons, in different vegetation types, and dur- ing different years. The directions taken ap- peared to be predictable if sufficient years and enough environmental variables are measured. Weather, shelter, food, and competition, in that order, will best predict wildlife population and community dynamics. ACKNOWLEDGMENTS My gratitude is extended to R. Madsen, J. Godlove, W. and N. Hale, R. and T. Anderson, and L. Page of WRSOC; P. Kung, B. Steele, R. Bayn, N. Preece, R. Miller, M. Sylvia, E. Parker, and S. VanderWall of Bio-Resources; and personnel of VIN, Inc., Aerovironment, Inc., and NPI, Inc., during monitoring of tracts Ua-Ub. LITERATURE CITED AMERICAN ORNITHOLOGISTS UNION (AOU). 1983. Check- list of North American birds. 6th ed. Allen Press, Inc., Lawrence, Kansas. 877 pp. ARMSTRONG, D. M. 1972. Distribution of mammals in Colorado. University of Kansas Mus. Nat. Hist. Monogr. 3: 1-415. BARBOUR, R. W., AND W. H. Davis. 1969. Bats of America, University of Kentucky Press, Lexington. 286 pp. BEHLE, W. H. 1981. The birds of northeastern Utah. Utah Mus. Nat. Hist. Occ. Publ. 2. 136 pp. BEHLE, W. H., AND M. L. Perry. 1975. Utah birds: check-list, seasonal and ecological occurrence charts and guides to bird finding. Univ. of Utah Mus. Nat. Hist., Salt Lake City. 142 pp. GRANT: OIL SHALE TRACT WILDLIFE 507 BocaN, M. A. 1974. Identification of Myotis californicus and Myotis leibii in southwestern North America. Proc. Biol. Soc. Washington 87: 49-56. Cook, A. G. 1984. Birds of the desert region of Uintah County, Utah. Great Basin Nat. 44: 584-620. Durrant, S. D. 1952. Mammals of Utah. Univ. of Kansas Mus. Nat. Hist. Publ. 6:1-549. EMLEN, J. T. 1971. Population densities of birds derived from transect counts. Auk 88: 323-342. FINDLEY, J. S., ANDC. JONES. 1964. Seasonal distribution of the hoary bat. J. Mammal. 45: 461-470. HASENYAGER, R. N. 1980. Bats of Utah. Utah Div. Wildl. Res. Publ. 80-15. 109 pp. Haywarp, C. L., C. Corram, A. M. Woopsury, AND H. H. Frost. 1976. Birds of Utah. Great Basin Nat. Mem. 1: 1-229. Haywakp, C. L., D. E. BECK, AND W. W. TANNER. 1958. Zool- ogy of the Upper Colorado River Basin. I. The biotic communities. Brigham Young Univ. Sci. Bull., Biol. Ser. 1(3): 1-74. KruTSZCH, P. H., AND C. A. HEPPENSTALL. 1955. Additional distributional records of bats in Utah. J. Mammal. 36: 126-127. Merriam, C. H. 1884. The mammals of the Adirondack re- gion. Trans. Linn. Soc. 1,2: 1-316. OLSEN, P. F. 1973. Wildlife resources of the Utah Oil Shale Area. Utah Div. Wildl. Res. Publ. 74-2. 147 pp. PARKINSON, A. 1979. Morphologic variation and hybridization in Myotis yumanensis sociabilis and Myotis lucifugus carissima. J. Mammal. 60: 489-504. PETERSON, R. T. 1961. A field guide to western birds. Houghton Mifflin Co., Boston. 309 pp. RANCK, G. L. 1961. Mammals of the East Tavaputs Plateau. Unpublished thesis, University of Utah, Salt Lake City. 230 pp. RossIns, C. S., B. BRUUN, AND H. S. Zim. 1966. Birds of North America. Golden Press, New York. 340 pp. SHANNON, C. E., AND W. WEAVER. 1964. The mathematical theory of communication. University of Illinois Press, Urbana. 125 pp. SmitH, H. M. 1946. Handbook of lizards. Comstock Publ. Assoc., Ithaca, New York. 557 pp. STEBBINS, R. C. 1966. A field guide to western reptiles and amphibians. Houghton Mifflin Co., Boston. 279 pp. STEEL, R. G. D., AND J. H. Torri. 1960. Principles and proce- dures of statistics. McGraw-Hill Book Co., New York. 481 pp. STEELE, B. B., R. L. BAYN, ANDC. V. GRANT. 1984. Environmen- tal monitoring using populations of birds and small mammals: analyses of sampling effort. Biol. Cons. 30: 157-172. STEELE, B. B., AND S. B. VANDER WALL. 1985. Aquatic birds of the White River, Uintah County, Utah. Great Basin Nat. 45: 113-116. STEELE, B. B., S. B. VANDER WALL, AND C. V. GRANT. Long- term variation in the structure of bird communities in the Uinta Basin, Utah. Submitted to Ecol. Monogr. TANNER, W. W. 1957. A study of the western subspecies of the milk snake. Trans. Kansas Acad. Sci. Tenaza, R. R. 1966. Migration of hoary bats on South Farallon Island, California. J. Mammal. 47: 533-535. Twomey, A. C. 1942. The birds of the Uinta Basin, Utah. Carnegie Mus. Ann. 28: 341-490. WALTERS, R. E., AND E. SORENSEN, eds. 1983. Utah bird distri- bution: Latilong study. Utah Div. Wild]. Res. 83-10. 97 pp. COMPARISON OF VEGETATION PATTERNS RESULTING FROM BULLDOZING AND TWO-WAY CHAINING ON A UTAH PINYON-JUNIPER BIG GAME RANGE J. Skousen*”, J. N. Davis’, and Jack D. Brotherson® ABSTRACT. —Two adjacent mechanically treated pinyon-juniper (Pinus spp. and Juniperus spp.) big game winter range sites in central Utah were sampled in 1981 to estimate vegetational differences and tree mortality from the two treatments. One site was treated by selectively bulldozing in 1957 and the other was double chained in 1965. Both treatments significantly reduced tree and litter cover, whereas significant increases were found for native grasses and shrubs compared to a nearby untreated site. Juniper cover for the untreated site was 35.5% compared to only 1.4% for the bulldozed area and 4.1% for the two-way chained area. Browse species densities were increased by the mechanical treatments. The use of different mechanical treatments on separate smaller portions of critical areas of big game winter range would help provide: (1) for both long-term and short-term use of a critical wintering area, (2) greater overall productivity and carrying capacity, and (3) greater diversity by creating more edge effect between the differently treated and untreated areas. Pinyon-juniper (Pinus spp. and Juniperus spp.) ranges cover roughly 30 million ha in the western United States (West et al. 1975). Since the mid-1950s, mechanical treatment of pinyon-juniper ranges to increase forage pro- duction has been extensive (Aro 1975, Phillips 1977, Plummer et al. 1968). Methods used to reduce tree competition include: cabling, one-way chaining, two-way chaining, bulldoz- ing, windrowing, tree crushing, and burning (Arnold et al. 1964, Aro 1975, Plummer et al. 1960, 1968, 1970, Stoddart et al. 1975, Vallen- tine 1980). This paper evaluates differences in vegetational patterns following bulldozing and two-way chaining and seeding on adjacent sites in central Utah. STUDY AREA AND METHODS The study area is a pinyon-juniper big game winter range 2 km east of Holden, Millard County, Utah. Elevation is approximately 1,600 m. The area is classified as an upland stony loam range site. Soils are slightly cal- careous with a pH of 6.9. Average yearly pre- cipitation is about 37.5 cm, with most of it coming during the winter months. Slope is relatively constant and averages 7%. The as- pect is southwesterly. Before treatment the area supported an open stand of juniper (Ju- niperus osteosperma ) with an intermixture of cliffrose (Cowania stansburiana), big sage- brush (Artemisia tridentata), broom snake- weed (Xanthocephalum sarothrae), and some antelope bitterbrush (Purshia tridentata). Cheatgrass (Bromus tectorum) is the most prominent understory spécies on the control site. Other grasses and forbs are infrequent and produce little forage (Christensen et al. 1964). The bulldozed site is owned and managed by the Utah Division of Wildlife Resources, and the two-way chained site is federally owned and managed by the USDI Bureau of Land Management (BLM). Bulldozing was used to eliminate trees while minimizing dis- turbance to cliffrose and big sagebrush on Utah Division of Wildlife Resources land (Christensen et al. 1964). These two shrub species are considered important winter browse for big game animals. The nine species seeded on the bulldozed site, listed in Table 1, were either broadcast seeded or hand seeded into the depressions left by uprooted juniper trees (Plummer et al. 1960). Only crested wheatgrass (Agropyron cristatum) was seeded on the BLM site; 9 kg/ha of seed was applied during the chaining operation. Grazing has varied on the bull- dozed site since treatment. It was rested from Utah State Division of Wildlife Resources, Shrub Sciences Laboratory, Provo, Utah 84601. Current address: Division of Plant and Soil Sciences, West Virginia University, Morgantown, West Virginia 26506. 3Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 508 } _- i 4 ‘ i \ | July 1986 SKOUSEN ETAL.: VEGETATION PATTERNS 509 TABLE 1. Percent cover by species on bulldozed and two-way chained sites and adjacent untreated pinyon-juniper ranges. Amount of broadcasted seed is for the bulldozed site only. Species Untreated ANNUALS Bromus tectorium 133" Total all annual species 1.33* GRASSES Agropyron cristatum 0.00* Agropyron spicatum 0.00° Agropyron tricophorum 0.00* Poa secunda 0.41° Total all grass species 0.46* SHRUBS Artemisia tridentata 2.43* Cowania stansburiana 3.00° Xanthocephalum sarothrae 0.42° Total all shrub species 5.85" | TREES | Juniperus osteosperma 35.47° | Quercus gambelii 6.64* | Total all tree species 42.11° | grazing from 1961 to 1962 and 1964 to 1965, | but when grazed the stocking rate was 3.0 \ha/per animal unit (AU) from 10 May to 15 \June. Since treatment the two-way chained site has had a stocking rate of 4.5 ha/per ani- mal unit (AU) from 1 May to 15 June. Both areas have been grazed by cattle throughout _ the study periods. Treated areas were sampled by a single transect running across both treatments. This transect was 800 m long, with 400 m within each treatment. A second transect of 400 m »was used to sample a nearby untreated area that was used as a control. Meter-square quadrats were placed at 10 m intervals along the transect. Total plant cover and cover by species were estimated using a variation of Daubenmire’s (1959) cover class estimation technique. The variation consisted of adding a ‘smaller cover class, 0% to 1%, so that the cover of small plants would not be overesti- mated. Percent cover of bare ground, litter, and rock were also estimated at each quadrat. Density of trees and shrubs was evaluated ‘by the use of a circular plot with an area of 50 ‘m°. This enlarged plot was centered on every third quadrat along the transect. In each en- larged plot, every shrub and tree was identi- Treatment ; l Seeding rate Bulldozed Two-way (kg/ha) 3.85° 2.97? 4.05" 3.31? 0.19° 4.96? 1.597 8.74? 1.68* 2.54? 0.00* 0.55 2.01? 0.798 13.68° See 10.92? 5.95° 0.137 4,94? 3.68" 0.01? 2.55? 9.65? Taz? 19.40° 1.42> 4.10° 2.64" 0.007 4.06? 4.10? | *Values in the same row with different letters are significantly different at P < .05. / 1 Other seeded species were Agropyron intermedium (.48), Medicago sativa (0.08), Atriplex canescens (.17), Chrysothamnus nauseosus (.14), and Purshia ) tridentata (.34). The seeding rate for two-way chained site was Agropyron cristatum at 9 kg/ha. | ?Some of the seed was hand-seeded into the depressions. fied and assigned to a height and stem diame- ter class. Diameter of all trees was measured at 10 cm aboveground, and the largest stem of each multistemmed shrub was measured at approximately the same height. At least five juniper trees were cut near each transect and aged by growth rings. Regression analysis was used to correlate stem diameter with age. Evaluated parameters include: changes in species cover and composition, changes in tree and shrub density, and tree mortality resulting from treatment. Swept pellet group transects were used to indicate big game ac- tivity within the areas of treatment. Composite soil samples were taken to a depth of 25 cm at every tenth quadrat along the transect. Soil texture, pH, and soluble salts were determined. Soils were also ana- lyzed for nitrogen, phosphorus, potassium, calcium, magnesium, sodium, zinc, iron, manganese, and copper content. RESULTS AND DISCUSSION Total plant and litter cover were signifi- cantly reduced by both treatments when com- pared to the control (Table 2). However, total plant, bare ground, litter, and rock cover 510 GREAT BASIN NATURALIST Vol. 46, No. 3 TABLE 2. Percent cover of surface characteristics on bulldozed and two-way chained sites and adjacent untreated pinyon-juniper ranges. Characteristic Untreated Total plant cover 48.87°° Bare ground 25.08°* Litter Qh225 Rock 4.47° Lifeform cover: Annual species 1.33* Forb species ol Seeded grass species .00° Native grass species .46° Seeded shrub species** 5.43" Native shrub species 42? Tree species 42.11* Treatment Bulldozed Two-way 37.90° 35.48° 27.07? 33.12? 15.33° 16.59° Rave BIG2 4.05° B.Bie 1.04? 36 2.73? 5.78? 10.95° 9..53° 12.43? 9.64? 2.64> 9.76° 4.06? 4.10° *Values in the same row with different letters are significantly different at P < .05. **Included in the seed mixture were shrub species that also occur naturally on the site. These are shown as seeded in the table so that comparisons could be made between treatments. were not significantly different between the two mechanical treatments. When these cover categories were compared, no signifi- cant differences existed between the two treatments, but visually and structurally they did appear quite different from each other. The treated sites supported significantly greater native grass and native shrub cover than the nearby untreated control site. Com- parison between treatments showed that sig- nificant cover differences existed for the total of all grass species (Table 1). Crested wheat- grass had greater cover on the two-way chain- ing, whereas bluebunch wheatgrass (Agropy- ron spicatum) and sandberg bluegrass (Poa secunda) both were native and not seeded and exhibited greater cover on the bulldozed site. Pubescent wheatgrass (Agropyron tri- cophorum) showed good establishment and persistence on the bulldozed area. Crested wheatgrass probably had greater cover because of its higher seeding rate on the two-way site, and the two-way chaining pre- pared an improved seedbed and covered the seed for better germination and establish- ment. Crested wheatgrass was also seeded on the bulldozed site, but it did not respond as well as the other two wheatgrasses after the bulldozing treatment. Bluebunch wheatgrass was probably suppressed by crested wheat- grass competition on the two-way chained site. The bulldozed site exhibited greater big sagebrush and cliffrose cover than the chained site, but the cover values were not signifi- cantly different between sites. Juniper density was significantly lower on the bulldozed site (Table 3). There were 42% more trees on the two-way chained area than the bulldozed site. This would indicate that | bulldozing to eliminate juniper in this area was probably more effective than chaining be- cause the bulldozed treatment was done eight years previous to the two-way chaining and | still had fewer juniper when sampled in 1981. In this study the bulldozing treatment killed 81% of the juniper, whereas double chaining killed only 54% when compared to the nearby control area. Aro (1975) reported a 95% to 100% kill when the trees were windrowed with a bulldozer and burned, whereas double | chaining averaged 60% kill. Arnold et al. (1964) stated that double chaining results in a 50% to 80% kill of juniper trees. Many researchers have noted that a major factor determining actual tree kill from me- chanical treatments is the age structure of the juniper stand before treatment (Aro 1975, Skousen 1982, Stevens et al. 1975). Trees on the control area indicated that the stand was relatively young at the time of treatment. Of the trees sampled there, 40% were small, | with stem diameters of 5 cm or less when the treatment took place. The presence of these small, flexible trees may explain why the chaining treatment was less effective than bulldozing. No significant differences were found between the sites for cliffrose and big sagebrush densities. ow Cliffrose cover was |} higher on the bulldozed site. This anomaly |, probably arose because surviving plants of oe i July 1986 SKOUSEN ET AL.: VEGETATION PATTERNS dll TABLE 3. Tree and shrub densities per hectare on bulldozed, two-way chained, and untreated pinyon-juniper ranges. Treatment Species Untreated Bulldozed Two-way TREES Juniperus osteosperma rial 428° 734° Quercus gambelii 454" 719? 305* Total trees 1225° 1147* 1039* SHRUBS Artemisia tridentata 1351° 2738° 1943°* Chrysothamnus nauseosus 21° 244° 30° Cowania stansburiana 429% 597* 963* Xanthocephalum sarothrae 389° 934* 5936" Total shrubs 2190° 4513* 8872° *Values in the same row with different letters are significantly different at P < .05. cliffrose were much larger on the bulldozed site. Two-way chaining appears to have uprooted or broken off all large cliffrose plants (Christensen et al. 1964). Cliffrose plants were intentionally avoided on the bulldozed project. Broom snake- weed was six times more dense on the chained site, suggesting that increased soil disturbance may have allowed this invader species to spread (Arnold et al. 1964). For example, broom snake- weed made up 7% of the shrub cover for the untreated area, 14% of the shrub cover for the bulldozed area, and a high 50% of the shrub cover for the two-way chained area. Only one soil factor was significantly different between untreated, bulldozed, and two-way chained sites (Table 4). Phosphorus concentra- ' tions were significantly lower on the bulldozed site than on the untreated or two-way chained / areas. Utah Wildlife Resource conservation officers ‘reported that swept pellet group transects | showed big game activity to be two to three | times heavier on the bulldozed site than on the ) two-way chained area. Some years, e.g. 1977, | big game use was 13 times heavier on the bull- \ dozed site than on the two-way chained areas (Brent Olsen, data on file). It has been suggested that the preferential use _ of the bulldozed site by deer was related to the _ greater height of the cliffrose. On the two-way § chained area, 75% of the cliffrose individuals were under | m tall, and all cliffrose plants were under 1.5 m tall. On the bulldozed site, 50% of _the cliffrose plants were under 1 m tall, 30% t were between 1 and 2 m, and 20% were over 2 m. The deer apparently preferred the bulldozed site where the larger cliffrose plants provided 'security cover as well as a variety of forage. Even though the study sites were treated eight years apart, some generalizations are worth noting. Although bulldozing costs are 1.5 to 2 times greater (depending on tree density) than two-way chaining (Bill Davis, personal communication), bulldozing was a practical and effective method for juniper removal on this site. Widemann and Cross (1981) found that bulldoz- ing light to moderate stands of juniper in Texas with a small, low-powered crawler tractor was an economical alternative. Their cost varied from $6 to $50 per ha. The trees on this area did not constitute a closed stand, but the bulldozing treatment left Gambel oak (Quercus gambelii), a sprouting species, undisturbed. Bulldozing also allowed minimal disturbance to desired under- story species. Cliffrose, an important cover and browse species for big game in this area, was left intact and big sagebrush populations were reju- venated following treatment and seeding. Be- cause big game showed a two- to threefold pref- erence for bulldozed versus the two-way chained site, restoration projects on pinyon-ju- niper big game ranges in this area should con- sider the feasibility of using this method. How- ever, because cliffrose density was increased by double chaining, the two-way chained area should become increasingly more valuable to big game in the future as the cliffrose becomes more mature. Within areas of critical winter range, it would be advisable to apply different mechanical treat- ments to different sections of the range. Bull- dozed areas could be used by wildlife immedi- ately because the understory is left minimally disturbed (assuming an understory of desirable remnant plants are present for subsequent re- lease and reseeding themselves), whereas 512 GREAT BASIN NATURALIST Vol. 46, No. 3 TABLE 4. Soil data on bulldozed, two-way chained, and adjacent untreated pinyon-juniper ranges. Characteristic Untreated Texture Clay loam Depth, cm 1G. pH 6.5° Soluble salts, ppm 608.0" Phosphorus, ppm 36.2* Potassium, ppm 300.7* Calcium, ppm 6756.0* Magnesium, ppm 381.2* Sodium, ppm 43.9* Zinc, ppm 1.6 Iron, ppm 76.0° Manganese, ppm 61.5° Copper, ppm Pils % Nitrogen 0.2° Treatment Bulldozed Two-way Loam Loam 15.0? 14.0? 6.8* 6.9? 590. 0* 560. 0* 80? fale 234.0" 228.0" 9500. 0* 10075.0* 300. 0* 275.0° 86.5° 83.0* 1.0° 1.67 57.3" 37.5" 49.0* 33.6° 1.0° LF 0.2 0.2? *Means within rows with the same letter are not significantly different at P < .05. chained areas would become more beneficial as the plant species mature. A regional ap- proach should be developed that would allow areas to be treated by different mechanical techniques (varying degrees of disturbance) while leaving some areas undisturbed for a better balanced use of the resource through time. These combined treatment effects would provide for: (1) both short-term and long-term use of the treated areas by various livestock and wildlife species because of the varying stages of community development that the mechanical treatments induce, (2) greater overall productivity and carrying ca- pacity, and (3) greater edge effect between differentially treated and undisturbed areas. ACKNOWLEDGMENTS This research was made possible with fed- eral funds for wildlife restoration provided through Pittman-Robertson Project W-82-R, Job 1, in cooperation with the Intermountain Research Station, USDA Forest Service, and the Utah Division of Wildlife Resources. LITERATURE CITED ARNOLD, J. F., D. A. JAMESON, AND E. H. REtp. 1964. The pinyon-juniper type of Arizona: effects of grazing, fire, and tree control. USDA Prod. Res. Rep. 84. Rocky Mtn. For. and Range Expt. Sta., Fort Collins, Colorado. 28 pp. Aro, R. S. 1975. Pinyon-juniper woodland manipulation with mechanical methods. Pages 67-75 in The pinyon-juniper ecosystem: a symposium. Utah State University, Logan. CHRISTENSEN, D. R., S. B. MONSEN, AND A. P. PLUMMER. 1964. Response of seeded and native plants six and seven years after eradication of Utah juniper by cabling and hula-dozing followed by pipe harrowing as an after treatment on portions of the major treatments. Un- published report by Utah State Fish and Game De- partment, Salt Lake City. 28 pp. DAUBENMIRE, R. 1959. A canopy-coverage method of vegeta- tional analysis. Northwest Sci. 33: 43-46. PuiLuirs, T. A. 1977. An analysis of some Forest Service pinyon-juniper chaining projects in Region 4— | 1954-1975. Range Improve. Notes. USDA Forest Service Intermt. Region, Ogden, Utah. 20 pp. PLUMMER, A. P., H. D. STAPLEY, AND D. R. CHRISTENSEN. 1960. Job completion report for game forage revegetation | project W-82-R-5. Utah State Dept. of Fish and Game _ Info. Bull. 1957-1960. 51 pp. PLUMMER, A. P., D. R. CHRISTENSEN, AND S. B. MONSEN. 1968. Restoring big game range in Utah. Utah State Div. of Fish and Game, Publ. No. 68-3. 183 pp. | PLUMMER, A. P., D. R. CHRISTENSEN, R. STEVENS, K. R. Jor. | GENSEN. 1970. Highlights, results, and accomplish- ments of game range restoration studies, 1970. Utah State Div. of Fish and Game, Publ. No. 70-3. 94 pp. SKOUSEN, J. G. 1982. Evaluation of pinyon-juniper chainings for big game in central Utah. Unpublished thesis, Brigham Young University, Provo, Utah. 74 pp. | STEVENS, R., B. C. GIUNTA, AND A. P. PLUMMER. 1975. Some | aspects in the biological control of juniper and pinyon. Pages 77-80 in The pinyon-juniper ecosystem: asym- | posium. Utah State University, Logan. Stoppakrt, L. A., A. D. SmirH, AND T. W. Box. 1975. Range | management. 3d ed. McGraw-Hill Book Co., New | York. 532 pp. VALLENTINE, J. F. 1980. Range development and improve- | ments. 2d ed. Brigham Young University Press, | Provo, Utah. 545 pp. i West, N. E., K. H. REA, AND R. J. Tauscu. 1975. Basic syneco- | logical relationships in juniper-pinyon woodlands. | Pages 41-54 in The pinyon-juniper ecosystem: a sym- | posium. Utah State University, Logan. | WIDEMANN, H. T., AND B. T. Cross. 1981. Low-energy grub- | bing for control of junipers. J. Range Manage. 34: | 235-237. | | | H | VERTEBRATE FAUNA OF THE IDAHO NATIONAL ENVIRONMENTAL RESEARCH PARK Timothy D. Reynolds’, John W. Connelly'?, Douglas K. Halford'?, and W. John Arthur’ ABSTRACT. —The relative abundance, habitat use, and seasonal occurrence are reported for the 6 fish, | amphibian, 9 reptile, 164 bird, and 39 mammal species recorded on the Idaho National Environmental Research Park in southeast- ern Idaho. The Idaho National Engineering Labora- tory (INEL) is an energy research and devel- opment site administered by the U.S. De- partment of Energy (DOE). The Energy Reorganization Act of 1974 ordered the U.S. Energy Research and Development Adminis- tration (precursor to DOE) to engage in envi- ronmental research related to the develop- ment of energy sources to advance the goals of restoring, protecting, and enhancing environ- mental quality. The INEL was designated the nation’s second National Environmental Re- search Park (NERP) in 1975 to satisfy this directive. The NERP provides a controlled, protected, outdoor laboratory for environ- mentally related research to help achieve na- tional environmental goals as stated in the National Environmental Policy Act (NEPA) of 1969. The NERP charter presents broad ob- jectives which, in part, require that each NERP serve as a bench mark for quantita- tively assessing and predicting the environ- mental impact of man’s activities. The compi- lation of baseline data, including species lists and identification of ecological communities, is fundamental to the NERP objectives. Much of the research on individual species of wildlife found on the Idaho NERP has been published in the scientific literature or pre- sented in theses, dissertations, or reports (vide Markham 1973, 1978, 1983). This paper represents a consolidation of those findings which, combined with unpublished informa- tion, provide baseline data on the abundance, distribution, habitat preference and seasonal occurrence of the vertebrate species recorded on the Idaho NERP. STuDY AREA The INEL was established by the U.S. Atomic Energy Commission as the National Reactor Testing Station in the late 1940s. Since 1949 the land composing the Idaho NERP has been closed to public access, thereby providing wildlife habitat that has re- mained relatively undisturbed for 35 years. The NERP occupies about 2,305 km? of sagebrush-dominated rangeland on the upper Snake River Plain in portions of Bonneville, Bingham, Butte, Clark, and Jefferson coun- ties. The INEL is approximately 48 km west of Idaho Falls, Bonneville County, Idaho. With the exception of two large buttes of volcanic origin near the southeastern boundary, and several small cinder cones, craters, and ex- posed lava ridges scattered over the site, the topography of most of the NERP is flat to gently rolling and typical of the Columbian Plateau Province (Atwood 1970). The mean elevation is 1,490 m ASL. Three predomi- nately north-south mountain ranges, rising to 3,370 m ASL, border the NERP to the north and west (Fig. 1). The intervening valleys and the Snake River Plain itself provide migration corridors to, or across, the Site. Soils are derived mostly from silicic volcanics and Pale- ozoic rocks from the surrounding mountains. These are primarily aeolian sandy loams and loess underlain by undifferentiated basalt ‘Radiological and Environmental Sciences Laboratory, U.S. Department of Energy, 785 DOE Place, Idaho Falls, Idaho 83402. Present address: Idaho Department of Fish and Game, 5205 South 5th, Pocatello, Idaho 83201. 3Present address: Waste Management Programs, EG&G Idaho, Inc., P.O. Box 1625, Idaho Falls, Idaho 83415. *Present address: Uranium Mill Tailings Remedial Action Project, U.S. Department of Energy, P.O. Box 5400, Albuquerque, New Mexico 87115. 513 514 GREAT BASIN NATURALIST Vol. 46, No. 3 Circular Butte Lost River Sinks Cinder Butte Lost River Range Diversion dam Spreading Area 0 5 10 Kilometers ) 26 | | Miles Big Southern Butte 5 0818 Fig. 1. Landmarks, roadways, and watercourses on the Idaho National Environmental Research Park and environs in southeastern Idaho. The area of the park open to livestock grazing is shaded. July 1986 atop a rhyolite foundation. The climate is characterized by hot summers and cold win- ters. Maximum and minimum temperatures recorded from 1950 to 1983 are 39 and —44 C, respectively. Annual precipitation over the past 34 years averaged 21.6 cm (Range: 11.4-36.6 cm), coming mostly from spring rain or snow storms in April, May, and June, with a smaller peak resulting from snowfall during December and January. Surface water is limited on the Idaho NERP. The Big Lost River enters the south- west corner of the INEL, flows northward about 50 km, and ultimately percolates into the Snake River aquifer at the sinks near Howe, Idaho (Fig. 1). When filled, the sinks provide about 260 surface ha of aquatic habi- tat. Because of annual variations in the snow- pack, offsite irrigation demands upstream, and the porosity of the stream bed, the Big Lost River does not flow across the NERP throughout every summer. In some years wa- ter never reaches the sinks. Water is also di- verted from the Big Lost River into four spreading areas in the southwest corner of the INEL (Fig. 1) to prevent ice jams and flooding of some INEL facilities during the winter and spring runoff. Thus a series of ephemeral shal- low lakes of up te 919 ha in total size is gener- ally present during the spring peak of bird migration. Other natural water sources on the Idaho NERP are the Little Lost River and Birch Creek. Both of these streams are cur- rently diverted offsite for agricultural irriga- tion and only rarely flow onto the INEL. Sev- eral 0.2—5.0 ha man-made ponds occur near facilities on site. In aggregate these provide approximately 15 ha of additional aquatic habitat. Although McBride et al. (1978) described 20 distinct vegetative cover types on the INEL, nearly 87% of the NERP is shrub- steppe habitat. About 7% is grassland, and about 3% is juniper woodland. Aquatic and riparian habitat and irrigated lawns at facility complexes account for the remaining 3%. The _ dominant and most conspicuous vegetation is _ big sagebrush (Artemisia tridentata). Other locally important shrubs are rabbitbrush | {Chrysothamnus nauseosus and C.. vicidi- florus), winterfat (Ceratoides lanata), shad- scale saltbush (Atriplex confertifolia), Nuttall saltbush (A. nutallii), and gray horsebrush REYNOLDS ET AL.: IDAHO VERTEBRATES 515 (Tetradymia canescens) (Anderson and Holte 1981, Harniss and West 1973). The more abun- dant grasses include bottlebrush squirreltail (Elymus elymoides: formerly Sitanion hystrix), needle and thread grass (Stipa comata), Indian ricegrass (Oryzopsis hymenoides), Great Basin wild rye (Leymus cinereus: formerly Elymus cinereus ), thickspike wheatgrass (Elymus lance- olatus : formerly Agropyron dasystachyum), and bluebunch wheatgrass (Elytrigia spicata: for- merly A. spicatum). Native tree species are lim- ited to junipers (Juniperus osteosperma and J. scopulorum) near the southeast and northwest portions of the NERP, and plains cottonwood (Populus deltoides), narrow-leaved cottonwood (P. angustifolia), and several species of willow (Salix spp.) along the Big Lost River and Birch Creek. Russian olive (Elaeagnus angustifolia) and several other introduced species of decidu- ous and evergreen trees have been planted near most of the INEL facilities. Lawns composed of Poa spp. are maintained at most of the facility complexes. Approximately 4,000 ha of the INEL were seeded with crested wheatgrass (Agropy- ron cristatum and A. desertorum) in the late 1950s and early 1960s. Reinvasion by native shrub-steppe vegetation has been slow, and these plantings persist mostly as monocultures (Marlette 1982). Jeppson and Holte (1978) listed a total of 389 vascular plant species on the INEL. None are classified as threatened or endangered at either the federal or state level (Cholewa and Henderson 1984). The periphery of the NERP, representing nearly 65% of the total area, is open to seasonal cattle and sheep grazing regulated by the U.S. Department of the Interior Bureau of Land Management (Fig. 1). The geology, topography, vegetation, cli- mate, and recent history of Idaho NERP previ- ously have been described in detail (Anderson and Holte 1981, Atwood 1970, Harniss and West 1973, Nace et al. 1972, 1975, McBride et al. 1978). Public access to the Idaho NERP is restricted. Except for scientific purposes and periodic predator control, the collection of flora and fauna is prohibited. METHODS Past and present ecological studies con- ducted on the INEL and correspondence and interviews with INEL researchers and per- 516 GREAT BASIN NATURALIST TABLE 1. Fishes recorded on the Idaho National Environmental Research Park. Taxa SALMONIFORMES Salmonidae Kokanee Salmon, Oncorhynchus nerka Rainbow Trout, Salmo gairdneri Brook Trout, Salvelinus fontinalis Mountain Whitefish, Prosopium williamsoni CYPRINIFORMES Cyprinidae Speckled Dace, Rhinichthys osculus PERCIFORMES Cottidae Shorthead Sculpin, Cottus confusus ‘See text for definitions of abundance terms. sonnel provided the data presented in this paper. Species inventories and population as- sessments on the Idaho NERP have been published for small mammals (Allred 1973), herpetofauna (Sehman and Linder 1978), and raptors (Craig 1979). Additional data on these and other taxa were gleaned from numerous sources. Those that provided limited but use- ful information germane to this paper and not cited elsewhere in the text included theses and dissertations (Craig 1977, Fuller 1981, Gleason 1978, Hoskinson 1977, Johnson 1977, Peterson 1982, and Reynolds 1978) and technical publications (Bailey 1974, Best and Peterson 1982, Craig 1978, Craig and Renn 1977, Halford and Millard 1978, Powers and Craig 1976, and Reynolds and Trost 1980). Moreover, current and previous NERP re- searchers were encouraged to record observa- tions of vertebrates encountered incidental to their research activities and, from their expe- rience, to indicate the relative abundance and distribution of these species on the Idaho NERP. Informal interviews with INEL Site services and security personnel provided ad- ditional information. The relative abundance, distribution, and habitat use for all native and introduced verte- brate species recorded on the Idaho NERP are presented. Abundance ratings were used to indicate relative numbers of a species dur- ing the season(s) it was recorded on the Idaho NERP. Sehman and Linder (1978) and Over- ton (1977) provided abundance and residence status for herpetofauna and fishes, respec- tively. Abundance ratings for terrestrial homeo- therms are often biased in favor of conspicu- Vol. 46, No. 3 Distribution Abundance’ Big Lost River Uncommon Big Lost River Common Big Lost River Uncommon Big Lost River Common Big Lost River Uncommon Big Lost River Common ous and diurnal species, especially when ob- servations incidental to other projects are the source of data. To reduce this bias, former and current INEL ecological researchers subjec- tively rated the abundance of each bird and mammal species, based on personal experi- ence. Ratings were from | to 10, 1 represent- ing a very rare sighting and 10 indicating an abundant species. A mean value for each spe- cies was calculated. Because the ratings for some species ranged widely (i.e., from 1 to 9), a category termed occasional or local was in- cluded (cf Stephens and Reynolds 1983). All abundance classes assumed that a qualified biologist exerted a reasonable effort to search or sample the proper habitat at the appropri- ate time of year. Abundance ratings were: 1. Abundant—Very numerous and certain to be seen or sampled: a mean rating of 8.1—10.0. 2. Common—Likely but not certain to be observed or sampled: a mean rating of 5. 1—8.0. 3. Uncommon—Found in limited numbers, not likely to be sampled or observed: mean rating of 2.1—5.0. 4. Occasional or local—A species that is not always present or is restricted in distribution: a difference between the high and low response of 7 or more. 5. Rare—A species that has a range including all or part of the Idaho NERP but has been documented S seven times on site: mean rating of 0. 1—2.0. 6. Vagrant or accidental—A species that is not expected to occur on the Idaho NERP but has been recorded there (e.g., Black-legged Kittiwake [Rissa _ tri- dactyla |). Habitats listed for each species closely fol- low those presented by Trost et al. (1977) and Stephens and Reynolds (1983). The designa- tion “sitewide” indicates that a particular spe- cies was ubiquitous on the Idaho NERP. Un- less otherwise noted, taxonomic nomen- July 1986 REYNOLDS ET AL.: IDAHO VERTEBRATES 517 TABLE 2. Herptiles recorded on the Idaho National Environmental Research Park. Distribution Taxa and habitat Abundance’ ANURA Pelobatidae Great Basin Spadefoot Toad, Spea intermontana’ Big Lost River and sinks Common SQUAMATA Iguanidae Leopard Lizard, Gambelia wislizenii* NE NERP; sandy areas Local Short-horned Lizard, Phrynosoma douglassi Sitewide; shrub-steppe Abundant Sagebrush Lizard, Sceloporus graciosus Sitewide; shrub-steppe Abundant Scincidae Western Skink, Eumeces skiltonianus South NERP Rare Boidae Rubber Boa, Charina bottae Unknown Accidental Colubridae Desert Striped Whipsnake, Masticophis taeniatus NE NERP; shrub-steppe Uncommon Gopher Snake, Pituophis melanoleucus Sitewide; shrub-steppe Common Western Garter Snake, Thamnophis elegans Sitewide; all habitats Uncommon Viperidae Western Rattlesnake, Crotalus viridis Sitewide; shrub-steppe Common 1See text for definitions of abundance terms. Collins et al. (1978) list this as Scaphiophus intermontanus. 3Collins et al. (1978) place this in the genus Crotaphytus. clature for fish, amphibians and reptiles, birds, and mammals follows Simpson and Wallace (1978), Nussbaum et al. (1983), American Ornithological Union (1983), and Jones et al. (1982), respectively. RESULTS AND DISCUSSION A total of 219 vertebrate species were recorded on the Idaho NERP. Only six spe- cies of fish (Table 1) were identified. Four of these were game fish (all salmonids), and two were nongame species. Species inventories of fish were incidental to a study on the density and distribution of salmonid fishes in the Big Lost River (Overton 1977) and a game fish salvage and transplant operation conducted by the Idaho Department of Fish and Game. Because both of those activities primarily con- cerned game fish, the records of nongame species actually present in the Big Lost River on the NERP may be incomplete. Contrari- wise, the low species richness may indeed be accurate, caused by annual and seasonal fluc- tuations in the flow rate of the river. Overton (1977) attributed decreases in salmonid popu- lations on the NERP directly to: (1) abnor- mally high discharge in the spring and (2) low or nonexistent flows during both the winter and summer. Diversion practices to control flooding typically reduce the flow in the win- ter months, whereas low summer flows result from upstream irrigation demands and are generally accompanied by elevated water temperatures. Additional efforts may be needed to completely inventory fish popula- tions on the Idaho NERP. One amphibian and nine reptiles were recorded on the Idaho NERP (Table 2). The spadefoot toad (Spea intermontana) was re- ported from the Big Lost River as well as the Lost River Sinks and the diversion spreading area (Fig. 1). There is only one confirmed record for the rubber boa (Charina bottae) on the Idaho NERP. Concentrations of leopard lizards (Gambolia wislizenii) and desert stiped whip- snakes (Masticophis taniatus) were only found in the northeast portion of the NERP, near Circular Butte and Cinder Butte, respec- tively (Linder and Sehman 1978) (Fig. 1). The former species is evidently restricted to areas of sandy soils. The latter is more widespread but limited in numbers away from Cinder Butte (Sehman and Linder 1978). The west- ern skink (Eumeces skiltonianus) is confined to isolated locations on the southern half of the NERP, possibly because of increased micro- habitat moisture requirements (Sehman and Linder 1978). The remainder of the herptiles species listed are either sitewide in distribu- tion or found in the shrub-steppe habitat that 518 GREAT BASIN NATURALIST Vol. 46, No. 3 TABLE 3. Avifauna recorded on the Idaho National Environmental Research Park. Abundance’, season, Taxa and breeding status” Habitat® GAVIIFORMES Gaviidae Common Loon, Gavia immer M5 w PODICIPEDIFORMES Podicipedidae Pied-billed Grebe, Podilymbus podiceps S5, M5 Ww Horned grebe, Podiceps auritus M5 w Eared grebe, P. nigricollis B5, M3, W3 Ww Western Grebe, Aechmophorus occidentalis * S5, M5 Ww PELECANIFORMES Pelecanidae American White Pelican, Pelecanus erythrorhynchos M5 Ww CICONIIFORMES Ardeidae American Bittern, Botaurus lentiginosus S5, M5 Ww Great Blue Heron, Ardea herodias S5, M5 w Great Egret, Casmerodius albus S5, M5 w Green-backed Heron, Butorides striatus S6, M6 w Threskiornithidae White-faced Ibis, Plegadis chihi” S5, M5 Ww ANSERIFORMES Anatidae Tundra Swan, Cygnus columbianus M5 w Snow Goose, Chen caerulescens M5 w Canada Goose, Branta canadensis S3, M3 w Wood Duck, Aix sponsa S6, M5 5 Ww Green-winged Teal, Anas crecca S5, M5 w Mallard, A. platyrhynchos B2, M2, W3 w Northern Pintail, A. acuta S3, M3 w Blue-winged Teal, A. discors B2, M3 w Cinnamon Teal, A. cyanoptera $3, M3 Ww Northern Shoveler, A. clypeata B3, M3 w Gadwall, A. strepera S3, M3 w American Wigeon, A. americana $3, M3 w Canvasback, Aythya valisineria B5, M5 Ww Redhead, A. americana S5, M5, W5 Ww Ring-necked Duck, A. collaris S5, M5 w Lesser Scaup, A. affinis S5, M3, W3 Ww Common Goldeneye, Bucephala clangula S5, M3, W3 w Barrow’s Goldeneye, B. islandica S6, M5 w Bufflehead, B. albeola S5, M3 w Common Merganser, Mergus merganser $3, M5 w Ruddy Duck, Oxyura jamaicensis B5, M3 Ww FALCONIFORMES Cathartidae Turkey Vulture, Cathartes aura S3, M3, W6 sw Accipitridae Osprey, Pandion haliaetus” M5 w Bald Eagle, Haliaeetus leucocephalus® M5, W3 sw Northern Harrier, Circus cyaneus R2 Sw Sharp-shinned Hawk, Accipiter striatus * S5, M5, W5 sw Cooper's Hawk, A. cooperii S3, M5, W5 sw Northern Goshawk, A. gentilis S5, M5, W5 sw Swainson’s Hawk, Buteo swainsoni B3, M3, W5 sw Red-tailed Hawk, B. jamaicensis B3, M3, W5 sw Ferruginous Hawk, B. regalis”’ B3, M3, W5 sw Rough-legged Hawk, B. lagopus S6, M2, W2 sw Golden Eagle, Aquila chrysaetos B3, M4, W2 sw Falconidae American Kestrel, Falco sparverius B2, M2, W3 ; Sw Merlin, F. columbarius® R5 sw July 1986 REYNOLDS ET AL.: IDAHO VERTEBRATES 519 Table 3 continued. Abundance’, season, Tra and breeding status” Habitat® Peregrine Falcon, F. peregrinus® S5, M5, W5 sw Gyrfalcon, F. rusticolus‘ M6 eer Prairie Falcon, F. mexicanus” R3 ary GALLIFORMES Phasianidae Gray Partridge, Perdix perdix R3 g, ss, f Chukar, Alectoris chukar R3 g, SS Ring-necked Pheasant, Phasianus colchicus R3 g, Ss Blue Grouse, Dendragapus obscurus S6 f Sage Grouse, Centrocercus urophasianus R2 SSaeaah GRUIFORMES Rallidae Sora, Porzana carolina B5, M5 w,f American Coot, Fulica americana R3 Ww CHARADRIIFORMES Charadriidae Killdeer, Charadrius vociferus B2, M2 sw Recurvirostridae American Avocet, Recurvirostra americana $2, M3 w Scolopacidae Greater Yellowlegs, Tringa melanoleuca M5 w Lesser Yellowlegs, T. flavipes $5, M5 Ww Solitary Sandpiper, T. solitaria S5, M3 Ww Willet, Catoptrophorus semipalmatus $3, M3 w, Ss Spotted Sandpiper, Actitis macularia $3, M3 Ww Long-billed Curlew, Numenius americanus *° S3, M3 W, SS Marbled Godwit, Limosa fedoa S3, M5 Ww Least Sandpiper, Calidris minutilla S5, M5 w Long-billed Dowitcher, Limnodromus scolopaceus M5 Ww Common Snipe, Gallinago gallinago S5, M5 w Wilson’s Phalarope, Phalaropus tricolor S3, M3 Ww Red-necked Phalarope, P. lobatus M5 w Laridae Franklin’s Gull, Larus pipixcan S3, M3 w, Ss Bonaparte’s Gull, L. philadelphia M5 Ww Ring-billed Gull, L. delawarensis S3, M3 Ww, SS, g California Gull, L. californicus S5, M3 w, SS Herring Gull, L. argentatus S3, M3 w, SS, & Black-legged Kittiwake, Rissa tridactyla W6 w Caspian Tern, Sterna caspia M5 Ww Forster's Tern, S. forsteri S5 w | Black Tern, Chlidonias niger S5, M5 Ww ~COLUMBIFORMES _ Columbidae Rock Dove, Columba livia R2 sw Mourning Dove, Zenaida macroura Bl, M3, W5 SW |) STRIGIFORMES Strigidae Great Horned Owl, Bubo virginianus R3 sw Snowy Owl, Nyctea scandiaca W5 sw | Burrowing Owl, Athene cunicularia” B3, M3, W6 Ss, | Long-eared Owl, Asio otus B4, M4 d | Short-eared Owl, A. flammeus* B3, M3 Sse | Northern Saw-whet Owl, Aegolius acadicus S6, M6, W6 sw |CAPRIMULGIFORMES Caprimulgidae Common Nighthawk, Chordeiles minor B2, M3 Sw APODIFORMES Apodidae White-throated Swift, Aeronautes saxatalis SS d 520 GREAT BASIN NATURALIST Vol. 46, No. 3 Table 3 continued. Abundance’, season, Taxa and breeding status” Habitat® Trochilidae Rufous Hummingbird, Selasphorus rufus S3, M3 d CORACIIFORMES Alcedinidae Belted Kingfisher, Ceryle alcyon $3, M3 w PICIFORMES Picidae Downy Woodpecker, Picoides pubescens B5, M5 d Northern Flicker, Colaptes auratus B3, M3 d PASSERIFORMES Tyrannidae Olive-sided Flycatcher, Contopus borealis S5, M5 d Western Flycatcher, Empidonax difficilis S5 d Say’s Phoebe, Sayornis saya B3, M3 ss, d, f, j Ash-throated Flycatcher, Myiarchus cinerascens S5 d Western Kingbird, Tyrannus verticalis B3, M3 f, d, j Eastern Kingbird, T. tyrannus B3, M3 1 Gla Alaudidae Horned Lark, Eremophila alpestris R2 g, Ss Hirundinidae Tree Swallow, Tachycineta bicolor B3, M3 d, j Violet-green Swallow, T. thalassina B4, M4 r d, j Northern Rough-winged Swallow, Stelgidopteryx serripennis B3, M3 d,.j Bank Swallow, Riparia riparia B5, M3 Gl. Cliff Swallow, Hirundo pyrrhonota B2, M2 d, j Barn Swallow, H. rustica B2, M3 d, j Corvidae Clark’s Nutcracker, Nucifraga columbiana S4, M4, W5 j Black-billed Magpie, Pica pica R2 sw American Crow, Corvus brachyrhynchos R3 sw Common Raven, C. corax R3 sw Troglodytidae Rock Wren, Salpinctes obsoletus B3, M3 ss Canyon Wren, Catherpes mexicanus S5, M5 ss House Wren, Troglodytes aedon R3 d Muscicapidae Ruby-crowned Kinglet, Regulus calendula M3, W6 d Western Bluebird, Sialia mexicana* S5, M5 Ss Mountain Bluebird, S. currucoides S3, M3 ss Townsend's Solitaire, Myadestes townsendi S5, M5 d American Robin, Turdus migratorius B2, M2 sw Varied Thrush, Ixoreus naevius W6 ss Mimidae Northern Mockingbird, Mimus polyglottos S6 j Sage Thrasher, Oreoscoptes montanus B2, M2 ss Motacillidae Water Pipit, Anthus spinoletta M5 ss Bombycillidae Bohemian Waxwing, Bombycilla garrulus S3, M2, W3 fd Cedar Waxwing, B. cedrorum S5, M3, W5 fd Laniidae Northern Shrike, Lanius excubitor M3, W5 sw Loggerhead Shrike, L. ludovicianus* B3 ss July 1986 REYNOLDS ET AL.: IDAHO VERTEBRATES 521 Table 3 continued Abundance’, season, Taxa and breeding status” Habitat® Sturnidae European Starling, Sturnus vulgaris R3 sw Vireonidae Warbling Vireo, Vireo gilvus S5, M5 d Emberizidae Yellow Warbler, Dendroica petechia* B5, M3 d Yellow-rumped Warbler, D. coronata S3, M3 d Townsend's Warbler, D. townsendi M5 d American Redstart, Setophaga ruticilla M6 f Common Yellowthroat, Geothlypis trichas S5 d Wilson’s Warbler, Wilsonia pusilla S5, M5 d Yellow-breasted Chat, Icteria virens S5 d Western Tanager, Piranga ludoviciana S3, M3 d Black-headed Grosbeak, Pheucticus melanocephalus S5, M5 sw Lazuli Bunting, Passerina amoena S5, M5 d Green-tailed Towhee, Pipilo chlorurus S3, M3 ss Rufous-sided Towhee, P. erythrophthalmus S3, M3 sw Chipping Sparrow, Spizella passerina M5 f, d, ss Brewer s Sparrow, S. breweri B2, M2 Ss Vesper Sparrow, Pooecetes gramineus B3, M3 g, ss Lark Sparrow, Chondestes grammacus S3, M5 sw Black-throated Sparrow, Amphispiza bilineata S5, M5 ss Sage Sparrow, A. belli B2, M2 ss Lark Bunting, Calamospiza melanocorys S5, M5 ss Savannah Sparrow, Passerculus sandwichensis S5, M3 d,g Song Sparrow, Melospiza melodia S5, M3 d White-crowned Sparrow, Zonotrichia leucophrys M4 ss Dark-eyed Junco, Junco hyemalis M3 sw Snow Bunting, Plectrophenax nivalis W5 g, ss Red-winged Blackbird, Agelaius phoeniceus B3, M3 W, SS Western Meadowlark, Sturnella neglecta B2, M2, W3 g, ss Yellow-headed Blackbird, Xanthocephalus xanthocephalus B4, M3 w,d Brewer's Blackbird, Euphagus cyanocephalus B2, M2, W5 sw Brown-headed Cowbird, Molothrus ater B3, M3 ss Northern Oriole, Icterus galbula S3, M3 d Fringillidae Rosy Finch, Leucosticte arctoa M5, Wo SS House Finch, Carpodacus mexicanus $3, M3 fd Pine Siskin, Carduelis pinus S5, M3 f,d American Goldfinch, C. tristis M5 d, ss Evening Grosbeak, Coccothraustes vespertinus S5, M3 d Passeridae House Sparrow, Passer domesticus B2, M1, W3 f,d 6. 2 3 4. 5. | 1Abundance code (see text for definitions of terms): IL, Abundant . Common . Uncommon Occasional or local . Rare Vagrant or accidental Breeding and seasonal code: R = Breeder and year-round resident B = Summer breeder M =Migrant W = Winter visitor S = Summer visitor; no breeding records 3Habitat code (multiple habitats listed for a particular species are given in descending order of preference): w = on or near water ss = shrub-steppe d = deciduous or riparian j = juniper woodland g = grassland sw = sitewide f = facility complexes ~ 4Audubon Blue List (Tate and Tate 1982) Sensitive Species (USDI Bureau of Land Management) 5Endangered (Federal Register 1976) “Species of Special Concern (Idaho Department of Fish and Game, 1977) 522 GREAT BASIN NATURALIST Vol. 46, No. 3 TABLE 4. Mammals recorded on the Idaho National Environmental Research Park. Distribution Taxa and Habitat Abundance’ INSECTIVORA Soricidae Merriam Shrew, Sorex merriami Sitewide; sagebrush-steppe Uncommon CHIROPTERA Vespertilionidae Little Brown Myotis, Myotis lucifugus Sitewide; roosts in buildings Common Small-footed Myotis, M. leibii Sitewide; rocky outcrops & lava Abundant Long-eared Myotis, M. evotis SE NERP; junipers Common Big-brown Bat, Eptesicus fuscus Sitewide; roosts in buildings & caves Common Hoary Bat, Lasiurus cinereus Patchy; riparian & junipers Uncommon Townsend's Big-eared Bat, Plecotus townsendii Sitewide; caves & lava tubes Abundant LAGOMORPHA Leporidae White-tailed Jackrabbit, Lepus townsendii Sitewide; sagebrush-steppe Occasional Black-tailed Jackrabbit, L. californicus Sitewide; sagebrush-steppe Abundant- occasional (cyclic) Nuttall’s Cottontail, Sylvilagus nuttallii Sitewide; sagebrush-steppe, facilities Common Pygmy Rabbit, S. idahoensis* Patchy; sagebrush-steppe & rocky outcrops Common RODENTIA Sciuridae Least Chipmunk, Tamias minimus Sitewide; sagebrush-steppe Abundant Yellow-bellied Marmot, Marmota flaviventris Sitewide; rocky outcrops Uncommon Townsend's Ground Squirrel, Spermophilus townsendii Sitewide; sagebrush-steppe, facilities Common Geomyidae Northern Pocket Gopher, Thomomys talpoides Patchy; sagebrush-steppe Occasional Heteromyidae Great Basin Pocket Mouse, Perognathus parvus Sitewide; sagebrush-steppe Uncommon Ord’s Kangaroo Rat, Dipodomys ordii Sitewide; sagebrush-steppe & grassland Common Castoridae Beaver, Castor canadensis Patchy; Big Lost River Local Cricetidae Western Harvest Mouse, Reithrodontomys megalotis Sitewide; sagebrush-steppe & grassland Common Deer Mouse, Peromyscus maniculatus Sitewide; all habitats Abundant Northern Grasshopper Mouse, Onychomys leucogaster Sitewide; sagebrush-steppe Occasional Bushy-tailed Woodrat, Neotoma cinerea Sitewide; rocky outcrops Common Montane Vole, Microtus montanus Sitewide; grassland & facilities Abundant- occasional (cyclic) Sagebrush Vole, Lagurus curtatus” Patchy; sagebrush-steppe Uncommon Muskrat, Ondatra zibethicus Patchy; aquatic Rare Muridae Norw, facilities Common poides Patchy; sagebrush-steppe Occasional idae in Pocket Mouse, Perognathus parvus Sitewide; sagebrush-steppe Uncommon CARNIVORA Canidae Coyote, Canis latrans Sitewide; all habitats Common Mustelidae Long-tailed Weasel, Mustela frenata Sitewide; sagebrush-steppe Common Badger, Taxidea taxus Sitewide; all habitats Uncommon Western Spotted Skunk, Spilogale gracilis Sitewide; rocky outcrops Rare Felidae Mountain Lion, Felis concolor Sitewide; transient Vagrant Bobcat, F. rufus Sitewide; sagebrush-steppe, juniper Uncommon July 1986 REYNOLDS ET AL.: IDAHO VERTEBRATES 523 Table 4 continued. Distribution Taxa and Habitat Abundance! as rare ee a ae eR EA ee EES YS eee nd oo ee wa, a ees Ce ARTIODACTYLA Cervidae Wapiti, Cervus elaphus Mule Deer, Odocoileus hemionus Moose, Alces alces Antilocapridae* Pronghorn, Antilocapra americana Bovidae Mountain Sheep, Ovis canadensis 1S ee text for definition of abundance terms. - 2Green and Flinders (1980) place this in the genus Brachylagus. Carleton and Musser (1984) place this in the genus Lemmiscus. dominates the area. Winter hibernacula for several species of snakes have been located and described (Sehman 1977). With the ex- ception of the rubber boa, none of the reptiles recorded on the NERP were unexpected. Two species that were expected but not recorded on the Idaho NERP were the leop- ard frog (Rana pipiens) and the western racer (Coluber constrictor). Both have been col- _ lected within a few kilometers of the site (Seh- _ man and Linder 1978). With the possible ex- ception of these two species, and the possibility of minor range extensions or the discovery of some species in different habi- _ tats, the inventory of the herpetofauna on the Idaho NERP is probably complete. A total of 164 species of birds were recorded _on the Idaho NERP (Table 3). One hundred _ thirty-five species were recorded during the summer months, but only 59 (49%) of these _ were reported breeding on the site. Ember- _ izids (mostly sparrows and blackbirds), rap- | tors, waterfowl, and swallows accounted for nearly 50% of the breeding species. Breeding records were lacking for 76 species classified as summer visitors. Most of these were water- fowl, shorebirds, and emberizids (mostly war- blers and sparrows) with 13, 15, and 15 spe- cies, respectively. Forty-six bird species were _ recorded on site during the winter. Forty of _ these were also observed on the NERP during _ the summer. Six species were recorded dur- | ing the winter only or during the winter and | migration. Nineteen species (mostly shore- ‘birds and emberizids) were observed only _ during migration. _ Thirty percent of the wintering species recorded on the NERP were raptors. The { { | Sitewide; transient Vagrant Sitewide; sagebrush-steppe, grassland Uncommon Sitewide; transient Vagrant Sitewide; sagebrush-steppe, facilities Abundant North NERP; transient Vagrant 4O’Gara and Matson (1975) do not recognize Antilocapridae as a valid family and place pronghorn in the family Bovidae. number of birds of prey wintering on the NERP is tightly coupled to fluctuations in the black-tailed jackrabbit (Lepus californicus) population (Craig et al. 1983). During winters of low jackrabbit abundance (i.e., 1974-75 and 1975-76), an average of fewer than 15 raptors were recorded biweekly along a 187 km survey route. Eighty percent of these were Rough-legged Hawks (Buteo lagopus). During the winter of 1981-82, when the rab- bit population was high, each survey along the same route averaged nearly 142 raptors. Rough-legged Hawks and Golden Eagles (Aquila chrysaetos) accounted for 48% and 30% of the sightings, respectively. As many as 108 Golden Eagles and 15 Bald Eagles have been observed on the NERP in a single day (Watson 1984). Five species of gallinaceous birds were recorded on the NERP. The Sage Grouse (Centrocercus urophasianus ) was, however, the only common upland game bird on the park (Table 3). The NERP supports both win- tering and breeding populations of Sage Grouse (Connelly 1982, Connelly and Markham 1983, Gates 1983). Wintering flocks include birds that are year-round site resi- dents and large numbers of grouse that sum- mer in the mountain valleys and agricultural areas adjacent to the NERP (Connelly 1982). Sixty-seven sage grouse leks have been identi- fied on the site (Connelly and Ball 1983). Trost et al. (1977) listed 305 bird species in southeast Idaho. Of these, 216 occur in habi- tats similar to those found on the Idaho NERP. Thus, the avifauna records presented here represent about 75% of the species that potentially could occur on the Idaho NERP. 524 Increased efforts, especially during the breeding season, are needed to completely inventory the NERP avifauna. Thirty-nine species of mammals were recorded on the Idaho NERP (Table 4). With the exception of a few transient species (i.e., moose, [Alces alces|, wapiti [Cervus ela- phus|, mountain sheep [Ovis canadensis |, and mountain lion [Felis concolor |), the ma- jority of the species were resident on the NERP throughout the year. Most chiropter- ans presumably migrate from the NERP in the winter, but some of the many caves and lava tubes present on site are used as winter hibernacula. The extent of the wintering bat population is currently under investigation. Several predator-prey (Johnson 1978, John- son and Hansen 1979, Laundré et al. 1978, MacCracken 1980, MacCracken and Hansen 1982), synecology (Fraley et al. 1982, Gates 1983, R. Johnson 1982), and autecology (Fisher 1979, French et al. 1965, Grant 1983, Wilde 1978) studies conducted on the Idaho NERP addressed rabbits and hares. Although all leporid populations may be cyclic, num- bers of black-tailed jackrabbits were the most variable of the four species found on the NERP. Their densities ranged from less than 0.5/km* during population lows to more than 142/km* during the peak phase of the cycle (Stoddart 1983). All four leporids occurred sitewide, but the pygmy rabbit (Sylvilagus idahoensis ) appeared to be most specific in its habitat preference (Wilde 1978). Many studies on the NERP directly or indi- rectly addressed rodent populations (e.g., Allred 1973, Groves 1981, Groves and Keller 1983, Halford 1981, Halford and Markham 1978, M. Johnson 1982, Johnson and Keller 1983, Reynolds 1980). Seventeen species rep- resenting seven families were recorded on site (Table 4). No rodent species were particu- larly novel in their presence or conspicuous in their absence on the Idaho NERP. Most ro- dents were recorded throughout the research park. The montane vole (Microtus montanus ) and the western harvest mouse (Reithrodon- tomys megalotis) preferred grassland habitat. Predictably, the porcupine (Erythizon dorsa- tum), beaver (Castor canadensis), and muskrat (Ondatra zibethicus) were most abundant along the Big Lost River, although the latter was occasionally recorded in settling GREAT BASIN NATURALIST Vol. 46, No. 3 ponds near facilities and, even less fre- quently, in sagebrush habitat and agricultural fields. Six species of carnivores were recorded on the NERP (Table 4). Striped skunks (Mephitis mephitis ) were recorded within 10 km of the southern and southwestern boundary of the site but to date have not been reported on the NERP. With the exception of the spotted skunk (Spilogale gracilis), all occurred throughout the site. Coyotes (Canis latrans) and weasels (Mustela frenata) were the most abundant mammalian predators. Coyotes on the NERP were the focus of several com- pleted and ongoing studies (e.g., Davison 1980, Johnson 1978, Laundré 1979, Stoddart 1983, Woodruff 1977). Two studies have di- rectly addressed bobcats (Bailey 1972, Knick and Ball 1983). Populations of both species track but lag behind cyclic jackrabbit popula- tions. Studies on the other carnivores of the park are lacking. Pronghorn antelope (Antilocapra ameri- cana) were the most abundant and conspicu- ous of the big game animals on the research park. Populations of pronghorn fluctuate sea- sonally (Hoskinson and Tester 1980, Reynolds and Rose 1978, 1982). More than 4,500 pronghorn were recorded on the park during the winter of 1977-78. During typical sum- mers, about 750 pronghorn used the research park. Little information was available on the mule deer (Odocoileus hemionus) of the NERP. Deer were infrequently observed by NERP biologists and other site personnel. Onsite population densities, seasonal move- ments, and local habitat preferences have not been studied in detail. No fishes, amphibians, reptiles, or mam- mals recorded on the Idaho NERP were clas- sified as threatened or endangered species (U.S. Fish and Wildlife Service 1982). How- ever, both the endangered Peregrine Falcon and Bald Eagle were recorded on the NERP. Sightings of the former were rare during all seasons. The Bald Eagle was a consistent, al- beit infrequent, winter visitor to the research park. Although most of the mammals and her- petofauna were recorded throughout the site, less than 30% of the bird species were regis- tered either in shrub-steppe habitat or sitewide in occurrence. Alternatively, on or | July 1986 near water was recorded as the preferred habitat for nearly 40% of the avifauna. Only two species (Clark's Nutcracker [Nucifraga columbiana| and Northern Mockingbird [Mimus polyglottos |) were recorded in the juniper stands only. Blue Grouse (Dendraga- pus obscurus) and American Redstart (Se- tophaga ruticilla), both accidental species, were the only birds recorded around facilities only. Although all habitats were not used by wildlife in direct proportion to their availabil- ity, all habitats in the park evidently con- tribute somewhat to the overall diversity of vertebrate species on the NERP. The Idaho NERP provides a protected environment for an unexpected diversity and density of resi- dent and migrant vertebrate species repre- sentative of the Snake River Plain and North- ern Great Basin ecosystem. Undoubtedly the large expanse of the Idaho National Environ- mental Research Park, the controlled access requirements, and the relatively undisturbed native habitat are major factors that attract large wintering populations of pronghorn, raptors, and Sage Grouse. Although many ecological studies have been completed on the site since its NERP designation, contin- ued study is required to fully inventory some taxa and to answer some population and distri- bution questions about others. ACKNOWLEDGMENTS We thank J. E. Anderson, H. W. Browers, T. H. Craig, E. H. Craig, G. Devoe, R. J. Gates, D. L. Genter, J. C. Grant, C. R. Groves, M. K. Johnson, B. L. Keller, S. T. Knick, J. G. MacCracken, R. S. McCarty, K. L. Peterson, L. R. Powers, C. H. Trost, and J. W. Watson for aid in preparation of this spe- cies list. O. D. Markham, B. L. Keller, and K. S. Moor provided critical reviews of various drafts of the manuscript. This is a contribution of the Idaho National Engineering Laboratory Radioecology and Ecology Programs, funded by the Office of Health and Environmental Research, U. S. Department of Energy. LITERATURE CITED ALLRED, D. M. 1973. Small mammals of the National Reactor Testing Station. Great Basin Nat. 33: 246-250. REYNOLDS ET AL.: IDAHO VERTEBRATES 525 AMERICAN ORNITHOLOGISTS UNION. 1983. Check-list of North American birds. Allen Press, Lawrence, Kansas. 877 pp. ANDERSON, J. E., AND K. E. HOLTE. 1981. Vegetation de- velopment over 25 years without grazing on sage- brush-dominated rangeland in southeastern Idaho. J. Range Manage. 34: 25-29. ATWOOD, N. D. 1970. Flora of the National Reactor Tes- ting Station. Brigham Young Univ. Sci. Bull., Biol. Series 11(4): 1-46. BaILEy, T. N. 1972. Ecology of bobcats with special refer- ence to social organization. Unpublished disserta- tion, University of Idaho, Moscow. 82 pp. ———. 1974. Social organization in a bobcat population. J. Wildl. Manage. 38: 435-446. BEsT, L. B., AND K. L. PETERSON. 1982. Effects of stage of _ breeding cycle on sage sparrow detectability. Auk 99: 788-791. CARLETON, M. D., AND G. C. Musser. 1984. Muroid ro- dents. Pages 289-379 in S. Anderson and J. K. Jones, eds., Orders and families of recent mam- mals of the world. John Wiley and Sons, New York. 686 pp. CHOLEWA, A. F., AND D. M. HENDERSON. 1984. A survey and assessment of the rare vascular plants of the Idaho National Engineering Laboratory. Great Basin Nat. 44: 140-144. Couns, J. T., J. E. HUHEEYy, J. L. KNIGHT, AND H. M. SMITH. 1978. Standard common and current scien- tific names for North American amphibians and reptiles. Soc. Study Amph. Rept. Misc. Publ., Herp. Circ. 7: 1-36. CONNELLY, J. W. 1982. An ecological study of sage grouse in southeastern Idaho. Unpublished dissertation, Washington State University, Pullman. 84 pp. CONNELLY, J. W., ANDI. J. BALL. 1983. Sage grouse on the Idaho National Environmental Research Park. Pages 347-356 in O. D. Markham, ed., 1983 Pro- gress Report, Idaho National Engineering Labora- tory Radioecology and Ecology Programs. DOE/ ID-12098. Nat. Tech. Info. Serv., Springfield, Virginia. CONNELLY, J. W., AND R. J. Gates. 1981. First record of a black-legged kittiwake in Idaho. Condor 83: 272-273. CONNELLY, J. W., ANDO. D. MARKHAM. 1983. Movements and radionuclide concentrations of sage grouse in southeastern Idaho. J. Wildl. Manage. 47: 169- ie T. H. 1977. Raptors of the Idaho National Engi- neering Laboratory Site. Unpublished thesis, Idaho State University, Pocatello. 102 pp. . 1978. A car survey for raptors in southeastern Idaho. Raptor Res. 12: 40—45. . 1979. Raptors of the Idaho National Engineering Laboratory Site. IDO-12089. U.S. Dept. of En- ergy. Nat. Tech. Info. Serv., Springfield, Virginia. 27 pp. Cralc, T. H., E. H. Craic, AND L. R. Powers. 1983. Raptor studies on the Idaho National Engineering Labo- ratory. Pages 309-315 in O. D. Markham, ed., 1983 Progress Report, Idaho National Engineer- ing Laboratory Radioecology and Ecology Pro- grams. DOE/ID-12098. Nat. Tech. Info. Serv., Springfield, Virginia. CRAIG, 526 Cralc, T. H., AND F. RENN. 1977. Recent nesting of the Merlin in Idaho. Condor 57: 392. Davison, R. P. 1980. The effect of exploitation on some parameters of coyote populations. Unpublished dis- sertation, Utah State University, Logan. 139 pp. FISHER, J. S. 1979. Reproduction in the pygmy rabbit in southeastern Idaho. Unpublished thesis, Idaho State University, Pocatello. 33 pp. FRALEY, L., JR, G. C. BOWMAN, AND O. D. MARKHAM. 1982. Iodine-129 in rabbit thyroids near a nuclear fuel reprocessing plant in Idaho. Health Phys. 43: 251-258. FRENCH, N. R., R. MCBRIDE, AND J. E. DETMER. 1965. Fertility and population density of the black-tailed jackrabbit. J. Wildl. Manage. 29: 14-26. FULLER, R. K. 1981. Habitat utilization, invertebrate con- sumption, and movement by salmonid fishes un- der fluctuating flow conditions in the Big Lost River, Idaho. Unpublished thesis, Idaho State University, Pocatello. 76 pp. GaTEs, R. J. 1983. Sage grouse, lagomorphs, and pronghorn use of a sagebrush grassland burn site on the Idaho National Engineering Laboratory. Unpublished thesis, Montana State University, Bozeman. 135 pp. GLEASON, R. S. 1978. Aspects of the breeding biology of burrowing owls in southeastern Idaho. Unpub- lished thesis, University of Idaho, Moscow. 47 pp. GRANT, J. C. 1983. Black-tailed jackrabbit movements and habitat utilization at the Idaho National Engineer- ing Laboratory Radioactive Waste Management Complex. Pages 278-282 in O. D. Markham, ed., 1983 Progress Report, Idaho National Engineer- ing Laboratory Radioecology and Ecology pro- grams. DOE/ID-12098. Nat. Tech. Info. Serv., Springfield, Virginia. 434 pp. GREEN, J. S., AND J. T. FLINDERS. 1980. Brachylagus ida- hoensis. Mammalian species 125: 1—4. GROVES, C. R. 1981. The ecology of small mammals on the Subsurface Disposal Area, Idaho National Engi- neering Laboratory Site. Unpublished thesis, Idaho State University, Pocatello. 87 pp. GrovEs, C. R., AND B. L. KELLER. 1983. Ecological charac- teristics of small mammals on a radioactive waste disposal area in southeastern Idaho. Amer. Mid. Nat. 109: 253-265. HALForD, D. K. 1981. Repopulation and food habits of Peromyscus maniculatus on a burned sagebrush desert in southeastern Idaho. Northwest Sci. 55: 44-49, HALFoRD, D. K., AND O. D. MARKHAM. 1978. Radiation dosimetry of small mammals inhabiting a liquid radioactive waste disposal area. Ecology 59: 1047-1054. HALForD, D. K., AND J. B. MILLARD. 1978. Vertebrate fauna of a radioactive leaching pond complex in southeastern Idaho. Great Basin Nat. 38: 64-70. Harniss, R. O., AND N. E. WEsT. 1973. Vegetation pat- terns on the National Reactor Testing Station, southeastern Idaho. Northwest Sci. 47: 30-43. HOSKINSON, R. L. 1977. Migration behavior of pronghorn antelope and summer movements and fall migra- tions of pronghorn fawns in southeastern Idaho. Unpublished dissertation, University of Minne- sota, Minneapolis. 125 pp. GREAT BASIN NATURALIST Vol. 46, No. 3 HOSKINSON, R. L., AND J. R. TESTER. 1980. Migration be- havior of pronghorn antelope in southeastern Idaho. J. Wildl. Manage. 44: 132-144. IDAHO DEPARTMENT OF FISH AND GAME. 1977. A plan for future management of Idaho's fish and wildlife resources. Vol. 1, Goals, objectives, and policies. Idaho Dept. Fish and Game, Boise. 215 pp. JEPPSON, R. J., AND K. E. HOLTE. 1978. Flora of the Idaho National Engineering Laboratory Site. Pages 129-143 in O. D. Markham, ed., Ecological stud- ies on the Idaho National Engineering Laboratory Site, 1978 Progress Report. IDO-12087. Nat. Tech. Info. Serv., Springfield, Virginia. 370 pp. JOHNSON, M. K. 1978. Food habits of coyotes in southcen- tral Idaho. Unpublished dissertation, Colorado State University, Fort Collins. 75 pp. —___. 1982. Response of small mammals to livestock grazing in south-central Idaho. J. Range Manage. 35: 51-53. JOHNSON, M. K., AND R. M. HANSEN. 1979. Coyote food habits on the Idaho National Engineering Labora- tory. J. Wildl. Manage. 43: 951-956. JoHNsoN, R. D. 1982. Relationships of black-tailed jackrabbit diets to population density and vegetal components of habitat. Unpublished thesis, Idaho State University, Pocatello. 86 pp. JOHNSON, W. C. 1977. Examination of censusing tech- niques for small mammals in a high desert ecosys- tem. Unpublished thesis, Idaho State University, Pocatello. 95 pp. A JOHNSON, W.C., AND B. L. KELLER. 1983. An examination of snap-trapping techniques for estimating rodent density in high desert. Northwest Sci. 57: 194-204. Jones, J. K., Jk., D. C. Carter, H. H. GENoways, R. S. HOFFMAN, AND D. W. RICE. 1982. Revised check- list of North American mammals north of Mexico, 1982. Occas. Papers Mus., Texas Tech University 62: 1-17. KNICK, S.T., ANDI. J. BALL. 1983. Ecology of exploited and unexploited bobcat populations in southeastern Idaho. Pages 319-324 in O. D. Markham, ed., 1983 Progress Report, Idaho National Engineer- ing Laboratory Radioecology and Ecology Pro- grams. DOE/ID-12098. Nat. Tech. Info. Serv., Springfield, Virginia. 434 pp. LAUNDRE, J. W. 1979. A behavioral study of home range utilization by coyotes on the INEL Site in south- eastern Idaho. Unpublished dissertation, Idaho State University, Pocatello. 70 pp. LAUNDRE, J. W., R. Davison, M. K. JOHNSON, B. L. KELLER, AND D. B. WILDE. 1978. Coyote-prey as- sessment on a NERP site in southeastern Idaho. Pages 55-76 in J. T. Kitchings, and N. E. Tarr, eds., National Environmental Research Park Symposium: Natural resource inventory, charac- terization, and analysis. ORNL-5304. Oak Ridge National Laboratory, Oak Ridge, Tennessee. 184 pp: LINDER, A. D., AND R. W. SEHMAN. 1978. The herpeto- fauna of the Idaho National Engineering Labora- tory Site. J. Idaho Acad. Sci. 13: 47-50. MacCrackENn, J. G. 1980. Feeding ecology of coyotes on the upper Snake River Plain, Idaho. Unpublished thesis, Colorado State University, Fort Collins. 85 pp. July 1986 MaAcCRACKEN, J. G., AND R. M. HANSEN. 1982. Herba- ceous vegetation of habitat used by black-tailed jackrabbits and Nuttall cottontails in southeastern Idaho. Amer. Midl. Nat. 107: 180-184. MarkKHaM, O. D. 1973. National Reactor Testing Station environmentally related publications. IDO- 12078. AEC Idaho Operations Office, Idaho Falls. 11 pp. . 1978. 1978 Progress Report. Ecological studies on the Idaho National Engineering Laboratory Site. IDO-12087. Nat. Tech. Info. Serv., Springfield, Virginia. 370 pp. . 1983. Idaho National Engineering Laboratory Ra- dio-Ecological and Ecological Studies 1983 Pro- gress Report. DOE/ID-12098. Nat. Tech. Info. Serv., Springfield, Virginia. 434 pp. MARLETTE, G. M. 1982. Stability and succession in crested wheatgrass seedings on the Idaho National Engi- neering Laboratory Site. Unpublished thesis, Idaho State University, Pocatello. 100 pp. MCBRIDE, R., N. R. FRENCH, A. H. DAHL, AND J. E. DET- MER. 1978. Vegetation types and surface soils of the Idaho National Engineering Laboratory Site. IDO-12084. Nat. Tech. Info. Serv., Springfield, Virginia. 28 pp. Nace, R. L., M. DEUTSCH, AND P. T. VOEGELI. 1972. Physi- cal environment of the National Reactor Testing Station, Idaho—a summary. U.S. Dept. Int., Geol. Surv. Prof. Paper 725-A. NACE, R. L., P. T. VOEGELI, J. R. JONES, AND M. DEUTSCH. 1975. Generalized geologic framework of the Na- tional Reactor Testing Station, Idaho. U.S. Dept. Int., Geol. Surv. Prof. Paper 725-B. NussBaAuM, R. A., E. D. BRODIE, JR., AND R. M. STORM. 1983. Amphibians and reptiles of the Pacific Northwest. University of Idaho Press, Moscow. 330 pp. O Gara, B. W., ANDG. MATSON. 1975. Growth and casting of horns by pronghorns and exfoliation of horns by bovids. J. Mammal. 56: 829-846. OVERTON, C. K. 1977. Description, distribution, and den- sity of the Big Lost River salmonid populations. Unpublished thesis, Idaho State University, Pocatello. 51 pp. PETERSON, K. L. 1982. Breeding ecology of passerine birds in a sagebrush-dominated community. Unpublished the- sis, lowa State University, Ames. 66 pp. Powers, L. R., AND T. H. Craic. 1976. Notes on the status of nesting ferruginous hawks (Buteo regalis) in Little Lost River Valley and vicinity, southeastern Idaho. Murrelet 57: 46. REYNOLDS, T. D. 1978. The response of native vertebrate populations to different land management prac- tices on the Idaho National Engineering Labora- tory Site. Unpublished dissertation, Idaho State University, Pocatello. 105 pp. . 1980. The effects of some different land manage- ment practices on small mammal populations. J. Mammal. 61: 558-561. REYNOLDS ET AL.: IDAHO VERTEBRATES 527 REYNOLDS, T. D., AND F. L. Rose. 1978. Pronghorn an- telope use of the INEL National Environmental Research Park. Pages 219-233 in O. D. Markham, ed., Ecological studies on the Idaho National En- gineering Laboratory Site. I[DO-12087. Nat. Tech. Info. Serv., Springfield, Virginia. 372 pp. . 1982. Pronghorn antelope use of the Idaho Na- tional Environmental Research Park. Page 63 in O. D. Markham, ed., Idaho National Engineering Laboratory, Hanford Site, and Los Alamos Na- tional Laboratory symposium on radioecology and ecology. IDO-12095. U.S. Dept. of Energy, Idaho Operations Office, Idaho Falls. 63 pp. REYNOLDS, T. D., AND C. H. Trost. 1980. The response of native vertebrate populations to crested wheat- grass planting and grazing by sheep. J. Range Manage. 33: 122-125. SEHMAN, R. W. 1977. Hibernaculum dynamics of the great basin rattlesnake (Crotalus viridis lutosus). Unpublished thesis, Idaho State University, Pocatello. 66 pp. SEHMAN, R. W., AND A. D. LINDER. 1978. Amphibian and reptilian fauna of the Idaho National Engineering Laboratory Site. IDO-12086. U.S. Dept. of En- ergy. Nat. Tech. Info. Serv., Springfield, Virginia. 26 pp. SIMPSON, J.C., AND R. L. WALLACE. 1978. Fishes of Idaho. University of Idaho Press, Moscow. 237 pp. STEPHENS, D. A., AND T. D. REYNOLDS. 1983. Birds of southwestern Idaho. Great Basin Nat. 43: 728- 738. StToppakT, L. C. 1983. Relative abundance of coyotes, lagomorphs, and rodents on the Idaho National Engineering Laboratory. Pages 268-277 in O. D. Markham, ed:, 1983 Progress Report, Idaho Na- tional Engineering Laboratory Radioecology and Ecology Programs. DOE/IDO-12098. Nat. Tech. Info. Serv., Springfield, Virginia. 434 pp. TATE, J., JR., AND D. J. TATE. 1982. The Blue List for1982. Amer. Birds 36: 126-135. Trost, C. H., S. L. FINDHOLT, AND T. D. Ricu. 1977. Birds of Idaho field check-list. Dept. of Biology, Idaho State University, Pocatello. 2 pp. U.S. FISH AND WILDLIFE SERVICE. 1982. List of endan- gered and threatened wildlife and plants. Bureau of National Affairs, Inc. (50 CFR 17.11) Washing- ton, D. C. WarTSON, J. W. 1984. Rough-legged Hawk winter ecology in southeastern Idaho. Unpublished thesis, Mon- tana State University, Bozeman. 125 pp. WILDE, D. B. 1978. A population analysis of the pygmy rabbit (Sylvilagus idahoensis) on the INEL Site. Unpublished dissertation, Idaho State University, Pocatello. 172 pp. WooprvurfF, R. A. 1977. Annual dispersal, daily activity pattern, and home range of Canis latrans on the Idaho National Engineering Laboratory Site. Un- published thesis, Idaho State University, Pocatello. 125 pp. INFECTION OF YOUNG DOUGLAS-FIRS AND SPRUCES BY DWARF MISTLETOES IN THE SOUTHWEST Robert L. Mathiasen! ABSTRACT. —Mistletoe infection of Douglas-fir and spruce seedlings increased as the mean dwarf mistletoe rating of the overstory, seedling density, and total age of seedlings increased. Percent of spruce seedlings infected increased more rapidly than for Douglas-fir as overstory dwarf mistletoe ratings increased. However, the intensity of infection as measured by the mean dwarf mistletoe rating of seedlings, increased at about the same rate for spruce and Douglas-fir. Percent infection of seedlings less than 20 years total age was higher for spruce than for Douglas-fir. Douglas-fir and spruce seedling mortality was from two to three times greater in heavily infested stands than in healthy stands. Dwarf mistletoes (Arceuthobium spp.) are the most serious disease agent in southwest- ern mixed conifer forests (Jones 1974). These parasitic flowering plants reduce the growth of heavily infected trees, cause increased mor- tality, reduce cone and seed production, and may predispose trees to other diseases and insects (Hawksworth and Wiens 1972). The most prevalent and damaging dwarf mistle- toe, A. douglasii Engelm. It occurs through- out the range of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) in the Southwest. Andrews and Daniels (1960) estimated that approximately 50% of the Southwest's Dou- glas-fir type was infested with Douglas-fir dwarf mistletoe. Another damaging dwarf mistletoe in southwestern forests is the west- ern spruce dwarf mistletoe, Arceuthobium microcarpum (Engelm.) Hawksw. & Wiens. It parasitizes Engelmann and blue spruce (Picea engelmannii Parry and P. pungens En- gelm.) in several locations in Arizona, but it is only known from the Mogollon and Sacra- mento mountains in New Mexico (Hawks- worth and Graham 1963a, Hawksworth and Wiens 1972, Mathiasen and Jones 1983). Western spruce dwarf mistletoe is most prevalent in the Apache-Sitgreaves National Forest, Arizona, where it is a primary factor associated with mortality in infested spruce stands (Hawksworth and Graham 1963a). Douglas-fir and spruce are the most preva- lent and commercially important species in southwestern mixed conifer forests. Douglas- fir and spruce regeneration is frequently 'School of Forestry, Box 4098, Northern Arizona University, Flagstaff, Arizona 86011. 528 abundant in the understory of mixed conifer stands in the Southwest (Moir and Ludwig 1979, Gottfried and Embry 1977). When overstories are infested with dwarf mistletoes, spread to young regeneration perpetuates the infestation. Management of mixed conifer forests infested with dwarf mistletoes should minimize the spread of the disease from an infested overstory to the understory and par- ticularly to young regeneration (Jones 1974, Gottfried and Embry 1977). There are several factors that influence in- fection of regeneration. Natural infection of seedlings is not common because they repre- sent a relatively small target area for mistletoe seeds (Hawksworth 1958, 1961, Wicker and Shaw 1967), and several factors remove seeds from seedlings (Wicker 1965, 1967). Scharpf (1969) reported that only 7% of true fir seedlings less than three feet tall were in- fected in heavily infested stands and that in- fection intensified rapidly in regeneration taller than three feet. The duration of exposure to inoculum is an important factor influencing seedling infec- tion. Hawksworth (1958) found there is little infection in lodgepole pine (Pinus contorta Doug]. ex Loud.) stands less than 15 years old. Hawksworth and Graham (1963b) reported infection of lodgepole pine as young as 6 years in the intermountain region and 8 years in the | central Rocky Mountains. Weir (1918) re- ported the average age of 50 naturally infected Douglas-fir seedlings, used for assessing the | effects of dwarf mistletoe on seedling growth, |, July 1986 was 18 years, but he did not report the youngest age for the seedlings. Infection of ponderosa pine (Pinus ponderosa Laws.) by ~ southwestern dwarf mistletoe (Arceuthobium vaginatum subsp. cryptopodum (Engelm.) Hawksw. & Wiens) has been documented for 10-year-old seedlings (Gill and Hawksworth 1954, Hawksworth 1961). Infection of regeneration in dwarf mistle- toe—infested stands has important manage- ment implications, but information concern- _ ing infection of young regeneration is not available for most species growing in south- western mixed conifer stands. Therefore, this investigation was undertaken to provide data on the infection of young Douglas-fir and _ spruce seedlings in dwarf mistletoe-infested mixed conifer stands in the Southwest. MATERIALS AND METHODS / In 1979, 150 temporary 0.2 acre plots were established in mixed conifer stands in the Apache-Sitgreaves National Forest, Arizona. Of these, 123 were infested with Douglas-fir dwarf | mistletoe. For each Douglas-fir greater than 4.5 feet in height, diameter at breast height (dbh) | was measured to the nearest 0.1 inch. All re- | maining trees greater than 4.5 feet in height |) were measured at dbh to the nearest 2.0 inch. _ Dwarf mistletoe ratings (Hawks-worth 1977) were recorded for all living trees and for dead trees whenever possible. _ The dwarf mistletoe rating system used di- j vided the live crown of a tree into thirds, and each third was rated separately as: 0, no mistle- “toe infection; 1, less than 50% of the live H branches infected: 2, more than 50% of the live branches infected (Hawksworth 1977). The rat- ings for each third were totaled to obtain a dwarf mistletoe rating (DMR) for the tree. Mean over- jstory_and seedling dwarf mistletoe ratings ~ (DMR) were calculated by adding the DMRs for all live overstory trees (dbh greater than 1:0) and seedlings (dbh 0.11.0) in a stand and dividing the total by the number of trees or seedlings. Infection rates were based on the percentage of live trees in a stand or plot with one or more observable mistletoe infections. Infection inten- ‘sities were based on the DMR of the overstory of “seedlings. _ During 1980 and 1981, 237 temporary ‘rectangular plots ranging in size from 0.1 to 2 2525 rote: o> 25 2 roe res se 25 ro res re! 2 res oS 25 25 2S res ros ree re $33 ozs ose eee S333 33333 3533 $2505 <> 2S res 25 2! 3 ree 2 sess 23 3 2 5°, 3 2s sos aes 2525 ote. < 2 res x3 ree 2 55 ae 2S SS 2: eS ree 2 res rat res a5 = S $3 5 = . S35 2: <> ree = TITS Bases pesese 2; S555 ies veces rotet cemereret Soe 2; ies ret aS oo> tet = Sesesesets: tete%e = 3 3 ae: a. 23 hrererarsrsP SPIER Feoscors Soh ret res RS EStsz3 Peal scene: bs 0.9 o ua Oo o ° eee q BS oO fo) 0.1 0.2 0.3 0.4 Fig. 2A. Number of deer beds found at various slope positions within mesic areas. 1.0 = top of the mountain. 0.1 0.2 0.3 0.4 05 06 0.7 08 0.9 1.0 Fig. 2B. Number of deer beds found at various slope positions within xeric areas. 1.0 = top of the mountain. established trails when fleeing from danger or locating a resting spot than to forge through dense brush. Trails along which deer fre- quently bed, therefore, are the travel lanes connecting feeding, watering, bedding, and escape areas of deers home range (Dasmann and Taber 1956). Of the remaining beds that were not located on a trail, most were in dense cover close to trails. Significant differences were noted in char- acteristics of deer beds between xeric and mesic communities with respect to elevation of beds and percentage of cover projected over the bed. Beds in xeric habitats were on the average located at higher slope positions than those in mesic communities (Fig. 2). Analysis of variance indicated that the mean percentage of projected cover over the beds in xeric areas (12.7%, n = 12) was significantly less (P = .01) than in mesic areas (38%, n = 29). This seems obvious when considering the short vegetation of xeric communities, but it may explain why xeric beds are higher altitu- dinally. Deer that bed in xeric habitats com- pensate for lack of cooling shade by selecting | | | July 1986 bedding sites at higher elevations where up- drafting occurs and temperatures are cooler. Since bucks predominate in xeric communi- ties (King and Smith 1980), the possible ther- moregulatory function of antlers may help males compensate for the lack of shade in drier environments. This would be particu- larly true if wind currents play a role in heat dispersal. Bedding in more open higher places, without heavy concealing vegetation, provides deer with a visual advantage over predators. Many beds were located along the top of a ridge or just below the crest, where the bedded animal had commanding views of the surrounding areas. The percentage of cover variance on the uphill side of the bed between mesic and xeric areas indicated an important trend. There was a significant difference (P = .01, F = 12.37) between cover on the uphill side of beds lo- cated in xeric areas compared to those in more mesic areas. Mean uphill cover in xeric com- munities was 79.1% (n = 12) compared to 41.0% (n = 29) for mesic environments. Com- parisons of percent downhill cover and on both lateral sides of the beds showed no signif- icant differences between the two habitat types. The greater amount of protective uphill cover for more open, xeric beds not only broke up the outline of the deer but provided greater visual and physical protection from predators approaching the blind side of the deer. Predators, such as cougars, are forced to go around the obstacle provided by more dense cover instead of making a direct rushing attack on the resting deer. This gives bedded deer the advantage of a head start. Analyzing percent visibility that can be seen of the bed from the four compass points showed a common trend that holds for xeric and mesic communities. The lowest percent visibility is from the north (Table 1). This may afford greater protection from cold north winds and heat radiation to an open, air-circu- lating environment. There was also decreased visibility on the west side of the bed in both _ habitat types. This may be advantageous by | affording protection to bedded deer from the "i hot afternoon sun. Analysis of variance showed that the mean bed size does not vary significantly from xeric to mesic communities (F = 0.86, P = .01). This is a little surprising since Linsdale and SMITH ET AL.: MULE DEER BEDS 945 TABLE 1. Mean percent visibility from deer beds in north, south, east, and west directions for xeric and mesic communities. Xeric (%) Mesic (%) Visibility north 9.1 38.1 Visibility south 29.6 58.6 Visibility east 28.2 56.8 Visibility west 14.1 48.6 Tomich (1953) report that deer beds conform closely to the size and shape of the body of the reclining deer and more males were found in xeric habitats. It is likely that there is no sig- nificant size difference between females and males observed. A general observation was that no matter how steep the incline, deer beds are situated horizontally. Many beds were located in level spots created by the earth-leveling effect of tree roots. Linsdale and Tomich (1953) re- ported that deer bedded on steep slopes in which loose soil has been pushed downhill, creating a level site. Trails provided level rest- ing spots on steep hillsides upon which deer frequently bedded. Beds were selected in places where foliage was especially impenetrable and dense, providing almost complete concealment, or in high, open places where visibility was good and many escape routes were available. Does and fawns bedded more often in thickly vege- tated areas. They benefit from heavy cover and rely on concealment to avoid danger. Concealment is sometimes employed by ma- ture bucks. They often remain bedded rather than flush in response to heavy hunting pres- sure. In summarizing this study, more beds were found on mesic, north-facing slopes than xeric, south-facing slopes. North-facing slopes are cooler, more densely covered, and provide a food supply with higher water con- tent than xeric habitats. Such bedding sites provide thermoregulatory, concealment, and nutritional needs of deer, particularly fe- males, during summer months. Many beds, used predominantly by males, were located immediately above a precipice or below cliffs, especially where steep slopes run down from the base of the cliff. This provides protection from approach on one side, excellent vision below, and convenient escape routes. Other areas where beds were consistently located 546 included small benches or flat areas on moun- tainsides. These provide level places to lie on as well as excellent vantage points and backdrop cover. Included as favorite sites were shoulders or points of big ridges and patches of sagebrush or other short brush in open country. In some areas on Timpanogos, timberline is not consis- tent, but rather it extends in long fingers under- neath cliffs. Deer often bed near the crest of the slope where these narrow strips of timber project. LITERATURE CITED Bowyer, T. R. 1984. Sexual segregation in southern mule deer. J. Mammal. 65: 410-417. DarLING, F. F. 1937. A herd of red deer. Oxford University Press, London. 215 pp. DasMANN, R. F., AND R. D. TaBER. 1956. Behavior of Colum- bian blacktailed deer with reference to population ecology. J. Mammal. 37: 143-164. FLINDERS, J. T., AND C. L. ELuiorr. 1979. Abiotic characteris- tics of black-tailed jackrabbit forms and a hypothesis concerning form function. Encyclia 56: 34-38. Geist, V. 1981. Adaptive strategies in mule deer. Pages 157-223 in O. C. Wallmo, ed., Mule and black-tailed deer of North America. University of Nebraska Press, Lincoln. GREAT BASIN NATURALIST Vol. 46, No. 3 KiNG, M. M., AND H. D. Smiru. 1980. Differential habitat utilization by the sexes of mule deer. Great Basin Nat. 40: 273-281. KING, R. T. 1938. The essentials of a wildlife range. J. For. 36 (5): 457-464. LINSDALE, J. M., AND P. Q. Tomicu. 1953. A herd of mule deer. University of California Press, Berkeley and Los Angeles. 367 pp. McCu..oucu, D. R. 1979. The George Reserve deer herd: population ecology of a K-selected species. University of Michigan Press, Ann Arbor. 271 pp. McRae, B. 1980. The trouble with muleys. Outdoor Life 166 (4): 58-61. MILLER, F. L. 1968. Observed use of forage and plant communities by black-tailed deer. J. Wildl. Man- age. 32: 142-148. ____. 1970. Distribution patterns of black-tailed deer (Odocoileus hemionus columbianus) in relation to environment. J. Mammal. 51: 248-259. Moen, A. N. 1973. Wildlife ecology. An analytical ap- proach. W. H. Freeman and Co., San Francisco, California. 458 pp. SHort, H. L. 1981. Nutrition and metabolism. Pages 99-127 in O. C. Wallmo, ed., Mule and black- tailed deer of North America. University of Ne- braska Press, Lincoln. STONEHOUSE, B. 1968. Thermoregulatory function of growing antlers. Nature 218: 870-872. TRUMPETER SWAN (CYGNUS BUCCINATOR) FROM THE PLEISTOCENE OF UTAH Alan Feduccia' and Charles G. Oviatt” ABSTRACT.—A Trumpeter Swan (Cygnus buccinator) is reported from Pleistocene deposits in Utah. Among fossil bones recovered from a de- posit of Pleistocene age in Utah are numerous elements of the Trumpeter Swan (Cygnus buccinator)’. These fossils were collected by Oviatt from pre—Lake Bonneville marginal la- custrine deposits in an exposure along the West Side Canal in the NW 1/4, Sec 4, T12N, R2W, Cutler Dam, Utah, 7.5 minute quad- rangle. The exposure is approximately 140 ft above the Bear River at an altitude of about 4,400 ft. The bones were excavated from a silty clay unit that contained many gas- tropods, including the genera Helisoma, Lymnaea, and Valvata. Amino acid ratios on gastropods from this locality (W. D. McCoy, personal communica- tion, 1984) and available radiocarbon dates indicate that the marginal lacustrine deposits are approximately 40,000 to 65,000 years old and are part of a sequence of pre—Lake Bon- neville lacustrine beds well exposed in this area (Oviatt et al., 1985: 260). A soil profile at the top of the marginal lacustrine deposits is overlain by lacustrine deposits of the Bon- neville Alloformation. The marginal lacus- trine deposits were deposited near the shore of a lake that rose to a maximum altitude of slightly less than 4,400 feet. The presence ofa lake of this size in the Bonneville basin indi- cates that the climate was relatively cool or moist at the time the Trumpeter Swan bones were deposited. The elements recovered include parts of two humeri, a coracoid, radius, ulna, scapula, and pieces of vertebrae, all of which are assignable to C. buccinator on the basis of larger size and more robust nature when com- pared to similar elements of modern Cygnus columbianus , the Tundra Swan. The Trumpeter Swan, the largest living swan, lives on ponds, lakes, and marshes dur- ing the breeding season, when it occurs in Alaska, and in parts of western Canada and the United States, south from Saskatchewan to southeastern Oregon, eastern Idaho, and northwestern Wyoming. It bred formerly south to Nebraska, Iowa, Missouri, and Indi- ana. During the winter months this swan oc- curs over parts of Alaska and western Canada, south to California, and occasionally to Utah, New Mexico, and Colorado (American Or- nithologists’ Union, 1983: 63-64). It formerly wintered south to the Mexican border, the Gulf Coast of Texas and Louisiana, and the Mississippi Valley and on the Atlantic Coast from New Jersey and Pennsylvania to North Carolina (Banko 1960: 26). It is known from Pleistocene deposits from Oregon, Illinois, and Florida and from prehistoric sites from Alaska, Iowa, Illinois, and Ohio (Brodkorb 1964: 233). Thus, this is the first Pleistocene record of the Trumpeter Swan from Utah and could well be an additional indication of the more expansive range occupied by this spe- cies in the past, because it presently only occasionally visits Utah during the winter months. Trumpeter Swans feed primarily in shallow water, plunging the head and neck below the surface in their quest for aquatic plants grow- ing on the bottom. This type of habitat con- forms to the picture of the deposits from which these bones were recovered. 1Department of Biology, University of North Carolina, Chapel Hill, North Carolina 27514. 5 2: 4 ’ ON tess 2Utah Geological and Mineral Survey, 606 Black Hawk Way, Salt Lake City, Utah 84108. Present address: Department of Geology, Kansas State University, Manhattan, Kansas 66506. 3The swan bones are housed with the vertebrate paleontology collections at the Antiquities Section, Utah Division of State History, 300 Rio Grande, Salt Lake City, Utah 84101. Catalogue number UVPO99, collection locality number 42Bo049v. 547 LITERATURE CITED AMERICAN ORNITHOLOGISTS’ UNION. 1983. Check-list of North American Birds, 6th ed. Allen Press, Lawrence, Kansas. 877 pp. Banko, W. E. 1960. The Trumpeter Swan. N. Amer. Fauna no. 63. U.S. Dept. Inter., Fish and Wildlife Service, Washington, D.C. 214 pp. 048 GREAT BASIN NATURALIST Vol. 46, No. 3 Bropkorp, P. 1964. Catalogue of fossil birds: part 2 (Anseri- formes through Galliformes). Bull. Florida State Mus. 8(3): 195-335. Oviatt, C. G., W. D. McCoy, AND R. G. REIDER. 1985. Quater- nary lacustrine stratigraphy along the lower Bear River, Utah: evidence for a shallow early Wisconsin lake in the Bonneville Basin. Geological Society of America Abstracts with Programs 17(4): 260. NEW SPECIES AND A NEW COMBINATION OF MENTZELIA SECTION BARTONIA (LOASACEAE) FROM THE COLORADO PLATEAU H. Thompson’ and B. Prigge” ABSTRACT.—A new species, Mentzelia (sect. Bartonia) cronquistii, subfamily Mentzelioideae, is described and a new combination, M. (sect. Bartonia) marginata (Osterhout) is made. These two species of the Colorado Plateau are closely related to each other, but their relationships to other species in section Bartonia are obscure. The plants of Mentzelia sect. Bartonia in the Colorado Plateau of eastern Utah, western Colorado, and northern Arizona show very high morphological diversity that has not been easy to analyze by standard collection and herbarium methods. The opportunity for great morphological diversity has been pro- vided by the very high habitat diversity, the principle component of which is substrate di- versity. The Colorado Plateau is composed of a variety of strata, intruded by igneous rock and tilted with a northeast dip. The Colorado River has cut through the Plateau, against the dip, exposing the various strata. A second component of this habitat diversity is eleva- tion, for the various substrates are exposed over a gradient of over a thousand meters. A third component of diversity is rainfall, which varies geographically and with elevation. This paper factors two species from the diversity in Mentzelia (sect. Bartonia) in the Colorado Plateau, and, although recognition of these two species makes understanding the remain- ing diversity somewhat easier, there still exist some local endemics that need recognition, and we feel certain that additional endemics remain to be discovered. Plants of both Mentzelia cronquistii and M. marginata have been grown in the experi- mental garden at UCLA, where hybridiza- tions have been made using standard tech- niques of emasculation and pollen exclusion. Chromosomes have been observed with phase microscopes in squashed microsporo- cytes from buds fixed in 3:1 ethanol/ascetic acid. Seeds have been coated then examined and photographed with a scanning electron Sun City West, Arizona 85351. microscope. SEM photographs and voucher herbarium specimens for the hybridization and chromosome studies are deposited at RSA. Mentzelia marginata (Osterhout) Thomp- son & Prigge, comb. nov. Nuttallia marginata Osterhout. Bull. Torrey Bot. Club 49: 183. 1922. TypE: COLORADO, Mesa Co., DeBeque, on the hills north of town, August 22, 1918, Osterhout 5842 (Holo- type: RM!). Herbaceous perennial up to 25 cm tall; stems erect, single or much branched at base, white, pubescent with both glochidiate and pointed hairs; rosette leaves oblanceolate, crenate or sinuate; lower cauline leaves oblanceolate to broadly ovate, 3-6 cm long, 1-2 cm wide, with the lowermost petiolate, with upper ones smaller, oval, sessile, broad and somewhat clasping; lower leaf surfaces densely pubescent with long and_ short glochidiate hairs and scattered pointed hairs; upper leaf surfaces more sparsely pubescent with pointed hairs; leaf margins with large glochidiate hairs; bract at base of capsules lin- ear and entire; flowering period June through August; flowers opening in late afternoon, ca- lyx lobes 5-8 mm long; petals 5, yellow, 10-13 mm long, 2-5 mm wide, pubescent on outer surface, ovate to narrowly ovate, acute at apex; the next whorl within the petals 5 petaloid stamens, pubescent at base, similar in shape to the petals but smaller, only 3 mm wide and with functional anthers; stamens nu- merous, grading in length from 3 mm for the innermost to 10 mm for the outermost, with narrow filaments for innermost ones and "Department of Biology, University of California, Los Angeles, California 90024. 549 cronqu/stt/ n=10 @ @ 2 ad oe? of e® Ox m~ % | ! ! | GREAT BASIN NATURALIST Vol. 46, No. 3 ~40° Mmarginata n=|0 | 110° ! Fig. 1. Distribution map of Mentzelia cronquistii and M. marginata. broader filaments for outermost ones; styles 6-10 mm long; capsules cup shaped to sub- cylindrical, 8-12 mm long; seeds lenticular, oblong to somewhat ovate, 2.5—2.7 mm long, 1.7-1.8 mm wide, including the narrow wing, which is 0.2 mm wide; seed surface tan to dark grey, tessellate, strongly colliculate; seed coat cells with the radial walls straight, with center of outer tangential walls raised with 10—20 papillae; chromosome number n = 10; self- incompatible. SPECIMENS EXAMINED: UTAH. Grand Co.: 2 mi E of Thompson, Ripley & Barneby 8659 (CAS, NY); 1 mi W of Cisco, Ripley & Barneby 9209 (CAS); Cisco, Thompson 3518 (LA, chromosome voucher n = 10), Thomp- son 3532 (LA). COLORADO. Mesa Co.: 15 mi W of Fruita, Rollins 1936 (GH); 12.8 mi W of Fruita, Thompson 3519 (LA, chromosome voucher n = 10); 5 mi W of Mack, Brown in 1938 (CS); De Beque, Osterhout 4284 (NY), Osterhout 4724 (GH); near Whitewater, Si- monds in 1948 (CS). Delta Co.: 5 mi NW of Eckert, Weber 7527 (COLO, WS); Paonia, Osterhout 4601 (NY, RM). Montrose Co.: Montrose, Shear 4810 (NY, US). Ouray Co.: near Colona, Payson 2334 (RM). Mentzelia marginata occurs in eastern Utah and western Colorado (Fig. 1) in open habitats in juniper woodland at elevations between 1,400 and 1,800 m. Populations are restricted to grey clay soils often associated with coal. Mentzelia cronquistii Thompson & Prigge, sp. nov. (Fig. 2) July 1986 THOMPSON, PRIGGE: NEW MENTZELIA 10 cm Fig. 2. Mentzelia cronquistii, drawn from Prigge 6643 (LA). 501 SIN NATURALIST Vol. 46, No. 3 Fig. 3. Scanning electron micrographs of seeds of Mentzelia cronquistii (A and B) and M. marginata (C and D). A and C are of whole seed (bar = 1 mm). B and D are of seed coat (bar = 0.05 mm). Similis M. marginata sed in lobis rhachidis _ petalis et staminibus petaloideis nullis, et angustioribus et lobis brevis, floribus cum 10 _ testa cellulis majoribus, parietibus radialium July 1986 undulatus leviter vel bono modo, et plus papillatis differt. Herbaceous perennial 15—40 (—50) cm tall; stems erect, single or much branched at base, reddish colored under the surface layer, pubescent with both glochidiate and pointed hairs; rosette leaves lanceolate to oblance- olate, shallowly and bluntly lobed; lower cauline leaves lanceolate, 2.5-8 cm long, lobed, with the rachis 2-5 mm wide and the lobes 2-10 mm long, thus the leaves 5-15 mm wide; upper leaves shorter and narrower, with the lobes shorter and more pointed; lower leaf surfaces densely pubescent with long and short glochidiate hairs and scattered pointed hairs; upper leaf surfaces more sparsely pubescent with pointed hairs and small glochidiate hairs; leaf margins with large glochidiate hairs; bracts at base of capsules linear and entire; flowering period late May to September; flowers opening in late afternoon; calyx lobes 6-9 mm long; petals 10, yellow, 7-13 (—15) mm long, 2-5 mm _ wide, pubescent on outer surface, narrowly ovate to narrowly obovate, rounded to acute at apex, with inner 5 shorter and narrower than outer 5; stamens numerous, grading in length from 5 mm for the innermost to 9 mm for the outer- most; with narrow filaments for inner ones and wide filaments, to 2.5 mm wide, for the outer ones; styles 6-10 mm long; capsules cup shaped, 6-13 mm long; seeds lenticular, oblong to ovate, 2.4-2.7 mm long, 1.8—2.0 mm wide, including the narrow wing, which is 0.25 mm wide; seed surface tan to dark grey, tessellate, strongly colliculate; seed coat cells with the radial walls slightly to moder- ately wavy and center of outer tangential walls raised with 15-25 papillae; chromosome num- ber n = 10; self-incompatible. Type: UTAH, San Juan Co., about 75 mi W of Blanding and 10 mi E of Hite, May 16, 1961, Cronquist 9030 (Holotype NY!; isotypes LA!, WS!, WTC!, UTC!). Seed from Cron- quist 9030 has been grown in the greenhouse at UCLA, mature plants showed a chromo- some number of n = 10, voucher Thompson 3296 (LA). ADDITIONAL SPECIMENS EXAMINED: More than 100 specimens were examined from the following herbaria: ARIZ, ASU, BRY, CAS, COLO, CS, GH, LA, MO, MICH, NY, POM, RM, RSA, SMU, UC, US, UT, WTU, THOMPSON, PRIGGE: NEW MENTZELIA 503 and WS. A list of these specimens is available on request from LA. Mentzelia cronquistii occurs in southeastern Utah, southwestern Colorado, northwestern New Mexico, and northeastern Arizona (Fig. 1) in open, often disturbed habitats in juniper woodland at elevations between 1,100 and 1,800 m. Populations are most well developed on fine- to medium-grained sand from red sandstone, but populations also occur on rocky talus and in sandy washes. Roadsides are good sites for Mentzelia cronquistii, and the range and abun- dance of the species may have increased with the development of roads. The specific epithet honors Dr. Arthur C. Cronquist, student of angiosperm phylogeny, systematics of the Asteraceae, and floristics of the western United States. Mentzelia cronquistii and M. marginata are more similar to each other than to any other species. They both possess petals that are pubescent on their outer surfaces, a charac- teristic possessed by no other species of Mentzelia. All species of Mentzelia, and prob- ably all species of Loasaceae, have petals with a few hairs at the apex of the petals, but pubescent petals such as occur in M. cron- quistii and M. marginata occur only in those two species and in some South American spe- cies of Loasa and Caiophora of subfamily Loa- soideae. The seeds of M. cronquistii and M. marginata are also very similar, differing only in that the seed coat radial walls of M. cron- quistii are more wavy than those in M. mar- ginata and the seed coat cells in M. marginata are somewhat smaller (Fig. 3), making the seeds of M. marginata appear smoother when viewed with 10X magnification. Mentzelia cronquistii differs in two conspic- uous characters—petal number and _ leaf shape. Mentzelia cronquistii has 10 petals, whereas M. marginata has only 5. In M. cron- quistii the whorl immediately within the petals is composed of stamens with narrow filaments, but in M. marginata the whorls within the petals are petaloid stamens, sta- mens with broad filaments, and then stamens with narrow filaments (Fig. 4). The second conspicuous difference between M. cronquts- tii and M. marginata is in leaf shape. The leaves of M. cronquistii have a narrow rachis with short or long lobes, whereas the leaves of M. marginata are broad with crenate margins. 504 (hs LE (0) Fig. 4. Elements of the floral whorls. A. Mentzelia cron- quistii from left to right: outer whorl petal, inner whorl petal, outermost stamen, stamen - - - innermost stamen, style. B. M. marginata from left to right: petal, petaloid stamen, outer- most stamen, stamen stamen - - - innermost stamen, style. GREAT BASIN NATURALIST Vol. 46, No. 3 We have grown individuals of M. cronquis- tii and M. marginata in the experimental gar- den at UCLA. Voucher specimens of parent plants, siblings, and F, hybrids are deposited at RSA. The data for these crosses are given with the specimens and should be consulted by anyone interested in these species. In sum- mary, these data show that both M. cronquis- tii and M. marginata are self-incompatible. Sibling plants are vigorous and fertile. The hybrids of marginata 2 X cronquistii ¢ and reciprocals are vigorous but sterile, producing less than 10% good pollen and forming no viable seed in both backcrosses and sibling crosses. ACKNOWLEDGMENTS We thank Dr. Stanley L. Welsh for provid- ing the latin diagnosis. NEW VARIETY OF MENTZELIA MULTICAULIS (LOASACEAE) FROM THE BOOK CLIFFS OF UTAH Kaye H. Thorne’ and Frank J. Smith? ABSTRACT. — Described is Mentzelia multicaulis (Osterh.) Goodman var. librina Thorne & F. J. Smith. eto “0 Fig. | Mentzelia multicaulis (Osterh.) Goodman var. librina Thorne & F. J. Smith: A, Branch. B, Seed. C, Habit. Mentzelia multicaulis (Osterh.)Goodmanis northeastern Emery counties in Utah and a suffrutescent perennial endemic to Uintah, western Rio Blanco County, Colorado. The southeastern Duchesne, eastern Carbon, and __ plant is characterized by a diffusely branched lLife Science Museum, Brigham Young University, Provo, Utah 84602. ?26 North 1 East, Smithfield, Utah 84335. 00 508 Recent work by Dr. H. J. Thompson (per- sonal communication) has suggested the pres- ence of a previously undescribed Mentzelia in Utah and eastern Nevada that has been vari- ously treated as M. pumila (Nutt.) T. & G. or as M. laciniata (Rydb.) Darlington. The plants show morphological similarity to the former in capsule shape and flower size, and to the latter in the deeply pinnatifid leaves with narrowly acute lobes. The undescribed material ap- pears to be more nearly allied to M. pumila than to M. laciniata, sharing the same base chromosome number (x = 11, fide H. J. Thompson 3513, Atsatt 646 and 649, Garfield Co., H. J. Thompson 3516, Wayne Co., LA) and habit of growth. The plants differ from M. pumila var. pumila in having basal leaves deeply pinnatifid, with lobes acute to obtuse, upper leaves narrowly lanceolate and pinnati- fid almost to the midvein, bracts remotely pinnatifid with 1 or 2 lobes or entire, seeds winged, gray 3-3.5 mm long, and cell walls of the seed coat undulate. The material is there- fore described as follows. Mentzelia pumila (Nutt.) T. & G. var. la- garosa Thorne var. nov. Similis var. pumila generalis sed in foliis basalium pinnatifidis profunde et lobis acutis vel obtusis foliis cauli- norum lanceolatis anguste fissis propre venis GREAT BASIN NATURALIST Vol. 46, No. 3 bracteis pinnatifidis remotis l-vel 2-lobatis seminibus 3-3.5 mm longis (nec minus 3) et cellula parietis undulatis. Type: USA: Utah. Uintah Co., T11S, R24E, S11, near Watson, Evacuation Creek, 10 miles 173 degrees from Bonanza, 1,708 m, on gravel, 1 August 1980, S. Goodrich & N. D. Atwood 14664 (Holotype BRY; 3 isotypes dis- tributed previously as Mentzelia). ADDITIONAL SPECIMENS: Utah. Duchesne Co., T5S, R3W, S5, West Tavaputs Plateau, in Antelope Canyon, 24 km 67 degrees from Duchesne, 1,769 m, calcareous hills, Uinta Formation, 25 August 1980, S. Goodrich 14972. Kane Co., 14 miles SE of Cannonville along Cottonwood Wash road, pass between Round Valley Draw and Butler Valley, on roadside cut, 1,830 m, 21 June 1983, B. Albee 5064. Piute Co., T30S, RIW, S32, Dry Fork Canyon, ca2km NE of Antimony, 2,288 m, in pinyon-juniper community, on sandy gravel, 28 July 1976, S. L. Welsh, K. Taylor, & G. Moore 14120. Nevada. Lincoln Co., T3N, R70E, S16, ca 2 km on road to Hamlin Valley, Eagle Valley, White Rocks Range, at 1,830 m, grassland—mixed desert shrub community on rocky limestone and clay exposed slopes, 22 August 1979, K. H. Thorne and J. Thorne 775 (all BRY). | | | | i | | AGATHOXYLON LEMONII SP. NOV., FROM THE DAKOTA FORMATION, UTAH William D. Tidwell’ and Gregory F. Thayn? ABSTRACT. —Petrified wood specimens from the Dakota Formation of Utah are here described as Agathoxylon lemonii. This new species is characterized by having distinct growth rings, araucarian tracheary pitting, resin plates or plugs in its tracheids, abundant axial parenchyma, low uniseriate rays and 1—4 small, slightly bordered pits with circular to oval included apertures per crossfield. This is the first report of petrified wood from the Dakota Formation of the western United States and the first record of Agathoxylon from North America. Although fossil plant remains are abundant in the Dakota Formation, reports of petrified plant material from this formation are rare. The study of fossil plants in the Dakota For- mation began in the midwestern United States. Initial collections of fossil plants from this formation were obtained during western territorial surveys for a proposed route for a transcontinental railroad in the 1850s and 1860s (Dilcher et al. 1978). Hayden, in 1853, was the first to obtain leaves from the Dakota Group of Nebraska. He and Meek in 1856-57 collected additional plant materials from these sediments that were subsequently sent to Professor Oswald Heer in Switzerland for study. Heer (1859) published descriptions of these materials. This publication represents the first authentic record of North American Cretaceous plant fossils. Well-preserved leaves were later collected in large numbers in Kansas during the 1860s and 1880s by various workers. These collec- _tions formed the basis for the first major publi- cations on this flora by Lesquereux (1874, 1883, 1892). Subsequent to Lesquereuxs publica- _ tions,very little work had been done on the flora of the Dakota Formation until recently, when considerable research on the reproduc- ‘tive structures and leaves of early an- \ giosperms from this formation in Kansas was | published (Dilcher et al. 1976, Dilcher et al. 1978, Dilcher 1979, Retallack and Dilcher 1981). The Dakota Formation extends throughout much of the southwestern and midwestern United States. Stanton (1905) in working with Jurassic and Cretaceous formations and the Dakota in southern Colorado, New Mexico, and Oklahoma, demonstrated that the Dakota Formation of this region, as originally de- fined, contains both Lower and Upper Creta- ceous strata. In Utah the Dakota Formation occurs between the Lower Cretaceous Cedar Mountain Formation and the Upper Creta- ceous Mancos Shale and is considered to be mid-Cretaceous in age. Fossil plants in the Dakota Formation of western Colorado and eastern Utah were first reported by Brown (1950) and later by Tidwell et al. (1967), Rush- forth and Tidwell (1968), and Rushforth (1970, 1971). Brown (1950) described a flora from the Burro Canyon Formation that is more or less equivalent to at least a portion of the Cedar Mountain Formation (Young 1960, Tschudy et al. 1984) and a flora from the Dakota Forma- tion near Naturita, Colorado. This flora, like the Dakota flora from Westwater, Utah, is atypical in that the percentage of ferns is high as compared with the number of angiosperms. The Dakota flora near Westwater is domi- nated by the ferns Astralopteris Tidwell, Rushforth, and Reveal, Matonidium Schenk, Gleichenia Smith, Hausmania Dunker, As- plenium L., Cladophlebis Brong., and Conio- pteris Brong. (Rushforth 1970, 1971). No coniferous foliage has been reported from this flora in Utah. However, Retallack and Dilcher (1981) noted that Sequoia-like foliage, cones, and cone scales were common conifer megafossil material in the Dakota Formation Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. Bureau of Land Management, Salt Lake City, Utah 84101. 509 560 GREAT BASIN NATURALIST Vol. 46, No. 3 Figs. 1-6. Agathoxylon lemonii: 1, Transverse section illustrating a growth ring and dark axial parenchyma. (33X) (Holotype). 2, Transverse section illustrating an axial parenchyma cell with dark contents and overlapping tracheids (495 X) (Holotype). 3, Radial section illustrating axial parenchyma, radial intertracheary pitting and resin plates (123X) (Holotype). 4, Radial section illustrating uniseriate and biseriate pitting. Note resin plates in lower portion of photograph (495X) (Holotype). 5, Radial section showing a close up of radial pits. Note both elliptic and circular apertures (495X) (Holotype). 6, Radial section illustrating the nature of the ray cell walls and contents. Note crossfield pitting in the right portion of the photograph and showing through dark cell contents in upper right side ray cell (495X) (Holotype). July 1986 Fig. 7-9. Agathoxylon lemonii: 7, Tangential section showing general size, shape, and arrangement of the rays (33X) (Holotype). 8, Tangential section illustrating ray size and shape. Note tangential tracheid pitting. (123X) (Holotype). 9, Tangential section showing outline of ray and ray cells (495X) (Holotype). in Kansas. They also noted the presence of Brachyphyllum, which, although not com- mon, was evidently a minor constituent of the fossil angiospermous swamp woodland flora of this formation. Possible conifer dominance in the inland vegetation of the Dakota flora is supported by a number of palynological stud- ies of this formation and rock units of equiva- lent age in North America (Pierce 1961, Agasie 1969, Romans 1975, Retallack and Dilcher 1981). TIDWELL, THAYN: FossiL PLANT SYSTEMATICS Coniferales Agathoxylon Hartig Agathoxylon lemonii sp. nov. Figs. 1-9 D1aGNosis.—Growth rings distinct, 2-6 mm wide; transition from early to late wood gradual; tracheids generally hexagonal in transverse section, some compressed and oval in outline, approximately 50 pm in diameter, some slightly elongated radially, walls approx- imately 10 um thick, tips of adjacent tracheids appearing in cross section as intercellular spaces often lending an appearance of col- lenchyma to the xylem; radial pitting gener- ally uniseriate, frequently biseriate, pits slightly appressed, circular to horizontally elongate, 12-14 wm diameter with 2.4—-7.8 wm apertures; tangential pitting uniseriate with pits same size as radial pitting, generally isolated; resin plates or plugs observed in many tracheids; axial parenchyma abundant, diffuse, with smooth thick walls and dark in- tercellular contents; rays uniseriate, homocel- lular, approximately six per millimeter, 20-70 wm wide, 10 (commonly 5 or 6) cells high, 30-150 ym high; ray cells vertically flattened and elongate horizontally, nearly twice as wide tangentially as high, often 50 ym wide tangentially by 30 pm in vertical dimension and 30-100 pm (average 70 fm) in radial di- mension; ray cells filled with dark cell con- tents, ray cell walls approximately 3 ym thick 562 and unpitted except at crossfield; crossfield pits commonly obscured by cell contents, ob- servable crossfields with 1—4, small (approxi- mately 5 wm), slightly bordered pits with cir- cular to oval included apertures. ReEposiTory: Brigham Young University, 5029 (Holotype). LOCALITIES: 7 mi (11.2 km) east of Ferron, Utah; U.S. Geol. Surv. Map: Desert Lake Quadrangle, SE 1/4, Sec 26, T20S, R8E (Holotype). Other specimen: 2.5 mi (4.02 km) north of Westwater, Utah. Horizon: Dakota Formation. AGE: Lower Upper Cretaceous (Cenoma- nian). EtyMOLoGy: The specific epithet is for Mr. and Mrs. Frank Lemon of Moab, Utah, who donated specimens for this study. DISCUSSION The presence of araucarian tracheary pit- ting and of resin cysts, plugs, plates or spools in Agathoxylon lemonii indicate that it is allied to the Araucariaceae. The Araucariaceae con- sist of two extant genera, Araucaria De Jussieu with 18 species (De Laubenfels 1972) and Agathis Salisbury composed of 13 species (Whitmore 1980). Fossil woods with araucar- ian structure are abundant in the geologic record. They range in age from Carboniferous to Recent and have been assigned to many genera, including Araucarites Presl, Pinites Lindl. & Hutton sensu Presl, Dadoxylon Endlicher, Araucarioxylon Kraus, Agathoxy- lon Hartig, and Araucariopitys Jeffrey. These woods may also be related to Cordaioxylon Gr. Eury, Cordaites Unger, or early conifers such as Lebachia Florin or Ernestodendron Florin. Knowlton (1899), Penhallow (1907), Stopes (1914), Holden (1914), and Jeffrey (1926) assigned Paleozoic araucarian woods to Dadoxylon and reserved the name Araucari- oxylon for Mesozoic and Tertiary materials. Further, Seward (1919) proposed adding qualifying terms such as Araucarioxylon or Cordaioxylon in parenthesis after Dadoxylon whenever evidence supports such designa- tion. Torrey (1923) agreed somewhat with us- ing Araucarioxylon for Mesozoic and Ceno- zoic woods and Dadoxylon for Late Paleozoic types. However, he also separated them by assigning Dadoxylon to fossil woods lacking GREAT BASIN NATURALIST Vol. 46, No. 3 wood (axillary) parenchyma and assigning woods with parenchyma to Araucarioxylon. Edwards (1921) proposed using Dadoxylon for araucariaceous wood that could not be related to either Agathis or Araucaria. Some authors, such as Krausel and Jain (1963), Sah and Jain (1963), and Vogellehner (1964) followed Gothan’s (1905) recommendation that all woods having an anatomical similarity to Araucarineae or Cordaiteae should be placed into Dadoxylon as a single nomenclatural unit. Krausel (1949) designated Araucarioxy- lon as the genus for all woods related to Agathis or Araucaria. Hartig (1848) instituted the genus Agathoxylon for fossil having “zellfasern” or axial parenchyma woods simi- lar to the living genus Agathis. The presence of axial or wood parenchyma is the only char- acter absent or rarely found in Araucaria. Krausel and Jain (1963), in discussing Agath- oxylon remarked that Hartig had not given any details of his specimen. Sah and Jain (1963) went further and suggested that the “zellfasern” of the Hartig specimen were per- haps resinous tracheids. According to Seward (1919), resin plates in tracheids are often in- terpreted as end walls of axial parenchyma, thus leading to erroneous reports of Agath- oxylon. Greguss (1955), Jane (1970), and Stockey (1982) considered the anatomical sep- aration between Agathis and Araucaria on the basis of wood anatomy to be not only difficult but nearly impossible. Greguss (1967) later reversed himself by pointing out that, of the living species in the Araucariaceae, only those in the genus Agathis contain true axial par- enchyma, and subsequently he assigned two species to Agathoxylon. The fact that the Da- kota wood under study bears axial par- enchyma places it in Agathoxylon. Members of the genus Araucariopitys bear short shoots, have resin ducts and abietinous crossfield pitting, and are, therefore, not re- lated to this wood from the Dakota Forma- tion. COMPARISON Of the living species of Agathis, Agathoxy- lon lemonii bears striking resemblance to the wood of Agathis hypoleuca. It differs from A. hypoleuca in having higher rays (1-10 cells high as opposed to 1—4 cells in A. hypoleuca), | July 1986 fewer pits per crossfield (1-4 in A. lemonii, 4-8 in A. hypoleuca), and more abundant axial par- enchyma. Dadoxylon septentrionale Gothan (1905) dif- fers from A. lemonii in having typically 2—4 ellip- tical to oblique crossfield pits and lacking tan- gential pitting. Dadoxylon eocenum Chitaley (1949) and Dadoxylon (Araucarioxylon) japonicum Shimakura (1936) have tangential pit- ting, but both lack the xylem parenchyma of A. lemonii. Dadoxylon agathoides Krausel & Jain from the Jurassic of India is similar to the Dakota specimen. However, it lacks axial parenchyma, has higher, narrower rays and has crossfield pits that are arranged in clusters. Araucarioxylon texense Torrey has axial parenchyma and pitting similar to the speci- men from the Dakota Formation, although A. texense has narrower rays and short shoots, which A. lemonii lacks. Araucarioxylon hop- pertonae Knowlton from the Cretaceous of the Black Hills is similar to A. lemonii in hav- ing few pits per crossfield and low rays, but it lacks the characteristic axial parenchyma. Holden (1914) mentioned an araucarian type wood from the Cretaceous of New Jersey (Rar- itan Formation) whose pith contains large masses of stone cells similar to those in living Agathis but lacks wood parenchyma. Dadoxy- lon noveboracense (Holl. & Jeff.) from the mid-Cretaceous beds of Staten Island lacks definite growth rings and has uniseriate tra- cheary pitting. These characters are similar to A. wyomingense Andrews and Pannell (1942) from Cretaceous strata of Gros Ventre Canyon in Wyoming. Araucarioxylon wyomingense lacks the wood parenchyma and tangential pitting that separates this species from A. lemonii. Among the species belonging to this genus described from the Mesozoic of Africa, the following four possess wood parenchyma (Gazeau 1969): D. (A.) aegyptiacum Unger (1859), D. (A.) paumieri Loubiere (1935), D. (A.) septatum Boureau (1951) from the Sahara Soudanais, and D. (A.) koufraense Batton (1965) from the continental series of Libya. They differ from A. lemonii, in general, by having different ray height, different tra- cheary pitting, and, with some, having septa- tions in their tracheids. Dadoxylon alpinum Lemoigne (1966) from the Jurassic of the Bassin de la Durance has TIDWELL, THAYN: FOSSIL PLANT 563 wood parenchyma, but it differs from A. lemonii in possessing septate tracheids, hav- ing large crossfield pits with large lumens, and lacking tangential pitting. Dadoxylon (Arau- carioxylon) breveradiatum (Lignier) Seward from the Cenomanian of Normandy has abun- dant resiniferous parenchyma, but it differs from A. lemonii by having higher rays (4—80, usually 10—40, cells high) and septate tra- cheids. Dadoxylon (Araucarioxylon) novaezee- landii (Stopes) Seward from the Cretaceous of New Zealand has well-marked growth rings, resin plates, and araucarian pitting. However, it differs from the Dakota Formation species in its lack of wood parenchyma and having thick-walled tracheids on each side of the rays. There are a few reported species of Dadoxy- lon or Araucarioxylon from Japan that are similar to Agathoxylon lemonii. Dadoxylon (Araucarioxylon) sidugawaense Shimakura (1936) from the Jurassic of Miyagi Prefecture is similar in possessing distinct growth rings, similar radial pitting, tangential pitting, and, in the presence of xylem, parenchyma. But D. (A). sidugawaense differs from the former species in having circular, alternate tangential pits and simple crossfield pits, whereas the tangential pits of A. lemonii are the same size as the pits on the radial walls. These tangential pits are isolated rather than contiguous, and the crossfield pits of the latter species are slightly bordered rather than simple. Two Lower Cretaceous species from Japan, Arau- carioxylon hujinamiense Ogura (1960) from Wakayama and Chiba and A. pseudo-huji- namiense Nishida and Oishi (1982) from the Kwanto Mountain, both differ from A. lemonii in possessing tylosislike structures in their tra- cheids and in lacking wood parenchyma. Araucarioxylon nihongii Nishida and Nishida (1984) is quite similar to this Dakota Forma- tion species. The Japanese species, however, has 3-5 rows of pits on its tracheid walls, has shorter parenchyma cells, and lacks tangential pitting. Agathoxylon species similar to Agathoxylon lemonii include A. australe Evans, A. hun- garicum (Andreanszky) Greguss, and A. mec- sekense Greguss. Agathoxylon australe is a Pliocene fossil from New Zealand that has vestured pits and lacks axial parenchyma. 564 Evans (1934) originally assigned this species to Agathis and later renamed it Agathoxylon australe (Evans 1937). Krausel and Jain (1963) noted that Agathoxylon australe is very simi- lar to living Agathis australis. Agathoxylon mecsekense from the Jurassic of Hungary has high, narrow rays and commonly has triseriate pitting. Although the description given for A. hungaricum is incomplete, it appears to have higher rays and more pits per crossfield than our specimen. PALEOECOLOGY The extant genus Agathis may be found growing in association with the fern Matonia in the tropical uplands of the Malay Peninsula and the island of Borneo (Seward 1899). Mor- phologically, the fossil genus Matonidium is similar to the extant Matonia (Berry 1919). Matonidium occurs in abundance in the Da- kota Formation near Westwater, Utah, a lo- cality quite close to a collecting site for Agath- oxylon lemonii. Mahabale (1954) considered matoniaceous ferns to be among a select group of ferns that served as reliable indicators of subtropical to tropical paleoclimate; the pres- ence of abundant Matonidium is, therefore, suggestive of a similarly warm, climatic regime during the deposition of this formation in this area. This reconstruction is supported by Roman's (1972) study of the schizeaceous spores of the Dakota Formation and other reconstructions that have emphasized the subtropical to tropical nature of the regional fossil assemblage (Rushforth 1971). The precise community assignment for A. lemonii is more elusive. Agasie (1969), Rush- forth (1971), and May and Traverse (1973) all suggested that the Dakota Formation was de- posited on a wet, low-lying landscape in close proximity to drier uplands. The Matonidium leaf material occurs in an ash layer associated with a coal bed at the Westwater site. The swamps of the Dakota Formation, which pro- duced the peat that is the original source of the coals of the region, apparently supported a diverse tree and shrub flora in which ferns were probable understory elements (Retal- lack and Dilcher 1981). In such situations, however, microtopographic variability can re- sult in a complex vegetation mosaic with somewhat drier sites, possibly dominated by GREAT BASIN NATURALIST Vol. 46, No. 3 angiosperms, in close spatial assocation with the actual swamp vegetation. The coarser clastics of modern distributary channels and coastal lagoons usually include a mixed assem- blage derived primarily from the lower coastal plain but also including wood that may have been transported for greater distances than is typically the case with the angiosperm leaves or conifer needles in such deposits. The coarser sandstone in which the specimens of Agathoxylon lemonii were collected suggests the possibility that they were transported from a well-drained habitat occurring some- where along the lower delta plain associated with the deposition of the Dakota Formation. ACKNOWLEDGMENTS The authors wish to express their apprecia- tion to S. R. Rushforth of Brigham Young University, Provo, Utah; S. R. Ash of Weber State College, Ogden, Utah; Greg Retallack of the Univesity of Oregon, Eugene, Oregon; and R. E. Taggart of Michigan State Univer- sity, East Lansing, Michigan, for their review of the manuscript. LITERATURE CITED AGASIE, J. M. 1969. 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Zur nomenklatur der fossilen Holzgattung Dadoxylon Endlicher 1847. Taxon 13: 233-237. WHITMORE, T. C. 1980. A monograph of Agathis. Plant Syst. Evol. 135: 41-69. YOUNG, R. G. 1960. Dakota Group of Colorado Plateau. Amer. Assoc. Pet. Geol. Bull. 44(1): 156-194. GRAZING AND PASSERINE BREEDING BIRDS IN A GREAT BASIN LOW-SHRUB DESERT Dean E. Medin! ABSTRACT. — Densities of passerine breeding birds were compared between four range pastures variously grazed by sheep over a 50-year period. The experimental pastures, located at the Desert Experimental Range in southwestern Utah, included three grazed and one ungrazed. Grazed pastures were each heavily stocked and grazed annually at one of three winter seasons (early, middle, or late). Important structural (physiognomic) and compositional differences existed in the vegetation of the experimental pastures. Horned Larks (Eremophila alpestris |Linnaeus]), numerically dominant in the pastures, apparently responded to those differences. Black-throated Sparrows (Amphispiza bilineata [Cassin]) and Loggerhead Shrikes (Lanius ludovicianus Linnaeus) were less common and found as breeding birds only in dry wash habitats. No significant differences were found between the pastures in estimates of total breeding bird populations, bird standing crop biomass, or bird species richness. The low-shrub desert is a mosaic of plant communities found throughout the plains, foothills, and valleys of the Great Basin. In some areas, perennial grasses share domi- nance with the low shrubs. The desert has been variously classified as the shadscale asso- ciation, the northern desert shrub formation, or the salt-desert shrub association (Holm- gren 1973). Many of the native plants are palatable and nutritious, and the deserts are used primarily for winter grazing by domestic sheep (Holmgren and Hutchings 1972). Physical environments of low-shrub deserts are rigorous. Climatically, they are cold deserts: winters are cold and summers are warm. Annual precipitation is generally low and occurs sporadically. Long-term irregular- ities in climatic patterns exist. The vegetation is structurally and floristically uncomplicated. Production rates are relatively low (Blaisdell and Holmgren 1984). These features of low- shrub desert environments and vegetation re- strict avifaunal development to small species assemblages of broadly distributed forms (Wiens and Dyer 1975). Vegetation structure is an important factor determining habitat selection in birds (re- viewed in Hildén 1965). As a result, habitat structure may affect the organization of the avian community as a whole (Willson 1974, Roth 1976, Rotenberry and Wiens 1980). Be- cause grazing can alter the structure of low- shrub desert vegetation (Hutchings and Stew- art 1953, Holmgren and Hutchings 1972), dif- ferent grazing treatments may have important consequences for avifaunal composition and abundance. The objective of this study was to compare breeding bird populations and community or- ganization between selected pastures vari- ously grazed by sheep for 50 years. The study was restricted to passerine bird species. DESERT EXPERIMENTAL RANGE The study was conducted at the Desert Ex- perimental Range in southwestern Millard County, Utah. It was established in 1933. Its 225 km’ are representative of about 160,000 km’ of winter grazing lands in the Great Basin physiographic province as well as adjacent parts of the Columbia and Colorado plateaus (Blaisdell and Holmgren 1984). About 75% of the Experimental Range is alluvial slope or flat valley bottom. The rest is steeper upland overlain by a shallow soil mantle and broken by ledges of hard Paleozoic sedimentary or Tertiary volcanic rock. There are no seeps, springs, or live streams. Numerous dry washes cross the alluvial fans and may flow for short periods following high-intensity sum- mer showers. Elevation ranges from 1,547 to 2.565 m. Soil textures are typically loams, sandy loams, or loamy sands (Holmgren 1973). 1termountain Research Station, Forest Service, U.S. Department of Agriculture, Boise, Idaho 83702. 567 568 During the 50 years from 1934 to 1984, temperature extremes varied from —40 to 40 C. The ground is frozen most of the time from mid-November into March. Snowfalls are usually light, seldom more than 5 cm deep. The average annual precipitation is 15.7 cm, about half of which falls during the five months from May through September (Holm- gren 1973). The vegetation on the Experimental Range is a mixture of low shrub and shrub-grass types. The dominant shrub species are win- terfat (Ceratoides lanata [Pursh] J. T. How- ell), bud sagebrush (Artemisia spinescens D. C. Eaton in Wats.), and shadscale (Atriplex confertifolia |[Torr. & Frem.] Wats.). Three perennial grasses—Indian ricegrass (Oryzop- sis hymenoides [R. & S.] Ricker in Piper), galleta (Hilaria jamesii [Torr.] Benth.), and sand dropseed (Sporobolus cryptandrus [Torr. | Gray)—are associated with shrubs on most soils. METHODS Grazing studies began on the Desert Ex- perimental Range during the winter of 1934-1935 (Hutchings and Stewart 1953). Sheep grazed 20 large (129 ha) range pastures at one of three intensities (light, moderate, or heavy) and one of three winter seasons (early, middle, or late). Grazing assignments for each experimental pasture remained unchanged. The winter of 1983-1984 marked 50 consecu- tive years of grazing in the pastures. The rest of the area was divided into 14 units. Over the years, 11 have been grazed by sheep and 2 by cattle. One unit had not been grazed. Three heavily grazed pastures, one each grazed in early, middle, or late winter, and part of the ungrazed unit (hereafter referred to as the ungrazed pasture) were selected for study. The experimental pastures were chosen for maximal contrasts in seasonal grazing assignments and minimal differences in soil-site characteristics. Grazed pastures were contiguous, and all of the pastures selected for study were covered with similar plant types when the experiments started in 1934 (Hutchings and Stewart 1953). To avoid dissimilarities in soil and site characteris- tics, only the northern half of the late-winter grazed pasture and the eastern half of the un- grazed pasture were studied. GREAT BASIN NATURALIST Vol. 46, No. 3 Four 9-ha plots were randomly located in each of the four selected experimental pas- tures and censused for breeding birds using the Williams spot-map method (International Bird Census Committee 1970). The square plots were surveyed and gridded in a Carte- sian coordinate system with points flagged and numbered with stakes at 50-m intervals. Seven census visits were made to each plot from 5 April to 1 June 1984. Most of the work was done from sunrise to late morning. To ensure complete coverage, a plot was cen- sused by walking within 25 m of all points on the grid. Census routes through each plot were varied. Observations extended well be- yond plot boundaries. At the end of the sampling period, concen- trated groups of observations and coded activ- ity patterns were circled as indicating areas of activity or approximate home ranges. Frac- tional parts of boundary territories were rec- ognized. Results were converted to the num- ber of pairs of breeding birds per 40 ha. Species richness (N) was expressed as the total number of breeding bird species observed or. a plot. Vegetation and other features of experi- mental pastures were measured from 11 June to 2 August 1984. A m° quadrat was located at each of 49 equally spaced stakes that defined the coordinates of the 9-ha bird census grids. Canopy coverage (Daubenmire 1959) was oc- ularly estimated for each plant species and recorded as the midpoint of one of eight per- cent coverage classes (0-1, 1-5, 5-10, 10-25, 25-50, 50-75, 75-95, and 95-100). Percentages of litter, rock, and bare ground were similarly estimated. Distances to the nearest grass plant, forb, and shrub in each quadrant were measured from the center of each quadrat. The patchiness of the vegetation was esti- mated using Roth's (1976) index of hetero- geneity (D) calculated as D = 100 SD/x where SD is the standard deviation and x is the mean of the point-to-plant distances. The height of each grass plant, forb, and shrub nearest the center of each quadrat was mea- sured with a pocket tape. Vegetation data and attributes of avian com- munity organization between experimental pastures were compared using analysis of vari- July 1986 MEDIN: GREAT BASIN BIRDS 569 TABLE 1. Vegetation and other features of grazed and ungrazed pastures, Desert Experimental Range, Utah, 1984. ae ee Feature Heavy, early winter Ground cover (%) Bare ground 72.82 a Litter 2.36 ab! Rock” 4.26 a Grass Aristida purpurea 0.28 ab Bromus tectorum 0.47 a Hilaria jamesii 2.15 ab Oryzopsis hymenoides 2.09 a Sitanion hystrix 0.17 a Sporobolus contractus 0.04 a Sporabolus cryptandrus 6.51 a Others 0.02 a Totals 78 a Forb Chaenactis macrantha 0.02 a Descurainia pinnata 0.01 a Halogeton glomeratus 0.03 a Lappula occidentalis 0.39 a Lepidium montanum 0.20 a Machaeranthera canescens 0.06 a Phacelia corrugata 0.01 a Salsola iberica 0.02 a Sphaeralcea grossulariifolia 0.59 a Townsendia florifer 0.04 a Others 0.10 a Totals 1.47 a Shrub Artemisia spinescens 2.32 ab Atriplex canescens 0.09 a Atriplex confertifolia 3.94 ab Ceratoides lanata 2.49 a Chrysothamnus viscidiflorus 0.06 a Xanthocephalum sarothrae 0.97 a Others 0.50 a Totals 10.37 a Vegetation height (m) Grass 0.16 a Forb 0.05 a Shrub O.ll a Patchiness index (%)* Grass 107 a Forb 109 + ab Shrub 94° a Experimental pasture SEA SE ee a ae ed Heavy, Heavy, Ungrazed middle winter late winter 74.30 a 74.26 a 68.11 a WAIL Bh 3.23 ab 4.49 b 3.62 a 2.08 a 5.01 a 0.02 a 0.02 a 0.93 b 10l a Omsma 0.52 a 0.42 b 0.95 b 3.66 a 4.43 ab 10.19 ¢ 4.82 b 0.13 a 0.02 b 0.64 c 0.58 ab 1.65 b 0.00 a 5.25 ab 2.41 b 0.22 c 0.00 a 0.00 a 0.21 a 11.84 a 15.97 a 11.00 a 0.05 a 0.00 a 0.05 a 0.00 a 0.00 a 0.10 a 0.07 ab 0.25 b 0.29 ab 0.40 a 0.15 a 0.55 a 0.45 b 0.02 c 0.06 c 0.06 a 0.04 a 0.63 b 0.07 ab 0.00 a 0.18 b 0.03 a 1.54 b 0.10 a 0.19 b 0.23 b 1.10 ¢ 0.06 a 0.01 a 0.01 a 0.33 a 0.04 a 0.19 a 1.7la 2.28 ab 3.26 b 15224 0.02 c 2.53 b 0.00 a 0.00 a 0.21 a 6.05 b 1.68 c 2.98 ac 0.84 b 1.10 ab 2.95 a 0.09 a 0.02 a 3.82 b 1.38 a 0.28 b 0.03 b 0.51 a 0.23 a 0.07 a 10.09 a 3.33 b 12.59 a 0.18 ab 0.23 ¢ 0.21 be 0.04 a 0.04 a 0.12 b 0.12 a 0.12 a 0.15 a 107 a 116 a 120 a 108 ab 122 b 89 a 102 a 114 a 120 a 'Dissimilar letters in rows denote differences (P < 0.10) in means between pastures. Exposed bedrock and rock particles on the surface of the ground greater than 2.5 cm in diameter. 3Roth’s (1976) index of heterogeneity. ance for a 1-factor design. Multiple compari- sons of means followed Gabriel (1978). Arcsin transformation was used for percentage data. Tests of significance were at P < 0.10. Authorities for plant names are from Welsh et al. (1981). Bird nomenclature is from the 1983 AOU Check-list (American Ornitholo- gists Union 1983). RESULTS Vegetation Structural and compositional differences existed in the vegetation of the experimental pastures (Table 1). The pasture grazed in late winter each year differed from the others mainly in its reduced shrub ground cover, 570 GREAT BASIN NATURALIST Vol. 46, No. 3 TABLE 2. Mean density (pairs/40 ha), standing crop biomass, and other attributes of passerine birds breeding on grazed and ungrazed pastures, Desert Experimental Range, Species Foraging Nesting weight” Species category' substrate” (g) Horned Lark GGO G 31.3 (Eremophila alpestris ) Black-throated Sparrow EEO B 14.0 (Amphispiza bilineata) Loggerhead Shrike GFC B 48.1 (Lanius ludovicianus ) Total pairs/40 ha Total individuals/km? Standing crop biomass (g/ha) Species richness (N) 1GGO = ground-gleaning omnivore, GFC = ground-feeding carnivore. 2G = ground nester, B = bush nester. Utah, 1984. Experimental pasture Heavy, Heavy, Heavy, early winter middle winter late winter Ungrazed 19.7 a’ 18.2 ab 14:9) beaeleseab 18) 9.6 a 4.3 a 46a +° + 0.4 a 0.8 a 27hGea PHS) & 19.6 a 239200 137.8 a 139.0 a 93 Avaeeellorona! 36.3 a ST By” 27.4 a 32.9 a 2.0 a 2.0 a 7 a lia Species weights from Schoener (1968), Rotenberry (1980), and Wiens and Rotenberry (1980). ‘Dissimilar letters in rows denote differences (P < 9.10) in means between pastures. °+ indicates bird observed infrequently. which was less than a third that of the other pastures. The reduced shrub cover and the slightly larger percentage of grass cover gave the late-winter grazed pasture a grassy aspect. Also, the average grass height was higher than those pastures grazed earlier in the winter. Bud sagebrush, common in the other pas- tures, was a minor part of the plant cover in the late-winter grazed pasture. Indian rice- grass dominated the grass component, and Russian thistle (Salsola iberica Sennen & Pau) was most abundant among the forbs. Few features of the ungrazed pasture dif- fered from those pastures that were grazed. The ungrazed area had slightly less bare ground and slightly more rock cover than grazed pastures, but neither component was significantly different from grazed pastures. There was significantly more litter cover in the ungrazed pasture when compared to the pasture grazed in the middle of the winter. Forbs made up a larger percentage of the ground cover and were taller in the ungrazed pasture. Of the more common perennials, squirreltail (Sitanion hystrix [Nutt.] J. G. Sm.), galleta, and globemallow (Sphaeralcea grossulariifolia [H. & A.] Rydb.) made up a larger percentage of the plant cover on un- grazed sites. Sand dropseed, common in grazed pastures, was a minor component in the ungrazed pasture. Among the shrubs, only low rabbitbrush (Chrysothamnus viscidi- florus [Hook.| Nutt.) was more abundant in the ungrazed pasture. The pastures grazed in the early- and mid- dle-winter periods were comparable in nearly all the features measured. Only winterfat and globemallow had larger cover values in the early-grazed pasture; pepperweed (Lepidium montanum Nutt. in T. & G.) was more impor- tant in the pasture grazed in midwinter. Birds Only three passerine breeding bird species occurred in the experimental pastures (Table 2). The most common of these was the Horned Lark (Eremophila alpestris [Linnaeus]). A permanent resident, this broadly distributed bird was found throughout the variously grazed pastures. Less common, and found in more restricted habitats, were two summer residents, the Black-throated Sparrow (Am- phispiza bilineata [Cassin]) and the Logger- head Shrike (Lanius ludovicianus Linnaeus). Wide-ranging raptorial birds of prey, al- though commonly seen, were not included in the analysis. Large numbers of transient spe- cies were also excluded. Mean breeding bird density ranged from 19.6 to 27.8 pairs/40 ha in the experimental pastures (Table 2). From 65% to 76% of the average total bird density in each pasture was accounted for by the Horned Lark. Black- throated Sparrow and Loggerhead Shrike numbers were highly variable. Among the individual species, only the Horned Lark dif- fered significantly in density; the highest av- erage density was in the early-winter grazed | July 1986 pasture and the lowest in the late-winter grazed pasture. Although not significant, there was a tendency toward intermediate bird density and bird biomass values in the ungrazed pasture. Species richness means were slightly higher in the early- and middle-winter grazed pastures. No significant differences were found in esti- mates of total bird numbers, bird standing crop biomass, or bird species richness between the various pastures. DISCUSSION Horned Larks were distributed throughout the experimental pastures. Few locations within the 9-ha bird census plots were not included within the territorial boundaries of a nesting pair of Horned Larks. Breeding terri- tories were contiguous and frequently over- lapped. Densities, however, differed be- tween pastures. Horned Lark density was highest in the early-winter grazed pasture and lowest in the late-winter grazed pasture (Table 2). Important differences occurred in the floristics and the physiognomy of those two pastures (Table 1). The late-grazed pas- ture had the lowest shrub cover, the highest grass cover, and the tallest average grass height. Grasses, mostly Indian ricegrass, dominated the aspect. The early-grazed pas- ture had the lowest forb cover, the lowest average grass and shrub heights, and gener- ally lower grass and shrub patchiness indexes. The visual impression was that of an open, low-growing, mixed grass-shrub habitat. Black-throated Sparrows and Loggerhead Shrikes were locally distributed within the experimental pastures. As breeding birds, they were largely restricted to the dry washes that cross the alluvial fans and bajadas of the Desert Experimental Range. Dry washes were generally only a few meters wide, and they contained taller shrubs, mostly desert peachbrush (Prunus fasciculata [Torr. | Gray), interspersed among shorter vegetation. Den- sities of Black-throated Sparrows and Logger- head Shrikes within the larger boundaries of an experimental pasture (Table 2) were appar- ently a function of the size and linear extent of the dry washes and the taller vegetation they contained. The distribution of breeding birds within the experimental pastures is perhaps best ex- MEDIN: GREAT BASIN BIRDS 571 plained by their different nesting require- ments. Other investigators, working in a vari- ety of habitats and locations, have noted the apparent preference of Horned Larks for open and low-growing vegetation as nesting sites (Fautin 1946, Wiens 1973, Owens and Myres 1973, Krementz and Sauer 1982, Castrale 1982). In this study, Horned Lark nests were found only in open habitats and always on the ground. Nests were placed in a shallow exca- vation partly beneath or beside a low shrub or grass tussock. Fautin (1946) first noted that the Black- throated Sparrow seems to prefer an open type of vegetation within which there are oc- casional larger shrubs. Raitt and Maze (1968) found Black-throated Sparrows and Logger- head Shrikes nesting exclusively in dry wash (arroyo) habitats. Similarly, I found Black- throated Sparrows and Loggerhead Shrikes nesting above the ground in the shrubs of the dry washes. Black-throated Sparrow nests were placed near the ground (<1 m) in small to medium-sized shrubs. Loggerhead Shrike nests were placed higher (>1 m) and in larger, more thickly foliaged shrubs. Taller shrubs in the dry washes were fre- quently used as perches by each of the breed- ing bird species. Horned Larks sang from the ground, while perched, or from the air during nuptial flight displays. Black-throated Spar- rows sang from elevated perches and some- times foraged in the foliage of shrubs; they occasionally hawked insects from exposed perches. Loggerhead Shrikes used tall shrubs as observation posts. Both the Loggerhead Shrike and the Black-throated Sparrow were often seen coursing up and down the dry washes. Agonistic encounters between the two were occasionally seen in dry wash habi- tats. The Loggerhead Shrike was the aggres- sor in those encounters. Several workers (e.g., Cody 1968, Wiens 1969, Rotenberry and Wiens 1980) have shown that the physical structure of the habi- tat can affect relationshps between grassland and shrubsteppe birds. In this study, signifi- cant structural differences were found in the vegetation of range pastures variously grazed by sheep for 50 years. One passerine breeding bird species, the Horned Lark, apparently responded to those structural differences. Other passerines, the Black-throated Sparrow 572 and the Loggerhead Shrike, nested only in the taller vegetation that occurred in dry wash habitats. On the other hand, no significant differences were found in bird community at- tributes between the experimental pastures. The results of this study suggest that winter grazing by sheep, to the degree that it can alter the structure of low-shrub desert vegeta- tion, has the potential to alter breeding bird populations and their distribution. ACKNOWLEDGMENTS I gratefully acknowledge the field assis- tance of G. Medlyn and J. Martin. J. Kinney, R. C. Holmgren, and W. P. Clary of the Shrub Sciences Laboratory at Provo, Utah, provided support and contributed in many other ways. J. Verner made constructive com- ments on an earlier draft of this manuscript. LITERATURE CITED AMERICAN ORNITHOLOGISTS UNION. 1983. Check-list of North American birds. 6th ed. American Ornithologists’ Union, Washington, D.C. 877 pp. BLAISDELL, J. P., AND R.C. HOLMGREN. 1984. Managing inter- mountain rangelands—salt-desert shrub ranges. USDA For. Serv. Gen. Tech. Rep. INT-163. Inter- mountain For. and Range Expt. Sta., Ogden, Utah. CASTRALE, J. S. 1982. Effects of two sagebrush control meth- ods on nongame birds. J. Wildl. Manage. 46: 945-952. Copy, M. L. 1968. On the methods of resource division in grassland bird communities. Amer. Nat. 102: 107-147. DAUBENMIRE, R. 1959. A canopy-coverage method of vegeta- tional analysis. Northwest Sci. 33: 43-64. FautTin, R. W. 1946. Biotic communities of the northern desert shrub biome in western Utah. Ecol. Monogr. 16: 251-310. GABRIEL, K. R. 1978. A simple method of multiple compari- sons of means. J. Amer. Stat. Assoc. 73: 724-729. HILDEN, O. 1965. Habitat selection in birds. Ann. Zool. Fenn. 2: 53-75. HOLMGREN, R. C. 1973. The Desert Experimental Range: de- scription, history, and program. Pages 18—22 in Arid shrublands—proceedings of the third workshop of the U.S./Australia rangelands panel. Soc. Range Man- age., Denver, Colo. GREAT BASIN NATURALIST Vol. 46, No. 3 HOLMGREN, R. C., AND S. S. HUTCHINGS. 1972. Salt desert shrub response to grazing use. Pages 153-165 in Wildland shrubs—their biology and utilization. USDA For. Serv. Gen. Tech. Rep. INT-1. Inter- mountain For. and Range Expt. Sta., Ogden, Utah. HUTCHINGS, S. S., AND G. STEWART. 1953. Increasing for- age yields and sheep production on intermountain winter ranges. USDA Circ. 925. INTERNATIONAL BIRD CENSUS COMMITTEE. 1970. An inter- national standard for a mapping method in bird census work. Audubon Field Notes 24: 722-726. KREMENTZ, D. G., AND J. R. SAUER. 1982. Avian communi- ties on partially reclaimed mine spoils in south central Wyoming. J. Wildl. Manage. 46: 761-765. OWENS, R. A., AND M. T. Myres. 1973. Effects of agricul- ture upon populations of native passerine birds of an Alberta fescue grassland. Can. J. Zool. 51: 697-713. Ratt, R. J., AND R. L. Maze. 1968. Densities and species composition of breeding birds of a creosotebush community in southern New Mexico. Condor 70: 193-205. ROTENBERRY, J. T. 1980. Dietary relationships among shrubsteppe passerine birds: competition or op- portunism in a variable environment? Ecol. Monogr. 50: 93-110. ROTENBERRY, J. T., AND J. A. WIENS. 1980. Habitat struc- ture, patchiness, and avian communities in North American steppe vegetation: a multivariate analy- sis. Ecology 61: 1228-1250. Rotu, R. R. 1976. Spatial heterogeneity and bird species diversity. Ecology 57: 773-782. SCHOENER, T. W. 1968. Sizes of feeding territories among birds. Ecology 49: 123-141. WELSH, S. L., N. D. ATwoop, S. GooprRIcu, E. NEESE, K. H. THORNE, AND B. ALBEE. 1981. Preliminary in- dex of Utah vascular plant names. Great Basin Nat. 41: 1-108. WIENS, J. A. 1969. An approach to the study of ecological relationships among grassland birds. Ornithol. Monogr. 8: 1-93. ____.. 1973. Pattern and process in grassland bird com- munities. Ecol. Monogr. 43: 237-270. WIENS, J. A.. AND M.I. Dyer. 1975. Rangeland avifaunas: their composition, energetics, and role in the ecosystem. Pages 146-182 in Proceedings of the symposium on management of forest and range habitats for nongame birds. USDA For. Serv. Gen. Tech. Rep. WO0-1. WIENS, J. A., AND J. T. ROTENBERRY. 1980. Patterns of morphology and ecology in grassland and shrub- steppe bird populations. Ecol. Monogr. 50: 287-308. WILLSON, M. F. 1974. Avian community organization and habitat structure. Ecology 55: 1017-1029. Purchased by the USDA Forest Service for official use. CHLOROPLAST ULTRASTRUCTURE IN THE DESERT SHRUB CHRYSOTHAMNUS NAUSEOSUS SSP. ALBICAULIS Craig E. Coleman! and William R. Andersen! ABSTRACT. — Ultrastructure of the chloroplasts of white rubber rabbitbrush (Chrysothamnus nauseosus (Pallas) Britt. ssp. albicaulis ) was observed with electron microscopy. In addition, leaf anatomy was observed with light microscopy. Previously, it had been reported that the leaves of this desert shrub exhibited a relatively high rate of photosynthesis when compared to other C; plants. Comparisons with chloroplasts of other C, and C, plants demonstrated a reduced amount of granal stacking in the rabbitbrush. However, the classification of rabbitbrush as a C; plant is confirmed. RUBP-carboxylase concentration is reported at about 450 mg - ml’ stromal space based on the estimation of 1 mg of chlorophyll per 25 ul of stromal space in a normal C; chloroplast and data from an assay to determine the ratio of RUBP-carboxylase to chlorophyll. Recent interest in the desert shrub, Chrysothamnus nauseosus (Pallas) Britt. ssp. albicaulis (white rubber rabbitbrush), has led to a number of studies related to its ecology and physiology (McArthur et. al. 1979). Of particular interest has been its study as a non- traditional source of rubber. Ostler (1980) re- ported rubber acquisitions as high as 6% rub- ber per unit dry weight from the plant. As a result of this finding, studies are being con- ducted to discover some of the factors con- trolling the production of rubber. Recently work was done to determine some basic as- pects of the photosynthetic characteristics of the plant. It was discovered that, under non- stressed conditions, white rubber rabbitbrush exhibits a relatively high rate of photosynthe- sis when compared to other C; plants (Davis et. al., 1985), and it was felt that an electron microscopic analysis of the chloroplast ultra- structure might shed some additional light on the problem. This paper, therefore, presents the result of this analysis, along with anatomi- cal data obtained from light microscopy. Addi- tionally, we report the approximate RUBP- carboxylase concentration in the stromal space based on an estimate for the concentra- tion of chlorophyll in the stromal space. MATERIALS AND METHODS Leaves were removed from young branches of white rubber rabbitbrush plants growing in the greenhouse at Brigham Young University. The leaves were cut into sections approxi- mately 1-2 mm long and placed immediately in 0.2 M sodium cacodylate buffered (pH 7.3) 2% glutaraldehyde 3% acrolein (v/v) solution for two hours for fixation. After being washed for one hour with a 1:1 solution of buffer and distilled water, the material was stained with 2% osmium tetroxide (w/v) diluted 1:1 with the sodium cacodylate buffer. The material was again washed with the buffer solution for one hour and subsequently dehydrated using an ethyl alcohol series. The material was then embedded in Spurr’s resin (Spurr 1969) by first rinsing three times in 100% acetone. It was allowed to stand for one hour each in first a 25% resin to acetone solution (v/v), then a 75% solution before fi- nally embedding in 100% resin. Sections were obtained using a glass knife in a Porter-Blum MT-2 ultra-microtome. For electron mi- croscopy, sections were mounted on copper grids previously coated with formvar and a light layer of carbon. Lead citrate was used as a poststain as previously described (Reynolds 1963). The sections were observed and pho- tographed with a Phillips EM 400 transmis- sion electron microscope. For light mi- croscopy, sections were taken from the same resin-embedded material as used for electron microscopy. These sections were mounted on a glass slide and poststained with a 1% Tolu- idine Blue, 1% Azure II, and 1% NaHCO, ‘Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. O73 574 GREAT BASIN NATURALIST Vol. 46, No. 3 co «a Fig. 1. Light micrograph of the transverse section of a rabbitbrush leaf showing bundle sheath cells (bs), mesophyl! cells (m), stoma (s), vascular bundle (bs)(X170). stain. An enzyme assay to determine the ratio of RUBP-carboxylase to chlorophyll was per- formed as described previously (Davis et. al. 1985). RESULTS LIGHT MICROSCOPpYy.—Transverse sections were viewed with the light microscope (Fig. 1) with the intent to compare the anatomy of the white rubber rabbitbrush with that of known C, and C, plants. Chloroplasts are located exclu- sively in the mesophyll cells and are lacking in the bundle sheath cells. This indicates the pres- ence of C; metabolism (Laetsch 1974). ELECTRON MICROSCOPY.—Typical chloro- plasts observed with the electron microscope are shown in Figures 2 and 3. It should be noted that in all the chloroplasts observed there exsisted a uniformity of structure, in other words, a complete lack of dimorphism, which is usually exhibited in chloroplasts of C, plants (Laetsch 1968). Furthermore, chloro- plast size and distribution do not seem to vary from one section of the leaf tissue to another. The enzyme assay to determine the ratio of RUBP-carboxylase to chlorophyll in the chloroplast yielded a result of 11.24 + 0.81 mg RUBP-carboxylase. DISCUSSION To determine if the chloroplasts of the white rubber rabbitbrush were unusual in any way, their ultrastructure was compared to the ultrastructure of the chloroplasts from several C; and C, plants reported in the literature. Thus, comparisons were made with chloro- plasts from radish (Rufner et. al. 1984), barley (Robertson and Laetsch 1974), rye (Huner 1984), tobacco (Laetsch and Stetler 1965, Kasperbauer and Hamilton 1984), bean (Weier and Thomson 1962), rice (Miyake and Maeda 1976), spinach, and tomato (Rufner and Barker 1984). Although typical of C, anatomy, we note that the grana in the chloro- plasts of the white rubber rabbitbrush seem to be less stacked in comparison to chloroplasts of mesophyll cells in both C; and C, plants. This, however, could be due to a variety of July 1986 COLEMAN, ANDERSEN: CHLOROPLAST ULTRASTRUCTURE O75 Fig. 2. Transmission electron micrograph of a rabbitbrush chloroplast lacking starch grains showing cell wall (cw)(X40,000). factors, including the age of the leaf (Robertson and Laetsch 1974), the amount of light (Apel 1983), and the quality of light (Kasperbauer and Hamilton 1984) being absorbed by the leaves of the plants, or the absence of vital nutrients and mineral in the soil, such as iron (Rufner and Barker 1984). A most likely explanation for the _ reduced stacking is the age of the chloroplasts, _ since the leaves, although fully expanded, were _ taken from young shoots. Heldt (1979) estimated the chlorophyll con- tent in a normal C; chloroplast to be about 1 mg per 25 ul of stromal space. Data from an enzyme assay reported previously indicate the ratio of RUBP-carboxylase to chlorophyll in rabbitbrush to be 12.93 + 0.74 mg RUBP-carboxylase - mg” chlorophyll (Davis et. al. 1985). This assay was repeated for the purpose of this paper and, as already stated, yielded a value of 11.24 + 0.81 mg - mg’. Assuming Heldt’s estimation of stro- mal space to be reasonable in this case, the con- centration of RUBP-carboxylase in rabbitbrush chloroplasts is about 450 mg - ml’. This is a high concentration of the enzyme as compared to sev- eral other C, plants (values, on the average, are reported around 200-250 mg - ml’) (Ashton 1982, Kawashima and Mitake 1969; Lyttleton and Ts‘o 1958; Molin et. al. 1982) and may be responsible for the increased rate of photosyn- thesis as observed in the plant. ACKNOWLEDGMENTS We wish to express gratitude to Dr. W. M. Hess and the staff of the Electron Optics Labora- tory for their assistance in completing the mi- croscopy for this paper. LITERATURE CITED APEL, K. 1983. The light-dependent control of chloroplast development in barley (Hordeum vulgare L.). J. Cell Biol. 23(1—4): 181-189. ASHTON A. R. 1982. A role for ribulose-1,5—bisphosphate carboxyl ase as a metabolite buffer. FEBS Lett. 145(1): 1-7. Davis, T. D., N. SANKHLA, W. R. ANDERSON, E. J. WEBER AND B. N. Smiru. 1985. High rates of photosynthe- sis in the desert shrub Chrysothamnus nauseosus ssp. albicaulis. Great Basin Nat. 45: 520-526. HELpr, H. W. 1979. Light-dependent changes of stromal H* and Mg”* concentrations controlling CO, fixa- tion. Pages 202-207 in: Vol. 6, Encyclopedia of plant physiology (Springer), new series. 576 GREAT BASIN NATURALIST Vol. 46, No. 3 Fig. 3. Transmission electron micrograph of a rabbitbrush chloroplast with a prominent starch grain showing cell wall (cw), starch grain (sg)(X4500). HUNER, N. P. A., B. ELEMAN, AND M. KROL. 1984. Growth and development at cold-hardening temperatures. Chloroplast ultrastructure, pigment content, and composition. Canadian J. Bot. 62: 53-60. KASPERBAUER, M. J., AND J. L. HAMILTON. 1984. Chloroplast structure and starch grain accumulation in leaves that received different red and far-red levels during de- velop ment. Plant Phsiol. 74: 967-970. KAWASHIMA, N., AND T. MITAKE. 1969. Studies on protein metabolism in higher plants. IV, Changes in ribulose diphosphate carboxylase activity and fraction I protein content in tobacco leaves with age. Agric. Biol. Chem. 33(4): 539-543. Laetscu, W. M. 1968. Chloroplast specialization in dicotyle- dons possessing the C,-dicarboxylic acid pathway of photosyn thetic CO, fixation. Amer. J. Bot. 55: 875-883. ____.. 1974. The C, syndrome: A structural analysis. Ann. Rey. Plant Physiol. 25: 27-52. LAETSCH, W. M., AND D. A. STETLER. 1965. Chloroplast struc- ture and function in cultured tobacco tissue. Amer. J. Bot. 52(8): 798-804. LYTTLETON, J. W., AND P. O. P. Ts'o. 1958. The localization of fraction I protein of green leaves in the chloroplasts. Arch. Bio. Biophys. 73: 120-126. McArtuur, E. D., A. C. BLaugrR, A. P. PLUMMER, AND R. STEVENS. 1979. Characteristics and hybridization of important inter mountain shrubs. II., Sunflower family. U.S. Dept. of Agriculture, Forest Service Re- search Paper INT-220. Miyake, H., AND E. MAEDA. 1976. Development of bundle sheath chloroplasts in rice seedlings. Canadian J. Bot. 54(7): 556-565. MOLIN W. T., S. P. MEYERS, G. R. BAER, AND L. E. SCHRADER. 1982. Ploidy effects in isogenic populations of alfalfa. Plant Physiol. 70: 1710-1714. OSTLER, W. K. 1980. Selection of acquisitions of Chrysotham- nus for rubber production. Native Plants Inc., Salt Lake City. NSF Report. REYNOLDS, E. S. 1963. The use of lead citrate at high pH as an electron opaque stain in electron microscopy. J. Cell Biol. 17: 208-212. ROBERTSON, D., AND W. M. Laetscu. 1974. Structure and function of developing barley plastids. Plant Physiol. 54: 148-159. RUFNER, R., AND A. V. BARKER. 1984. Ultrastructure of zinc induced iron deficiency in mesophyll chloroplasts of spinach and tomato. J. Amer. Soc. Hort. Sci. 109(2): 164-168. RUENER, R., A. V. BARKER, J. P. BOUCHER, W. KROLL AND T. A. HosMER. 1984. Effects of nitrapyrin and nitrogen fer- tilizers on ultrastructure of mesophyll chloroplasts of radish. J. Amer. Soc. Hort. Sci. 109(2): 139-144. Spurr, A. R. 1969. A low viscosity epoxy resin embedding medium for electron microscopy. J. Ultrastruct. Res. 26: 31-43. WEIER, T. E., AND W. W. THoMsON. 1962. Membranes of mes- ophyll cells of Nicotiana rustica and Phaseolus vul- garis with particular reference to the chloroplast. Amer. J. Bot. 49(8): 807-820. GENETIC VARIATION OF WOODRATS (NEOTOMA CINEREA) AND DEER MICE (PEROMYSCUS MANICULATUS) ON MONTANE HABITAT ISLANDS IN THE GREAT BASIN William T. Mewaldt'” and Stephen H. Jenkins! ABSTRACT.—Seventeen loci were examined for polymorphism in four populations of Neotoma cinerea and Per- omyscus maniculatus on isolated mountain ranges in the Great Basin, one population of each in the Sierra Nevada, and one of each in the Rocky ponies All Peromyscus populations had higher levels of heterozygosity than syntopic Neotoma populations. Results cate; interstriae 1 moderately elevated, very slightly higher than interstriae 3, with a median Populations of species restricted to terres- trial habitat islands may be similar to those on oceanic islands in patterns of gene frequency change over time (Kilpatrick 1981). For exam- ple, Glover et al. (1977) studied pikas (Ochotona princeps), which are generally re- stricted to talus slopes and other rocky habi- tats at high elevations. Their findings of low heterozygosity within populations are consis- tent with patterns on oceanic islands reviewed by Soule (1976) and Kilpatrick (1981), who concluded that the stochastic processes of founder effect and genetic drift may cause unpredictable shifts in gene frequency and reduced heterozygosity. The distribution of montane habitats in the Great Basin of the western U.S. provides an opportunity for “island-mainland” comparisons of montane mammals. In the Great Basin, isolated patches of forested habitat act as refugia for many populations of mammals that apparently cannot live in or cross the intervening deserts (Brown 1971). As recently as 8,000 years ago, | climatic conditions were such that forests (in- terspersed with pluvial lakes) existed continu- ously across the Great Basin from the Sierra | Nevada to the Rocky Mountains. Since that | time, the climate has become drier, resulting in isolation of montane habitat at higher eleva- tions of the mountain ranges (Wells 1983). We analyzed allozymes in several populations of two cricetid rodent species, Neotoma cinerea acraia (Elliot) (bushy tailed woodrat) and Per- omyscus maniculatus sonoriensis (LeConte) Department of Biology, University of Nevada, Reno, Nevada 89557. (deer mouse), to test some genetic predictions of the hypothesis that stochastic effects should be more pronounced for isolated island populations than for “mainland” populations. Peromyscus maniculatus sonoriensis is distributed continu- ously across the Great Basin (Hall 1946), whereas Neotoma cinerea acraia is found in iso- lated populations on most of the Basin ranges (Brown 1971). Biochemical variation within at least 20 spe- cies of Peromyscus has been reported (e.¢g., Se- lander et al. 1971, Kilpatrick and Zimmerman 1976, Zimmerman et al. 1978, Avise et al. 1979, Gill 1980). Of special interest here are the stud- ies by Avise et al. (1979) and Gill (1980), both of which included P. m. sonoriensis. Avise et al. reported mean heterozygosity (H) values for populations of this subspecies ranging from 0.074 to 0.124, whereas Gill found an H of 0.118. Electrophoretic studies of Neotoma are few. Mascarello (1978) studied three chromosomal races of Neotoma lepida in the southwest but did not report H values, whereas Zimmerman and Nejtek (1977) reported H values for three semi- species of Neotoma (N. albigula, N. micropus, and N. floridana) in southern North America ranging from 0.024 to 0.140, with an average of 0.078. Heterozygosity measures for populations of N. cinerea have not been reported. MATERIALS AND METHODS Six mountain ranges with populations of N. c. acraia and P. m. sonoriensis were chosen *Present address: Western Nevada Community College, Fallon, Nevada 89406. O77 578 TABLE 1. Allele frequencies for polymorphic loci of Neotoma cinerea GREAT BASIN NATURALIST Vol. 46, No. 3 and Peromyscus maniculatus, and mean heterozygosity per locus over all loci sampled (H).* Population Sample ES-B ES-C AAT-1 GDH PGD H size Neotoma cinerea Sierra Nevada 16 100 100 100 (.93) 84 (.04) 100 0.013 (Carson Range) 120 (.07) 100 (.96) Shoshone 17 82 (.025) 100 100 (.80) 84 (.05) 92 (.025) 0.034 100 (.975) 120 (.20) 100 (.925) 100 (.975) 118 (.025) Toiyabe 14 100 94 (.18) 100 (.93) 84 (.21) 100 0.046 100 (.82) 120 (.07) 100 (.79) Toquima 15 100 94 (.07) 100 (.87) 84 (.10) 100 0.033 100 (.93) 120 (.13) 100 (.90) Snake 14 100 100 (.93) 100 (.77) 100 (.97) 92 (.03) 0.037 104 (.07) 120 (.23) 118 (.03) 100 (.97) Rocky Mountains 15 100 94 (.31) 100 (.97) 100 100 0.030 (Tushar Range) 100 (.69) 120 (.03) Population Sample ES-A ES-C AAT-1 GDH LDH-2 PGD H size Peromyscus maniculatus Sierra Nevada 18 100 (.91) 100 (.74) 100 (.76) 89 (.15) 100 100 0.075 (Carson Range) 118 (.09) 106 (.26) 120 (.24) 100 (.85) Shoshone 15 100 (.97) 100 (.63) 100 (.77) 100 (.97) 100% 100 0.064 118 (.03) 106 (.27) 120 (.23) — 111 (.03) 110 (.10) Toiyabe 15 100 (.97) 100 (.73) 100 (.67) 89 (.13) 100 83 (.13) 0.108 123 (.03) 102 (.03) 114 (.10) 100 (.70) 100 (.87) 106 (.20) 120 (.23) 111 (.17) 110 (.03) Toquima 18 100 (.94) 100 (.58) 100 (.53) 89 (.14) 100 83 (.14) 0.112 118 (.06) 106 (.39) 120 (.47) 100 (.80) 100 (.83) 110 (.03) 111 (.06) 112 (.03) Snake 21 100 100 (.64) 100 (.57) 89 (.045) 91 (.02) ~—- 83 (.02) ~—«*0.079 106 (.34) 120 (.41) 100 (.91) 100 (.98) 100 (.98) 110 (.02) 138 (.02) 111 (.045) Rocky Mountains 13 100 (.96) 100 (.81) 100 (.65) 89 (.08) 100 100 0.064 (Tushar Range) 118 (.04) 110 (.19) 120 (.35) 100 (.92) *Alleles designated according to proportional electrophoretic mobility relative to the most common allele (100). Frequencies given in parentheses. H calculated according to formula (3) in Nei (1978), for 17 scorable loci for Neotoma and 16 scorable loci for Peromyscus . for analysis (Table 1). All mountain ranges are over 80 km long and have peaks of over 3,000 m. Animals were collected in the summers of 1978 and 1979. Total sample sizes were 91 for N. c. acraia and 100 for P. m. sonoriensis (Table 1). Techniques of tissue (liver and kidney) preparation and enzyme staining for all loci except esterases were modified from Selander et al. (1971) and Gabriel (1971). Esterase anal- ysis using napthol-AS-D-acetate as a substrate followed Van Deusen and Kaufman (1978). All electrophoresis was done in polyacrylamide slab gels, with both homogeneous and gradi- ent type gels run in a water-cocled Pharmacia electrophoresis tank. Use of polyacrylamide rather than starch gels permitted better reso- lution of separate bands for some loci, such as the esterases. In all, seventeen presumed loci were ana- lyzed for each species. These included car- boxylesterase (ES-A, ES-B, and ES-C; E.C. 3.1.1.1), phosphoglucomutase (PGM; E.C. 5.4.2.2), malate dehydrogenase (MDH-1 and MDH-2; E.C. 1.1.1.37), malate dehydroge- nase (oxaloacetate-decarboxylating) (NADP’) July 1986 MEWALDT, JENKINS: WOODRAT VARIATION 579 TABLE 2. Genetic distances* between pairs of Neotoma cinerea populations (above diagonal) and pairs of Peromyscus maniculatus populations (below diagonal) on mountain ranges in the Great Basin. Shoshone Sierra Toiyabe Toquima Snake Rocky Nevada Mountains Sierra Nevada .00043 .00263 0 .00107 .00532 Shoshone 0 .00316 0 0 .00711 Toiyabe .00093 .00288 .00034 .00409 .00262 Toquima .00401 .00473 .00240 .00022 .00387 Snake .00217 .00093 .00331 0 .00645 Rocky Mountains .00258 .00236 .00371 .00793 .00477 *Formula for genetic distance (D) from Nei (1978). (MDH-3; E.C. 1.1.1.40), cytosol aminopepti- dase (CAP; E.C. 3.4.11.1), L-lactate dehydroge- nase (LDH-1 and LDH-2; E.C. 1.1.1.27), glu- cose dehydrogenase (GDH; E.C. 1.1. 1.47), isocitrate dehydrogenase (NADP’) (IDH; E.C. 1.1.1.42), aspartate aminotransferase (AAT-1 and AAT-2; E.C. 2.6.1.1), xanthine de- hydrogenase (XDH; E.C. 1.1.1.204), phospho- gluconate dehydrogenase (PGD; E.C. 1.1.1.43), and superoxide dismutase (Sod; E.C. 1.15.1.1). Choice of enzymes for analysis was based on our ability to obtain reproducible and unambiguous results from among those enzymes listed and studied by Selander et al. (1971). Enzyme nomenclature and E..C. numbers are from Moss (1982) and International Union of Biochemistry (1984). RESULTS AND DISCUSSION Twelve of the loci analyzed were monomor- phic in Neotoma and 10 were monomorphic in Peromyscus. For both species, the same allele was fixed in all 6 populations at each of the monomorphic loci. Five loci were polymor- phic in one or more populations of Neotoma ; 7 were polymorphic in one or more populations of Peromyscus (Table 1). Esterase-B was so variable in Peromyscus that it could not be scored accurately; this locus is omitted from Table 1. Extremely low interpopulation genetic dis- tances, D (Nei 1978), and proportionately large standard errors showed no clear patterns in either species (Table 2). The mean D was 0.0028 for all pairs of P. maniculatus popula- tions and 0.0025 for all pairs of N. cinerea populations. There was no significant correla- tion between genetic distance and geographic distance for either species (r = 0.48 for N. cinerea, 0.05 < P < 0.10; r = 0.12 for P. maniculatus ; P > 0.50). Heterozygosity (H) was significantly great- er for P. maniculatus (mean for the 6 popula- tions = 0.084) than for N. cinerea (mean = 0.032; P = 0.03 by randomization test for matched pairs). The difference between spe- cies would be even greater if we had not ex- cluded the highly variable ES-B locus from calculations for Peromyscus. Heterozygosity values were greater for all the central Great Basin populations of N. cinerea than for either the Sierra or Rocky Mountain population. The same was true for all but one of the P. manicu- latus Great Basin populations (Table 1). Our results in general are not consistent with the expectation that gene flow should be less and genetic drift greater for Neotoma cinerea populations than for Peromyscus maniculatus populations on isolated mountain ranges in the Great Basin. One or both of two factors could account for this inconsistency. First, N. cinerea populations might not be as isolated in montane habitats on Great Basin ranges as we initially assumed. Indeed, N. cinerea are occasionally captured below the lower tree line in some parts of Nevada (Hall 1946, personal observations). Sufficient inter- population gene flow could forestall genetic divergence. Second, the populations studied may have been too large or the time since isolation of their forested habitats too short for measurable genetic drift to have occurred. Our finding of significantly greater het- erozygosity for populations of P. maniculatus than for the syntopic populations of N. cinerea is consistent with a general pattern of greater genetic variablity of P. maniculatus than is found in most other rodent species that have been examined (Smith et al. 1978, Avise et al. 1979, Smith 1981). The niche-width variation hypothesis (Nevo 1978) might account for this pattern, since P. maniculatus is more general- ized in both diet and habitat than N. cinerea 580 and many other rodents. The high reproduc- tive rate of P. maniculatus may also contribute to its ability to maintain relatively high levels of heterozygosity (Smith 1981). Populations of N. cinerea and P. manicula- tus on isolated mountain ranges in the Great Basin were generally more heterozygous than populations in Sierra Nevada or Rocky Moun- tain sites. The latter sites were near the range limits for both N. cinerea acraia and P. manic- ulatus sonoriensis, the subspecies which were used in our study (Hall 1981). McClen- aghan and Gaines (1981) documented greater genetic variability for central than for mar- ginal populations of Sigmodon hispidus; our results exhibit a similar pattern at the sub- specific level. Although our initial predictions were not verified for the particular populations we studied, the Great Basin system of terrestrial habitat islands seems to be well suited for testing a variety of hypotheses about mam- malian populations genetics. The paleoecol- ogy of this area is well understood (Wells 1983), which provides a good foundation for such studies. ACKNOWLEDGMENTS We thank the University of Nevada Re- search Advisory Board for financial support, W. Welch for assistance with electrophoresis, and A. Gill, D. Hafner, and W. Welch for comments on the manuscript. LITERATURE CITED AVISE, J. C., M. H. SMITH, AND R. K. SELANDER. 1979. Biochemical polymorphism and systematics in the genus Peromyscus. VII. Geographic differentia- tion in members of the truei and maniculatus spe- cies groups. Journal of Mammalogy 60: 177-192. BROWN, J. H. 1971. Mammals on mountaintops: nonequi- librium insular biogeography. American Natural- ist 105: 467-478. GABRIEL, O. 1971. Locating enzymes on gels. Methods in Enzymology 22: 578-604. GILL, A. E. 1980. Evolutionary genetics of California is- lands Peromyscus. Pages 719-743 in D. M. Power, ed., The California islands. Santa Barbara Museum of Natural History, Santa Barbara, Cali- fornia. 787 pp. GLOVER, D. G., M. H. Smiru, L. AMES, J. JOULE, AND J. M. Dusacu. 1977. Genetic variation in pika popula- tions. Canadian Journal of Zoology 55: 1841-1845. GREAT BASIN NATURALIST Vol. 46, No. 3 HALL, E. R. 1946. Mammals of Nevada. University of Califor- nia Press, Berkeley, California. 710 pp. . 1981. The mammals of North America. 2d ed. John Wiley and Sons, New York. Vol 2: 601-1181 + 90. INTERNATIONAL UNION OF BIOCHEMISTRY. Nomenclature Committee. 1984. Enzyme nomenclature 1984. Aca- demic Press, Inc., Orlando, Florida. xx + 646 pp. KILPATRICK, C. W. 1981. Genetic structure of insular popula- tions. Pages 28-59 in M. H. Smith and J. Joule, eds., Mammalian population genetics. University of Geor- gia Press, Athens. 380 pp. KILPATRICK, C. W., AND E. G. ZIMMERMAN. 1976. Bio- chemical variation and systematics of Peromyscus pectoralis. Journal of Mammalogy 57: 506-522. MCCLENAGHAN, L. R., JR., AND M.S. GAINES. 1981. Genic and morphological variability in central and mar- ginal populations of Sigmodon hispidus. Pages 202-213 in M. H. Smith and J. Joule, eds., Mam- malian population genetics. University of Georgia Press, Athens. 380 pp. MASCARELLO, J. T. 1978. Chromosomal, biochemical, mensural, penile, and cranial variation in desert woodrats (Neotoma lepida). Journal of Mammal- ogy 59:477-495. Moss, D. W. 1982. Isoenzymes. Chapman and Hall, Lon- don. 204 pp. Nel, M. 1978. Estimation of average heterozygosity and genetic distance from a small number of individu- als. Genetics 89:583-590. Nevo, E. 1978. Genetic variation in natural populations: patterns and theory. Theoretical Population Biol- ogy 13: 121-177. SELANDER, R. K., M. H. SMITH, S. Y. YANG, W. E. JOHNSON, AND J. B. Gentry. 1971. Biochemical polymor- phism and systematics in the genus Peromyscus. I. Variation in the old-field mouse (Peromyscus polionotus ). Studies in Genetics 6: 49-90. SMITH, M. F. 1981. Relationships between genetic vari- ability and niche dimensions among coexisting species of Peromyscus. Journal of Mammalogy 62: 273-285. SMITH, M. H., M.N. MANLOVE, AND J. JOULE. 1978. Spatial and temporal dynamics of the genetic organization of small mammal populations. Pages 99-113 in D. P. Snyder, ed., Populations of small mammals under natural conditions. University of Pitts- burgh, Pittsburgh, Pennsylvania. xiii + 237 pp. SOULE, M. 1976. Allozyme variation: its determinants in space and time. Pages 60-77 in F. Ayala, ed., Molecular evolution. Sinauer Associates, Sunder- land, Massachusetts. x + 277 pp. VAN DEUSEN, M., AND D. W. KaurMaNn. 1978. Esterase-1 as a genetic marker for Peromyscus. Journal of Mammalogy 59: 185-186. WELLS, P. V. 1983. Paleobiogeography of montane islands in the Great Basin since the last glaciopluvial. Ecological Monographs 53: 341-382. ZIMMERMAN, E. G., AND M. E. NEJTEK. 1977. Genetics and speciation of three semispecies of Neotoma. Jour- nal of Mammalogy 58: 391—402. ZIMMERMAN, E. G., C. W. KILPATRICK, AND B. J. HART. 1978. The genetics of speciation in the rodent genus Peromyscus. Evolution 32: 565-579. BREEDING RECORDS FOR CLARK'S GREBE IN COLORADO AND NEVADA Richard L. Bunn! ABSTRACT.— Described as a new breeding species is Clark’s Grebe in Colorado and Nevada. The Western Grebe (Aechmophorus occi- dentalis) and Clark's Grebe (A. clarkii) were originally named and described by Lawrence (1858 in Baird 1858) as two species based on plumage, bill structure, and color, and size. Coues (1874) reduced the Clark’s Grebe to subspecific rank, which was upheld by the American Ornithologists Union (in Nu- echterlein 1981). Recently, a decision by the AOU Committee on Classification and Nomenclature (AOU 1985) restored the Clark’s Grebe to specific rank, based on stud- ies of assortative mating (Storer 1965, Ratti 1979), spatial segregation (Ratti 1979), and, within sympatric populations, advertising calls and ecological segregation (Nuechterlein 1981). Breeding populations of Clark's Grebe are known in Oregon, California, Utah, North Dakota, and South Dakota in the United States; Manitoba and Saskatchewan in Canada; Chihuahua, Durango, Zacatecas, Na- yarit, Jalisco, Michoacén, Guanajuato, San Luis Potosi, State of Mexico, and Guerrero in Mexico (Storer 1965, Ratti 1979, 1981, Nuechterlein 1981, Williams 1982). This note reports the nesting of Clark’s Grebe in Colo- rado and Nevada. Two courting pairs of Clark's Grebe were ob- served 21 June 1978 at Barr Lake, Adams County, Colorado (B. Andrews, personal com- munication). Since then Clark’s Grebe has been sighted in small numbers in eastern Colorado in the South Platte River drainage. They are appar- ently far less abundant there than the Western Grebe during migration and during the nesting season (personal observation). However, in the Arkansas River valley and areas in the San Luis Valley, the Clark’s Grebe represented approxi- mately half or more of the breeding Aechmopho- rus grebe population on several reservoirs from 1983 to 1985. On 4 July 1983 Charles Chase III and I found 20 adult and 5 juvenile Clark’s Grebes in Saguache County, Colorado, southwest of Russell Lakes State Wildlife Area. Four juve- niles occurred singly and were each accompa- nied by one adult bird; one juvenile was ac- companied by two adults. The grebes were seen on two reservoirs that had extensive bull- rush stands along their western edge. The advertising calls of the Clark's Grebe were heard throughout the visit, but Western Grebes were neither seen nor heard on either reservoir. On 17 June 1984 I saw 33 adult and 8 juve- nile Clark's Grebes at Stillwater National Wildlife Refuge, Fallon, Nevada. The juve- niles occurred singly with either 1 or 2 accom- panying adults on Lead and Goose lakes. Only adult Clark's Grebes were seen on Stillwater Point Reservoir. ACKNOWLEDGMENT I thank Bob Andrews for the early records of Clark’s Grebe in Colorado and Diana F. Tomback, Charles Chase III, and Steve Bissel for helpful criticisms of the manuscript. LITERATURE CITED AMERICAN ORNITHOLOGISTS UNION. 1985. Thirty-fifth supplement to the AOU check-list of North Amer- ican birds. Auk 102: 680-686. BairD, S. F. 1858. Birds. Pages 894-895 in Reports of explorations and surveys to ascertain the most practicable and economical route for a railroad from the Mississippi River to the Pacific Ocean, vol. 9, part 2. Washington, D.C., House of Repre- sentatives, 33d Congress, 2d Session. Ex. Doc. No. 91. Coues, E. 1874. Birds of the Northwest: a handbook of ornithology of the region drained by the Missouri River and its tributaries, Washington, D.C. Department of Biology, University of Colorado at Denver, Denver, Colorado 80202. 581 582 GREAT BASIN NATURALIST Vol. 46, No. 3 NUECHTERLEIN, G. L. 1981. Courtship behavior and re- |. 1981. Identification and distribution of Clark’s productive isolation between Western Grebe Grebe. Western Birds 12: 41—46. color morphs. Auk 98: 335-349. STORER, R. W. 1965. The color phases of the Western Grebe. Living Bird 4: 59-63. Ratti, J. T. 1979. Reproductive separation and isolating — Wiitiams, S. O., III. 1982. Notes on the breeding and mechanisms between sympatric dark- and light- occurrence of Western Grebes on the Mexican phase Western Grebes. Auk 96: 573-586. Plateau. Condor 84: 127-130. NOTICE TO CONTRIBUTORS Manuscripts intended for publication in the Great Basin Naturalist or Great Basin Natural- ist Memoirs must meet the criteria outlined in paragraph one on the inside front cover. They should be directed to Brigham Young University, Stephen L. Wood, Editor, Great Basin Naturalist, 290 Life Science Museum, Provo, Utah 84602. Three copies of the manuscript are required. 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No. 7 Biology of desert rodents. $8. ‘No. 8 The black-footed ferret. $10. TABLE OF CONTENTS North American stoneflies (Plecoptera): systematics, distribution, and taxonomic ref- erences. B. P. Stark, S. W. Szczytko, and R. W. Baumann. ................. 383 Three new records for diatoms from the Great Basin, USA. Samuel R. Rushforth, Lorin'E- Squires; and Jefirey, A. Johansents)-= 042. can eee ee 398 Movements by small mammals on a radioactive waste disposal area in southeastern Idaho: Craig: R. Groves‘and Barry. Keller. 7.52 45220450 oe te eee 404 Roll of three rodents in forest nitrogen fixation in western Oregon: another aspect of mammal-mycorrhizal fungus-tree mutualism. C. Y. Li, Chris Maser, Zane Maser; and Bruce A= Caldwell... 2205) 3. eee et ee ee eee ee 411 Canids from the late Pleistocene of Utah. Michael E. Nelson and James H. Madsen, Jr. Note on food habits of the Screech Owl and the Burrowing Owl of southeastern Oregon. Barbara A. Brown, John O. Whitaker, Thomas W. French, and Chris Maser is tic aioe arta Gio iy Pie det trate ghia) Van dai ay an NS) Se gay rae 421 Diseases associated with Juniperus osteosperma and a model for predicting their occurrence with environmental site factors. E. D. Bunderson, D. J. Weber, and Di Th. Nelsonse 202328 Bo aa we Bega cio oe 427 Status and distribution of the Fish Creek Springs tui chub. Thomas M. Baugh, John W. Pedretti, and James E-,Deacon®’.. sis... es ee Ge 44] Ponderosa pine conelet and cone mortality in central Arizona. J. M. Schmid, J. C. Mitchelli”and: Si An Mata: Ste ee bos oe sans heh ee 445 Number and condition of seeds in ponderosa pine cones in central Arizona. J. M. Schmid), SA: Matavand)f)(G: Mitchellh 33 522550 es see eee 449 New species of Protocedroxylon from the Upper Jurassic of British Columbia, Canada. David A. Medlyn and William D. Vidwelln”) 3... 5-2). 954.555. oe 452 New taxa and nomenclatural changes in Utah Penstemon (Scrophulariaceae). Eliza- bethiG: Neeseen ee a oes Bie ook led Sten oy AUS ee 459 Relict occurrence of three “American” Scolytidae (Coleoptera) in Asia. Stephen L. Wood and Elui-fen' Vine) 22). 225A. ee hook Lae ee 461 New genus of Scolytidae (Coleoptera) from Asia. Stephen L. Wood and Fu-sheng PR nam i ee A Sd le SU ae ra SU gery AO Sa ee 465 New Pseudoxylechinus (Coleoptera: Scolytidae) from India. Stephen L. Wood. ....... 468 Wildlife distribution and abundance on the Utah Oil Shale Tracts, 1975-1984. C. Val Grant Pie yO ae aaa nas ee ae OI 469 Comparison of vegetation patterns resulting from bulldozing and two-way chaining on a Utah pinyon-juniper big game range. J. Skousen, J. N. Davis, and Jack D. Brothersone 8s $21 7a hk ee a ae ne eke ee ea 508 Vertebrate fauna of the Idaho National Environmental Research Park. Timothy D. Reynolds, John W. Connelly, Douglas K. Halford, and W. John Arthur. ...... 513 Infection of young Douglas-firs and spruces by dwarf mistletoes in the Southwest. Robert Is.,,Mathiasen. 222. 2.45 session ee 528 Habitat relationships of saltcedar (Tamarix ramosissima) in central Utah. Jack D. Brotherson:and Von Winkel) 05.0.0 905 5 Yee eee ee 535 Characteristics of mule deer beds. H. Duane Smith, Mark C. Oveson, and Clyde L. Pritchett: 25030 G 255 Wa coe ee ieee OA EN Sees ee ee 542 Trumpeter swan (Cygnus buccinator) from the Pleistocene of Utah. Alan Feduccia and Charles'G: ©viatt. 04 oe ec ll er 2 cc 547 New species and a new combination of Mentzelia section Bartonia (Loasaceae) from the Colorado Plateau. H. Thompson and B. Prigge. ...............-++000+e 549 New variety of Mentzelia multicaulis (Loasaceae) from the Book Cliffs of Utah. Kaye H. Thorne and Frank J. Smith. .:<5.).). 8 gr) 22) ets Aes Oe eee 555 New variety of Mentzelia pumila (Loasaceae) from Utah. Kaye H. Thome. ........... 557 Agathoxylon lemonii sp. nov., from the Dakota Formation, Utah. William D. Tidwell and Gregory, F:Chaynr 0.1% of individuals) were examined during snow-free months in Convict Creek, a permanent snowmelt- and spring-fed stream in the Sierra Nevada of California. The communities were highly diverse. The most abundant taxa in the periphyton were diatoms (Achnanthes minutissima, Cocconeis placentula lineata, Cymbella microcephala, C. sinuata, Fragilaria construens, F. crotonensis, Navicula spp., Synedra acus, and S. rumpens), except in late spring and summer when several seasonal blue-green algae (Chamaesiphon incrustans, Lyngbya spp. and Oscillatoria spp.) are at their maximum densities. Most common periphyton taxa vary systematically in abundance with season, but relative abundances of taxa also appear to be influenced by streambed scouring and by concentrations of ambient nutrients. Data on population densities and length frequencies of larval and nymphal stages of common benthic insects and occurrences of pupal and adult stages were examined to determine life history patterns. Taxa hatching in winter and spring and abundant as immatures in late spring include ephemeropterans (Epeorus longimanus, Drunella flavilinea, and Caudatella heterocaudata), plecopterans (Calineuria californica, Doroneuria baumanni, and Pteronarcys prin- ceps) and dipterans (Cryptolabis sp.). Common taxa hatching in late spring or summer are the plecopteran Malenka (californica?) and the trichopterans Arctopsyche grandis and Rhyacophila acropedes. Several bivoltine and multi- voltine ephemeropterans (Baetis devinctus and B. tricaudatus) and dipterans (Simulium spp. and Chironomidae) have summer cohorts. Taxa hatching in late summer or autumn and most abundant in autumn include ephemeropterans (Baetis spp., Ephemerella infrequens, Epeorus dulciana, Ironodes lepidus , and Paraleptophlebia pallipes ), trichopter- ans (Hydropsyche oslari, Lepidostoma spp., Glossosoma califica, Micrasema sp., Brachycentrus americanus, Neophy- lax sp., and Rhyacophila vaccua) and dipterans (Antocha monticola, Pericoma sp., and Chironomidae). Major recurring events that may influence life history patterns and structure of the benthic insect community are (1) near-freezing, nighttime winter water temperatures and occasional anchor ice, (2) a prolonged period of high discharge in late spring and early summer (3) a brief summer, and (4) a prolonged period of moderate stream discharge in autumn when the substratum is stable and food is abundant. Despite early biogeographical research on aquatic insects of the Sierra Nevada in Califor- nia (see Usinger 1956), phenological data and information on structure of benthic insect communities in streams of this range are very limited. There is a similar lack of information on production of stream fauna and on the spe- cies composition and seasonality of stream pe- riphyton. We report results of a four-year in- vestigation of Convict Creek, a permanent snowmelt- and spring-fed stream draining a 41 km’ watershed on the steep eastern escarp- ment of the Sierra Nevada. The study area includes both mesic and xeric terrestrial vege- tation (Orr 1981). Objectives of this study were to determine (1) the species composition of periphyton and benthic insect communities of Convict Creek, (2) the temporal variation in population densities of common taxa, (3) basic life histories and annual production of the common benthic insects, and (4) effects of hydrologic extremes on community structure. The ecology of Convict Creek is compared to that of other streams at comparable altitudes in mountain ranges bordering the Great Basin of the western United States. STUDY AREA Convict Creek (Lat. 37° 37’ N, Long. 118° 50’ W) is in the Inyo National Forest in Mono County, California. The study area lies within Convict Creek basin at an altitude of 2,185 m in the reserve of the Sierra Nevada Aquatic Research Laboratory. The geology of the up- per part of the basin is unusual for the Sierra Nevada in that metamorphic rather than granitic rocks predominate. The lower part of the basin, including the study area, is com- posed of alluvium and morainal materials from erosion and glaciation of the area above. The metamorphic rock is of two major types, 1U.S. Geological Survey, Water Resources Division, 345 Middlefield Road, Menlo Park, California 94025. 595 596 siliceous hornfels and siliceous calc-hornfels (Rinehart and Ross 1964). The granitic rock is primarily quartz monzonite and granodiorite. Sandstones in the vicinity of Convict Creek are primarily quartz sandstone and calcareous quartz sandstone. Mineralogic analyses of fine sand and silt (fraction <74 wm) in the sedi- ments of Convict Creek indicate an abun- dance of feldspar, quartz, and calcite. A Great Basin sagebrush community domi- nated by big sagebrush, Artemisia tridentata Nutt., rabbit brush, Chrysothamnus nau- seosus (Pall.) Britt., and antelope bitterbrush, Purshia tridentata (Pursh) DC., covers much of the study area (Orr 1981). High desert ri- parian woodland occurs along the stream; dominant species of this community are quak- ing aspen, Populus tremuloides Michx., water birch, Betula fontinalis Sarg. and willows, Salix spp. Riparian meadow vegetation char- acterized by sedges, Carex spp., rushes, Jun- cus spp., and various grasses occur along the banks in low-lying areas of poorly drained soils. Convict Creek is a perennial riffle-pool stream. Fluctuations in discharge are moder- ated by Convict Lake 4 km upstream and a bypass channel permitting diversion of flood water around the study area. The streambed is rocky, ranging primarily from coarse sand and pebbles to cobbles. The water is clear; suspended sediment concentrations are typi- cally < 1 mg/liter. The stream is oligotrophic, with nitrate concentrations typically less than 10 wg/liter NO,;—N and ortho-phosphate con- centrations not exceeding 0.2 wg/liter PO} —P (Leland and Carter 1985). Silica con- centrations range from 7.1 to 8.3 mg/liter SiO,. The water is alkaline (pH 7.9 to 8.5) and always at or near saturation with respect to dissolved oxygen. Major ionic constituents (in mg/liter) determined during a period of base flow were: Ca”, 23; Mg”*, 0.3; K*, 0.7; Na’, 1.4; SOz , 11; and Cl, 0.2. The study area is the 340 m reach of Convict Creek that was used as a control section in experimental studies of Leland and Carter (1984, 1985) and Leland et al. (1986). For a map of the area, see Leland and Carter (1984). MATERIALS AND METHODS Water temperature and stream discharge in the study area were monitored continuously. The temperature sensor was located 5 cm be- GREAT BASIN NATURALIST Vol. 46, No. 4 low the surface of the streambed. A water stage servo-manometer with nitrogen-purge system (bubble gage—U.S. Geological Sur- vey 1962) was used to determine discharge from 1978 through 1980. Stream discharge in 1977 was estimated from records maintained by the Los Angeles Department of Water and Power at a site approximately 2 km upstream of the study area. Degree-day estimates were calculated from 1 January. Samples of benthic algae were scraped from 5x5 cm delineated areas of the upper surfaces of three separate cobbles (approximately 8 to 12 cm diameter) from the middle of the stream in unshaded riffles. Six samples (three per site) were taken from the same two riffles (each approximately 3 m by 30 m) each date. Sampling was approximately monthly from June through November in 1979 and 1980. Only algae with chloroplasts and intact cell walls were included in the counts (Leland and Carter 1984). Numbers of individuals record- ed for each taxon were cells for free and colo- nial species and filament fragments for fila- mentous forms (according to the methods of Greeson et al. 1977, section 7.3). Most di- atoms were identified to species, whereas green and blue-green algae were identified to genus. Sampling of aquatic insects was conducted monthly to bimonthly from late spring through autumn of the years 1977 through 1980. The same riffle (approximately 3 m by 50 m) was sampled each date. An invertebrate box sampler (Ellis-Rutter)” with a net of 0.35 mm mesh and sampling area of 0.1 m* was used. Sampling was from the middle of the stream and included both upstream and downstream areas of the riffle. Three samples were taken each sampling date to permit esti- mation of mean population densities. Samples were preserved in 70% ethanol and sorted in the laboratory with the aid of sugar flotation (Anderson 1959). All aquatic insects were identified to the lowest taxonomic level prac- tical (genus or species). Observations on time of adult emergence, composition of the drift, and rearing of later instars were conducted to provide additional life history information. Abundance data are expressed as popula- tion densities (individuals/em* of benthic al- gae and individuals/0. 1 m° of benthic insects). 2Use of brand names is for identification only and does not constitute endorsement by the U.S. Geological Survey. October 1986 LELAND ET AL.: SIERRA NEVADA AQUATIC INSECTS 597 [ 1977 1978 Temperature 20b ae ---- Discharge lo rh wal W Nov Ir All \ Peak flow 4 {\ | ~~ [Vv \. /) \ 10 NY Un + as al Nh V "| n\ \ 1 S -— — — — _ —| eJ e | I ! 3 Ww A) /\ \ Sich flow period *— as a \ Ay imurty fer = ath ~ A > i y Po 2 of a ce ee eee eee ea Baie Tis one ree li eee 1 N90 c | 1979 1980 < Qa 2 0 ar 22 = : Q ; mt jf i “yf Wh \ a KA 7 te : | 41 N \\, | 4 You seen l ce ace SIN = el J FP WM AS Med JI A& 8 © N D MONTH Fig. 1. Mean daily water temperature and mean daily discharge of Convict Creek. Abundant taxa are defined as those with den- sities greater than about 2% of the individuals of all taxa. Common taxa are those with densities greater than 0.1% of the individuals of all taxa. Secondary production was estimated using the size-frequency method (Hynes and Cole- man 1968, Hamilton 1969, Benke 1979, Krueger and Martin 1980), with computations based on body lengths and dry weights of common taxa. Cohort production intervals, in days from hatching to the attainment of the largest aquatic size class, were determined from life history and abundance data summa- rized in Figure 4. Sampling began in August 1979 and continued until June 1980. Intervals between samples were 24, 38, 41, 166, and 39 days. RESULTS AND DISCUSSION Environmental Variables Continuous records of stream discharge and water temperature were obtained for most of 1977 through 1980 (Fig. 1). Convict Creek is typically at or near base flow in win- ter (December to mid-March). The stream discharge increases in late spring due to snowmelt higher in the basin. This annual period of high discharge begins in May or June and lasts one or two months. Peak dis- charge (in m’/s) was 0.50 in 1977, 1.30 in 1978, 1.15 in 1979, and 2.14 in 1980. A second high discharge period occurred in early September 1978 due to unseasonably heavy rains accom- panying Hurricane Norman. Stream dis- charge generally declines during late summer and autumn and reaches baseflow in October. Mean daily water temperatures in winter ranged from 0.6 to 3.6 C in 1979 and 1980 (Fig. 1). Winter air temperatures were gener- ally lower in 1978 and 1979 than in 1977 (a mild winter) or in 1980 (when an exceptionally heavy snow cover was present). Anchor ice formed more frequently during the winters of 1978 and 1979 than during the other two years. Mean daily water temperatures during summer (July-August) were higher in 1977 and 1979 than in 1980. Daily means in Octo- ber did not vary substantially among years. Periphyton CoMMUNITY ComposITION.—The periphy- ton of Convict Creek is highly diverse (Table 1). A total of 30 genera (at least 104 species) of diatoms, 24 genera of green algae, and 29 genera of blue-green algae were identified GREAT BASIN NATURALIST TABLE 1. Benthic algae of Convict Creek CHLOROPHYTA Ankistrodesmus falcatus (Corda) Ralfs Carteria spp. Chaetophora spp. Characium spp. Cladophora spp. Closterium spp. Coelastrum spp. Coleochaete spp. Cylindrocapsa spp. Dictyosphaerium spp. Draparnaldia spp. Elakatothrix spp. Gloeocystis spp. Mougeotia spp. Oedogonium spp. Oocystis spp. Pediastrum spp. Protoderma spp. Rhizoclonium spp. Spirogyra grantiana Transeau Stigeoclonium spp. Ulothrix spp. Zygnema spp. EUGLENOPHYTA Euglena spp. Trachelomonas spp. PYRROPHYTA Ceratium hirundinella (O.F. Mull) Dujardin CHRYSOPHYTA Xanthophyceae Vaucheria spp. Chrysophyceae Dinobryon cylindricum Imhof ex Ahlstrom D. sertularia Ehr. Bacillariophyceae Achnanthes bergiani Cleve-Euler A. exigua Grun. A. lanceolata Breb. ex Kutz. A. linearis (W. Sm.) Grun. A. minutissima Kutz. Amphora ovalis (Kutz.) Kutz. A. perpusilla (Grun.) Grun. Amphipleura pellucida Kutz. Asterionella formosa Hass. Ceratoneis (=Hannaea) arcus Ehr. Kutz. Cocconeis pediculus Ehr. C. placentula euglypta (Ehr.) Cl. C. placentula lineata (Ehr.) V.H. Cyclotella comta (Ehr.) Kutz. C. kutzingiana Thwaites C. stelligera Cl. and Grun. Cymbella affinis Kutz. . brehmii Hust. . lanceolata (Ag.) Ag. . mexicana (Ehr.) Cl. . microcephala Grun. minuta minuta Hilse ex Rabh. . perpusilla Cl. . prostrata (Berk.) Cl. . sinuata Greg. . tumida (Breb ex Kutz.) V.H. DIDISIGISISAiGi@ Table 1 continued. Denticula spp. Diatoma hiemale (Lyngb.) Heib. D. vulgare Bory Diploneis ovalis oblongella (Naeg.) Cl. Epithemia adnata adnata (Kutz.) Breb. E. sorex Kutz. E. turgida (Ehr.) Kutz. E. turgida granulata (Ehr.) Brun Fragilaria brevistriata inflata (Pant.) Hust. F. construens (Ehr.) Grun. F. construens binodis (Ehr.) Grun. F. crotonensis Kitton F. leptostauron (Ehr.) Hust. F. pinnata Ehr. Frustulia spp. Gomphoneis herculeana (Ehr.) Cl. Gomphonema acuminatum Ehr. G. dichotomum Kutz. G. olivaceum (Lyngb.) Kutz. G. parvulum (Kutz.) G. subclavatum (Grun.) Grun. G. truncatum Ehr. Hantzschia spp. Melosira varians Ag. Navicula arvensis Hust. . aurora Sov. . bacillum Ehr. . capitata Ehr. . cocconeiformis Greg. ex Grev. . cryptocephala Kutz. ~~ hambergii Hust. lenceolata (Ag.) Kutz. pupula Kutz. radiosa Kutz. . rhynchocephala Kutz. . salinarum Grun. . seminulum Grun. . tripunctata (O.F. Mull.) Bory Neidium spp. Nitzschia acicularis W. Sm. N. actinastroides (Lemm) v. Goor N. amphibia Grun. N. dissipata (Kutz.) Grun. N. frustulum (Kutz.) Grun. N. linearis W. Sm. N. palea (Kutz.) W. Sm. N. sigma (Kutz.) W. Sm. N. vitrea Norman Pinnularia nodosa (Ehr.) W. Sm. P. rupestris Hantz. Rhopalodia gibba (Ehr.) O. Mull. Stauroneis spp. Stephanodiscus spp. Surirella spp. Synedra acus Kutz. S. radians Kutz. S. rumpens Kutz. S. rumpens fragilaroides Grun. S. ulna (Nitz.) Ehr. S S zezzzzz2z222222 . ulna oxyrhynchus Kutz. . ulna spathulifera (Grun.) V.H. Tabellaria fenestrata (Lyngb.) Kutz. RHODOPHYTA Batrachospermum spp. Vol. 46, No. 4 October 1986 Table 1 continued. CYANOPHYTA Amphithrix spp. Anabaena spp. Anabaenopsis spp. Aphanocapsa spp. Calothrix spp. Chamaesiphon incrustans (Grun. ) Chroococcus spp. Coelosphaerium spp. Colosterium spp. Cylindrospermum spp. Dactylococcopsis spp. Dichothrix spp. Gloeotrichia spp. Hapalosiphon spp. Lyngbya spp. Merismopedia spp. Microcystis spp. Nodularia spp. Nostoc spp. Oscillatoria spp. Phormidium spp. Plectonema spp. Raphidiopsis spp. Rivularia spp. Schizothrix spp. Scytonema spp. Spirulina spp. Stigonema spp. Tolypothrix spp. during the study. All of the most abundant taxa are diatoms, except for a few blue-greens that attain high abundance in late spring— summer. The composition of benthic algae in Sierra Nevada streams is poorly known. Hoff- man (1978) provided a partial inventory (23 species) of diatoms in Martis Creek, a peren- nial stream in the Truckee River Basin. Twelve of the 23 species, all cosmopolitan in their distribution, are common in both Martis Creek and Convict Creek. Sanford (1972) listed the conspicuous benthic algae in streams draining Feeley Lake and Round Lake in Tahoe National Forest; the dominant taxa in these streams are not abundant in Con- _ vict Creek. SEASONAL ABUNDANCES OF COMMON Taxa.—Mean population densities of 22 com- mon benthic algae in Convict Creek from late spring through autumn are presented in Fig- ure 2. An ordination method (detrended cor- respondence analysis [Hill 1979, Hill and Gauch 1980, Leland and Carter 1986]) was used to objectively order the taxa. The order- _ ing emphasizes the seasonal progression from _ taxa most abundant in spring to taxa most LELAND ET AL.: SIERRA NEVADA AQUATIC INSECTS 599 abundant in autumn. The major exceptions were Fragilaria crotonensis, characteristi- cally a planktonic species and perhaps an op- portunist in the periphyton of Convict Creek (Convict Lake is 4 km upstream), and Fragi- laria construens. Both taxa were highly vari- able spatially. Late spring—summer: The principal species of Gomphonema (parvulum, subclavatum, and truncatum) in Convict Creek were appar- ently most abundant in winter and early spring. The dominant algae in late spring and summer were the diatom Achnanthes minutissima and the blue-green Lyngbya spp.; population densities of the two co-domi- nants declined markedly by early autumn. Densities of A. minutissima and Lyngbya spp. were 3 and 11 times higher, respectively, in late spring 1980 than in late spring 1979, which accounted for a higher standing stock (total number of individuals of all taxa) in 1980. Other diatoms that had population max- ima in late spring-summer are Synedra acus, S. rumpens, and S. ulna. These species are early colonizers of denuded surfaces in Con- vict Creek (Leland and Carter 1984). The highly invasive blue-green Chamaesiphon in- crustans was also most abundant in late spring-summer. Late suwmmer-—early autumn: Stream dis- charge was substantially higher in late spring—summer 1980 than during the same period in 1979. By early August between-year differences in periphyton assemblages were apparent. The blue-green Oscillatoria spp. was very abundant in the summer of 1979 but not in 1980. The lower population density of Oscillatoria spp. in summer 1980 may have been related to the higher stream discharge. However, the rate of primary production was lower in 1980 (Leland and Carter 1985). Pri- mary production (estimated from three-week accumulations of autotrophic biomass on arti- ficial substrates) ranged from 0.22 to 0.58 mg C/m?/hr in summer-autumn 1979, but it de- clined to 0.08 to 0.28 mg C/m’/hr after peak discharge in summer 1980, apparently due to phosphorus-limited growth. Oscillatoria spe- cies are generally abundant only in areas of nutrient enrichment (VanLandingham 1982). The decrease in density of this taxon in sum- mer 1980 may have been attributable to slower growth in a phosphorus-deficient envi- ronment. 600 GREAT BASIN NATURALIST Vol. 46, No. 4 Lyngbya spp. Cocconeis placentula eS ea =e —a Gomphonema spp. Cymbella sinuata Synedra acus Cymbella minuta Ceratonels arcus Calothrix spp. Nitzschia spp. Cymbella microcephala LOG, INDIVIDUALS / CM2 S Cladophora spp. Fragilaria construens .|—-—= nm” == Achnanthes minutissima , Fragilaria crotonensis PEE ee ae Oscillatoria spp. Navicula spp. Synedra rumpens Amphipleura pellucida m= A ee —<==s — > — Ne) alee tenes) ea oe) JJASON JJASON JJASON JJASON 1979 1980 1979 1980 Fig. 2. Mean population densities (n = 6) of common benthic algae on streambed cobble of Convict Creek. Densities were normalized by natural logarithmic transformation prior to calculating means and standard errors. Seasonal population trends identified in the text are based on mean densities differing by at least the sum of the standard errors of the means. October 1986 TABLE 2. Benthic insects of Convict Creek. EPHEMEROPTERA Siphlonuridae + Ameletus sp. Baetidae Baetis devinctus Traver Baetis tricaudatus Dodds + Baetis spp. + Callibaetis pacificus Seeman Heptageniidae Cinygmula sp. + Epeorus dulciana (McDunnough) * — Epeorus longimanus (Eaton) Tronodes lepidus Traver Rhithrogena sp. Ephemerellidae * Caudatella heterocaudata (McDunnough) * — Caudatella hystrix (Traver) Drunella doddsi (Needham) * — Drunella flavilinea (McDunnough) Drunella grandis (Eaton) Drunella pelosa (Mayo) Ephemerella infrequens McDunnough Serratella tibialis (McDunnough) Tricorythidae Tricorythodes minutus Traver Leptophlebiidae Paraleptophlebia pallipes (Hagen) * PLECOPTERA Pteronarcidae Pteronarcys princeps Banks Pteronarcella regularis Hagen * * Peltoperlidae Yoraperla brevis (Banks) Nemouridae Malenka sp., probably californica (Claassen) * Zapada cinctipes (Banks) Perlidae Calineuria californica (Banks) Doroneuria baumanni Stark & Gaufin Hesperoperla pacifica (Banks) * * Chloroperlidae * Sweltsa sp., probably pacifica (Banks) HEMIPTERA Belostomatidae Belostoma bakeri Montd. Corixidae Corisella inscripta Uhler Sigara washingtonensis Hungerford MEGALOPTERA Corydalidae Orohermes crepusculus (Chandler) TRICHOPTERA Philopotamidae * Wormaldia sp., probably gabriella Banks Polycentropodidae + Polycentropus sp. Hydropsychidae Arctopsyche grandis (Banks) * LELAND ETAL.: SIERRA NEVADA AQUATIC INSECTS Table 2 continued. * Hydropsyche oslari (Banks) Rhyacophilidae Rhyacophila acropedes Banks Rhyacophila angelita Banks Rhyacophila vaccua Milne + Rhyacophila sp. * * Glossosomatidae Agapetus taho Ross Glossosoma califica Denning Hydroptilidae + Agraylea saltesa Ross + Hydroptila spp. + Oxyethira sp. * Brachycentridae Brachycentrus americanus (Banks) + © Micrasema sp. * Lepidostomatidae Lepidostoma cascadense (Milne) Lepidostoma rayneri Ross + Lepidostoma sp. * * Limnephilidae + Dicosmoecus atripes (Hagen) + Neophylax sp. Leptoceridae Triaenodes sp. LEPIDOPTERA Pyralidae undetermined genus COLEOPTERA Dytiscidae Agabus obsoletus (LeConte) Bidessus sp. Deronectes striatellus (LeConte) Deronectes sp. Hydroporus sp. Laccophilus decipiens LeConte Rhantus binotatus Harris Hydrophilidae Ametor scabrosus (Horn) Berosus sp. Laccobius ellipticus LeConte undetermined genus Hydraenidae Hydraena vandykei d Orchymont Ochthebius interruptus LeConte Elmidae Cleptelmis addenda (Fall) + Lara avara LeConte Narpus sp. Optioservus divergens (LeConte) Optioservus quadrimaculatus (Horn) Zaitzevia parvula Horn DIPTERA Deuterophlebiidae * Deuterophlebia nielsoni Kennedy Blephariceridae Agathon comstocki Kellogg Tipulidae * — Antocha monticola Alexander 601 602 Table 2 continued. Cryptolabis sp. + Dicranota sp. Gonomyia sp. + Hexatoma sp. Pedicia sp. Tipula spp. + Psychodidae Pericoma sp. Maruina sp. + Ceratopogonidae Palpomyia or Bezzia spp. Simuliidae * — Simulium aureum Fries * — Simulium arcticum Malloch + Simulium spp. Prosimulium dicum Dyar & Shannon Chironomidae Tanypodinae Tanypus sp. Thienemannimyia group Diamesinae + Diamesa sp. Diamesa latitarsus (Goetghebuer) Pagastia sp. Prodiamesa sp. Orthocladiinae Chaetocladius spp. Corynoneura spp. + Cricotopus spp. Cricotopus bicinctus group + Eukiefferiella spp. Eukiefferiella bavarica group Eukiefferiella brevicalcar group Eukiefferiella potthasti group Krenosmittia sp. Nanocladius sp. Orthocladius spp. Orthocladius (Euorthocladius) sp. Pseudorthocladius sp. Thienemanniella sp. 1 Thienemanniella spp. Chironominae Dicrotendipes sp. Polypedilum laetum (Meigen) Polypedilum (Tripodura) sp. Polypedilum spp. Pseudochironomus sp. Empididae Chelifera sp. Te Wiedemannia sp. undetermined genus Muscidae Limnophora sp. * Species reported by Kennedy (1967) +Genus or family reported by Kennedy (1967) Common diatoms in Convict Creek with population maxima during late summer-—early autumn were Cocconeis placentula lineata, Cymbella microcephala, Cymbella sinuata, GREAT BASIN NATURALIST Vol. 46, No. 4 Nitzschia frustulum, and N. palea. The green algae Mougeotia spp. and Cladophora spp. were also abundant at this time. Cymbella microcephala and C. sinuata were early colo- nizers in late summer, whereas the other taxa were later successional species. Autumn: Common taxa with population maxima in autumn were the diatoms Cy- clotella comta, Navicula spp. (principally cryptocephala, rhynchocephala and arven- sis), and Amphipleura pellucida, the green alga Spirogyra spp. (principally grantiana), and the blue-green Calothrix spp. By mid- October 1979 Spirogyra spp. was a co-domi- nant but in 1980 it was never very abundant, whereas Lyngbya spp., Achnanthes minutis- sima, Cocconeis placentula lineata, Cymbella microcephala, C. sinuata, Fragilaria con- struens, Navicula spp. and Synedra rumpens were abundant both years. By mid-November A. minutissima, F. construens, and F. cro- tonensis were the most abundant diatoms. Calothrix spp. was abundant in late autumn 1979 but not in 1980. Spirogyra spp. was the principal green alga in late autumn 1979, whereas Cladophora spp. was more abundant in 1980. Benthic Insects COMMUNITY COMPOSITION.—Comprehen- sive lists of benthic insects exist for several streams of the eastern Sierra Nevada at ap- proximately the same altitude as Convict Creek. Three streams in the Truckee River Basin, Sagehen Creek (Gard 1961, Siegfried and Knight 1975), Berry Creek (Siegfried and Knight 1975), and Prosser Creek (Needham and Usinger 1956), have faunal compositions (comparing genera) similar to that of Convict Creek (Convict Creek taxa are listed in Table 2). However, the Truckee River Basin streams have higher densities of the eph- emeropterans Cinygmula and Rhithrogena, their species of Ephemerellidae are mostly different, and we did not collect the ple- copteran families Leuctridae, Capniidae, and Perlodidae (but Kennedy [1967] did report Leuctridae and Capniidae in Convict Creek). Maciolek and Tunzi (1968) listed the fauna of Laurel Creek, which is near Convict Creek but at a higher elevation. Fewer taxa are present in Laurel Creek, but benthic insect compositions of the two streams are similar. Kennedy (1967) listed the benthic insects of October 1986 Convict Creek during 1961-1963 but pre- sented little information on population densi- ties; most of the common taxa we observed in 1977-1980 were also found by Kennedy (Table 2). Some apparent differences in the stream fauna between the early 1960s and late 1970s are due to difficulties in identifying im- mature stages and differences in sampling method. Kennedy (1967) collected exten- sively only from riffles, whereas midriffle ben- thos and drift were sampled in the present study. Notable differences between our spe- cies list and that of Kennedy (1967) are (1) that Optioservus divergens, Ironodes lepidus, Doroneuria baumanni, and Malenka (califor- nica?) are common taxa now but were not reported in 1961-1963 and (2) that Kennedy (1967) found three winter stoneflies (Leuctri- dae and Capniidae) not observed in 1977— 1980. These stoneflies typically develop in winter-spring and may have been missed in our sampling program. SEASONALITY AND LIFE HISTORIES OF ComM- MON TAXA.—Mean population densities of 28 common benthic insects in Convict Creek are given in Fig. 3. The taxa are ordered by their location on the primary axis in ordination space (detrended correspondence analysis— see Leland et al. 1986). Seasonality of taxa is emphasized in the ordering. Data on popula- tion densities (which emphasize early and middle instars) are supplemented with obser- vations on the occurrences of late instars, pu- pae, and adults and on length-frequency data to determine life histories (Fig. 4). Late spring (400 to 1,200 degree-days): Taxa most abundant during May and June are considered late-spring fauna. Species in this assemblage include ephemeropterans (Epe- orus longimanus, Drunella flavilinea, and Caudatella heterocaudata), plecopterans (Calineuria californica, Doroneuria bau- manni, and Pteronarcys princeps), and dipterans (Cryptolabis sp. and Palpomyia spp.). The three ephemeropterans occur as middle to late instars in late spring. Their major period of growth and development is early to late spring, and adults appear during late spring and summer (Fig. 4). Nymphs are _ not common in autumn, so most early instars _ must first appear during winter (as early as December for D. flavilinea) or early spring. \ Caudatella heterocaudata develops some- _ what later than the other two species, and LELAND ETAL.: SIERRA NEVADA AQUATIC INSECTS 603 most individuals emerge later in the summer. Slow growth or a diapause in autumn and early winter for eggs and early instars, fol- lowed by a rapid development in spring, is suggested for all three species. This corre- sponds to the “fast seasonal” type of life history described by Hynes (1970). Similar life histo- ries have been reported for D. flavilinea in Idaho (Andrews and Minshall 1979) and E. longimanus in Alberta (Hartland-Row 1964). Early instars of the plecopterans C. califor- nica, D. baumanni, and P. princeps are most abundant in late spring. These species typi- cally have a two- or three-year life cycle, and egg development requires as long as 8 to 10 months (Siegfried and Knight 1977, Barton 1980). Hatching of C. californica and D. bau- manni extends over several months since early instars are also present in autumn. Rates of development of all three species are proba- bly highest during late spring and summer (Heiman and Knight 1975, Siegfried and Knight 1977). Cryptolabis sp. is the most abundant tip- ulid in Convict Creek and is especially preva- lent in late spring. Development occurs pri- marily between August and May and individuals overwinter as middle to late in- stars; pupation occurs in July. The relative scarcity of early instars indicates that at this stage Cryptolabis sp. may be hyporheic or inhabit regions of slower current. Two other common dipterans, Deuterophlebia nielsoni and Palpomyia spp., are most abundant dur- ing late spring, but their population densities are never high. Deuterophlebia nielsoni is multivoltine (Kennedy 1967); however, we did not observe larvae of this species after September. Summer (1,200 to 2,200 degree-days ): Pop- ulation densities of many common taxa in Convict Creek are lowest in early summer. This is generally true for temperate streams (Hynes 1970) and is due at least in part to the lag between spring adult emergence and hatch of the next generation. Nighttime air temperatures generally remain above freez- ing during late spring and early summer, and emergence of most species occurs at this time (see Fig. 4). Scouring of the streambed during periods of high discharge in late spring— summer also appears to contribute to popula- tion declines. Other authors (Gaufin 1959, Canton and Ward 1978, Minshall 1981) have 604 GREAT BASIN NATURALIST Vol. 46, No. 4 Epeorus longimanus Arctopsyche grandis = — — 1b>- —> Pp a Drunella flavilinea Glossosoma califica Se Pa «a rng Cryptolabis sp. Simulium spp. a —— PF PP pm— Calineuria californica Rhyacophila vaccua =—— —_— e= Gz —_ji «a —_> a Caudatella heterocaudata lronodes lepidus ae ——— =e C25) e=a aw Doroneuria baumanni Cleptelmis addenda =, = ee Ga =a @ — Pe Pteronarcys princeps Baetis spp. -_—-— @ 8 BD Palpomyla spp. Epeorus dulciana —_—_ = ee Ee 6 ee Sweltsa (pacifica?) Brachycentrus americanus Ea = Ss a —— @ -@ >-—Z LOG, INDIVIDUALS / 0.1 M? pe Optioservus divergens Lepidostoma sp. maeQtmww<«#<«—~< Malenka (californica?) Paraleptophlebia pallipes Rhyacophila acropedes Antocha monticola = — asf @ — Chironomidae Pericoma Sp. Magen»vpea —<—« - Hydropsyche oslari Ephemerella infrequens @Teei»nu#4»nwez—= <« @~<« JASON JJASON JASON MJJASON JJASON JJASON JJASON MJJASON 1977 1978 1979 1980 1977 1978 1979 1980 Fig. 3. Mean population densities (n = 3) of common benthic insects in riffle areas of Convict Creek. Densities were normalized by natural logarithmic transformation prior to calculating means and standard errors. Seasonal population trends identified in the text are based on mean densities differing by at least the sum of the standard errors of the means. October 1986 Epeorus longimanus A Drunella flavilinea ------- --—}]}— N A Cryptolabis sp. le—————————_~—Wse=—-=------ ———<$ $m _ P Me ee eg LS Cre Calineuria californica NH ——————— [Xo Earners Soe, Caudatella heterocaudata ———— A E Doroneuria baumanni NH —————— A eee ewe eee ee = K&——— Dicranota sp. L JRE, ad ae ee ee ee are Malenka (californica?) N --H A =--Ee==-= Rhyacophila acropedes Sd H May June July Aug Sept Oct Nov Dec N=Nymph_ A = Adult L = Larva Re=\Pupayeeiah) ih ae LELAND ET AL.: SIERRA NEVADA AQUATIC INSECTS Occurs commonly Occurs rarely (immature stages) or 605 Arctopsyche grandis L ee A oe Glossosoma califica lL ———__----_. ----H- PS a A lo Simulium spp. L Em ——— cE H A —E— =--E—— Rrvacophila vaccua Z |] P SEE WN) SS SSSESGSS5S505 lronodes lepidus N ——-------- = A -----= [E9505 Baetis spp. N ——------- _——$S $< _ A —E— —(2— b2 m pr 1 I m ' 1 ' 1 f 1 ' Paraleptophlebia pallipes N --- =H". A ---E-————- Antocha monticola l-——----. OO ——————— SS P Se A eeeoe ss Oe Ephemerella infrequens N =e} A =—E— Micrasema sp. a I OS t—--——>—$—$—$$—>—$—$—— A —— eee May June July Aug Sept Oct Nov Dec H = Major hatch period E = Major emergence period probable occurrence (adults) Fig. 4. Generalized life histories for common benthic insects of Convict Creek. attributed low summer populations in streams of the western United States to the same fac- tors. The plecopteran Malenka (californica?) completes most of its development during summer and autumn. The main cohort emerges in October, but some individuals overwinter as late instars. The trichopteran Arctopsyche grandis is a common summer species; it is also the largest caddisfly in riffles and is thus an important component of the summer biomass. Arctopsyche grandis devel- ops mostly between July and early Septem- ber. A rapid summer development has also been reported in Idaho (Cuffney and Minshall 1981) and coastal California (Furnish 1979). The uniform size of individuals on each sam- pling date in Convict Creek suggests a uni- voltine life cycle; however, Smith (1968) found the species to be semivoltine in Idaho. Hydropsyche oslari is also most abundant (as early instars) in late summer, but most growth occurs after that of A. grandis. Growth of H. oslari begins about September and continues through autumn. Rhyacophila acropedes is another abundant summer trichopteran. Several bivoltine and multivoltine taxa have summer cohorts. At least two species of the ephemeropteran genus Baetis develop between June and September. The dipterans 606 Simulium spp. and Chironomidae are usually most abundant from August through Novem- ber. These taxa include a large number of species (see Table 2 for a list of the Chironomi- dae and Kennedy 1967 for a list of Simulium species.) Because of our inability to identify early instars, these taxa were enumerated by family; therefore, life histories cannot be de- scribed. The family Simuliidae was repre- sented only by Simuliwm spp. in riffle sam- ples, but a few adult Prosimulium dicum appeared in drift samples. A major emergence of Simulium spp. occurred in September 1980, but emergence was not as synchronous in other years. Autumn (2,200 to 3,100 degree-days ): Peak densities of many common taxa in Convict Creek occur in autumn. The community con- sists largely of early instars of “slow seasonal” species (Hynes 1970), insects that hatch quickly, develop partially during autumn, and emerge in spring (Fig. 4). Although some taxa are probably detritivores (for example, Para- leptophlebia pallipes and Lepidostoma spp.) and may have life histories timed to periods of leaf fall (Anderson and Cummins 1979), most are generalists and relative amounts of de- tritus and algae ingested vary seasonally (Chapman and Demory 1963, Gray and Ward 1979). Food quality probably influences life history patterns, but a more significant factor in Convict Creek may be that autumn is a period when the stream is typically at or near base flow. The moderate stream velocities may enhance survival of early instars by providing suitable substratum. Ephemerella infrequens and Epeorus dul- ciana are common ephemeropterans in Con- vict Creek with autumn and early spring de- velopment. Ephemerella infrequens has a similar life history in Colorado (Ward and Berner 1980). Ironodes lepidus is less abun- dant in Convict Creek, but our limited data indicate a similar life history. Paralep- tophlebia pallipes grows most rapidly in au- tumn, but hatching is apparently delayed in some individuals since recruitment continues through autumn. Low numbers of these taxa in June indicate a spring emergence. Hydropsyche oslari is the dominant tri- chopteran by late autumn. Most individuals overwinter as third to fifth instars. This spe- cies overwinters at an earlier stage (first and second instars) in Montana (Hauer and Stan- GREAT BASIN NATURALIST Vol. 46, No. 4 ford 1982). Many other trichopterans also have peak abundances in autumn. Early in- stars of Lepidostoma spp. are common in riffles in autumn, but the taxon is rarely found there in spring. Larvae apparently move into areas of slower current in later developmental stages. Glossosoma califica and Micrasema sp. develop slowly during autumn, overwin- ter as early to middle instars, and pupate in late spring-summer. Brachycentrus ameri- canus and Neophylax sp. develop to middle instars by December, but emergence does not occur until late spring or summer. Rhya- cophila vaccua hatches during summer and autumn, overwinters primarily as middle in- stars, and emerges in early summer. Chironomidae are abundant throughout autumn. Two other dipterans, Antocha mon- ticola and Pericoma sp., also are at their maxi- mum densities in autumn but neither is ever abundant. Antocha monticola completes most of its development in late summer and au- tumn. Although larvae were uncommon in riffles in summer, a large number of adults in August 1980 emergence samples indicates that emergence occurs througheut the summer. Late winter/early spring (0 to 400 degree- days): Samples of benthic insects were not taken from January through April. This is ap- parently an active period for many species. Many late spring species must develop rapidly after overwintering as eggs or early instars. The autumn taxa are present primar- ily as later instars, and many of these emerge in the spring. “Aseasonal” taxa: Some taxa are abundant throughout the snow-free months of the year, and population densities do not show strong seasonal trends. Baetis spp. (principally B. devinctus and B. tricaudatus) is the most abundant ephemeropteran taxon most of the year. Both species are bivoltine, with emer- gences in May-June and September. The au- tumn cohort is larger. Approximately 1,400 degree-days accumulate during development of a generation. The elmid Optioservus diver- gens appears to have a two-year life cycle (as was also described for O. ampliatus in eastern Canada by Le Sage and Harper 1976). Peak numbers of early instars occur between Au- gust and October, and most of this cohort reaches middle-instar stages by December. Plecopterans in Convict Creek show less seasonal variation in abundance than do other October 1986 = 2000 Ponape = 1600 4 e 1200 4 Se: at pee = 0 mo einen neo ra J JASONDJASONJASONDMJ JASON = UST 1980 Fig. 5. Total densities (n = 3) of benthic insects in riffle areas of Convict Creek. Densities were normalized by natural logarithmic transformation prior to calculating means and standard errors. ISVS Wrs orders of benthic insects. Calineuria califor- nia, Doroneuria baumanni, and Pteronarcys princeps all are semivoltine. However, the population density of Sweltsa sp., which is probably univoltine, did not vary much ei- ther. This may be due to the presence of more than one species. YEARLY VARIATION IN RELATIVE ABUN- DANCES OF TAXA.—Population densities of many common benthic insects in Convict Creek varied substantially among years. Be- cause our sampling interval was wide and sea- sonal changes in population density were pro- nounced, only major among-year differences can be defined with certainty. The total number of individuals of all ben- thic insect taxa was unusually high in late autumn 1977 (Fig. 5), a year of low discharge during all seasons and relatively high summer water temperatures (Fig. 1). Population den- sities of Baetis spp., Paraleptophlebia pal- lipes, Epeorus dulciana, and Antocha monti- cola, which hatch in autumn, were highest in late autumn 1977. Reproduction and survival of early instars of these taxa were apparently favored by the relatively mild hydrologic and climatic conditions. Except for P. pallipes, these taxa are all clinging, epilithic grazers (Merritt and Cummins 1984) and thus benefit from nondisruptive stream flows. Pericoma sp., which typically resides in fine sediments or algal mats (Usinger 1956, Merritt and Cum- mins 1984), was also common in riffles in au- tumn 1977. Ephemerella infrequens , another autumn-hatching ephemeropteran, was less abundant in 1977 than other years. Sampling of benthic insects was not ini- tiated until August in 1978, so any effects of the severe winter conditions on winter and spring fauna could not be observed. The high LELAND ETAL.: SIERRA NEVADA AQUATIC INSECTS 607 discharge accompanying Hurricane Norman in early autumn 1978 probably caused the low autumn densities of Glossosoma califica, Paraleptophlebia pallipes, Epeorus dulciana, Cptioservus divergens , and some of the larger plecopterans. The severe winter conditions of early 1978 may have contributed to the popu- lation declines of the semivoltine O. diver- gens, Pteronercys princeps, and Doroneuria baumanni. Severe winters appear to result in a high mortality of benthic insects. Reimers (1957) found that populations of ephemerop- terans and elmid beetles in Convict Creek are reduced more by unusually cold winters than are populations of dipterans and trichopter- ans. Winter air temperatures remained low longer (through March) in 1980 than in other years, but a heavy snow cover and unusually high winter baseflow prevented much anchor ice formation. The late spring-summer period of high discharge began late and continued through early August. Summer water temper- atures were generally lower than in 1977 and 1979. Several taxa, including most plecopter- ans, were more abundant than in other years. These included Doroneuria baumanni, Cal- ineuria californica, Pteronarcys princeps, Rhyacophila vaccua, and possibly Micrasema sp., Palpomyia spp., Malenka (californica ?), and Sweltsa pacifica. The greater abundances of these taxa in autumn 1980 may be the result of a more effective sorting of streambed sedi- ments due to the unusually high discharge in late spring-summer (DeMarch 1976). The ab- sence of anchor ice in winter may have con- tributed to the increased survival rate of the 1979 cohorts of D. baumanni, C. californica, and P. princeps. Benthic Insect Production The total annual production of herbivorous and detritivorous benthic insects in Convict Creek riffle areas was an estimated 3.9 g/m” (dry weight) (Table 3). Not included in this estimate is the contribution of oligochaetes and molluscs, which is approximately 10%— 20% of the mean annual standing stock. The total annual production of benthic insect predators was an estimated 1.7 g/m’. These annual production estimates compare favor- ably with the values of Krueger and Waters (1983) for the Caribou River and Blackhoof River, streams in Minnesota with similar sub- 608 GREAT BASIN NATURALIST Vol. 46, No. 4 TABLE 3. Annual production, mean annual standing stock, and cohort production interval (CPI) for the more abundant benthic insects in 1979-80. Annual production Mean annual Estimated (g/m? dry wt.) standing stock CPI (days) Estimate 95% C.1. (g/m? dry wt.) PLECOPTERA Calineuria californica 021 .0O1— .041 .006 780 Doroneuria baumanni PBS) .073— .394 .075 780 Sweltsa (pacifica ?) 012 .003— .020 .002 300 EPHEMEROPTERA Drunella flavilinea 104 .O70— .139 014 210 Ephemerella infrequens JUG .082— .152 018 270 Paraleptophlebia pallipes 041 .020— .062 .007 300 Baetis spp. .937 .721-1.152 079 150 Epeorus dulciana py Wis) .061— .168 014 300 Epeorus longimanus .158 .079— .237 .009 150 TRICHOPTERA Hydropsyche oslari 5006 .320— .792 .057 300 Arctopsyche grandis 1.417 .903-1.932 150 300 Rhyacophila vaccua .261 .128— .393 .034 300 Glossosoma califica .087 .025— .149 014 300 Brachycentrus americanus .056 .020— .092 .008 300 COLEOPTERA Optioservus divergens .128 .098— .158 .069 660 Cleptelmis addenda .003 .001— .005 .002 660 DIPTERA Simulium spp. .099 .049— .149 007 120 Chironomidae 12) _- 012 — strate but higher discharge and nutrient con- centrations. Annual production of herbivores/ detritivores in the two streams was 4.6 g/m° and 6.3 g/m’ (corrected to dry weight), respec- tively, and the annual production of predators was 0.94 g/m’ and 1.1 g/m’. There are several potential sources of error in the production estimates. Some large in- sects, such as Pteronarcys princeps and cer- tain tipulids, may have contributed substan- tially to production in Convict Creek, but these individuals were not abundant enough to estimate production with confidence. Mul- tiple-species populations (Baetis spp., Sim- ulium spp.) and sexual dimorphism may have yielded overestimations because the size fre- quency method assumes that all individuals can reach the largest size class. Although non- linear growth has been shown not to affect production estimates severely (Hamilton 1969, Cushman et al. 1978), taxa such as Arc- topsyche grandis are less accurately esti- mated. Emigration from the sampled area (riffles) during some portion of the life cycle, such as with Paraleptophlebia pallipes (An- derson and Lehmkuhl 1968), also would give an inaccurate estimate. Finally, the produc- tion estimates may be biased due to unequal sampling intervals. Stream conditions during the study period (mid-August 1979 to mid-June 1980) were not extreme. There was little anchor ice during winter, and the annual high discharge in late spring—summer was delayed. Consequently, insect mortalities were probably not unusu- ally high. With most taxa hatching during summer and autumn, the period of highest mortality (early instars) was well sampled. Ex- ceptions were Epeorus longimanus and Drunella flavilinea, which hatch in late autumn—winter. Phenology This investigation was not designed as a watershed phenological study, but rather a discussion of major recurring climatic and hy- drologic events that influence life history pat- terns in Convict Creek. Winter is a harsh season lasting approximately four months (December through mid-March), when the stream is at or near baseflow, nighttime water temperatures are near freezing, and anchor ice forms occasionally. The dominant winter periphyton are diatoms (Achnanthes, Fragi- October 1986 laria, and Gomphonema) and primary pro- duction is low. It is a period of activity for many benthic insects, with taxa that repro- duce in autumn present primarily as later in- stars. Unusually cold winters cause a high but selective mortality of benthic insects (Reimers 1957). Despite low water temperatures, brown trout (Salmo trutta L.) feed throughout the season (Jenkins 1969). The water temperature begins to rise about mid-March and increases through June or July. Snowmelt in higher areas of Convict Creek Basin causes a substantial increase in discharge, beginning in May or June, which lasts for one to two months. In early summer a dense riparian canopy dominated by willows and quaking aspen develops in some stream reaches. The high discharge in late spring— summer is accompanied by declines in stand- ing crops of periphyton and benthic insects. Succession in the periphyton community is interrupted by scouring of the streambed in late spring and early summer. Relative abun- dances of periphyton taxa in late summer and autumn appear to reflect the magnitude of discharge in late spring-summer and the availability of nutrients (see also Leland and Carter 1985). When stream discharge in late spring was exceptionally high and primary productivity relatively low, the community was dominated by early-colonizing diatoms during summer; diatoms and _ filamentous green and blue-green algae were all abundant in autumn. When stream discharge in late spring was only moderately high and the rate of production relatively high, blue-green al- gae dominated in summer and remained abundant in autumn. There is a total emergence (hence repro- ductive) period of at least seven months (April—October) for benthic insects, but most species emerge in late spring and early sum- mer. Low population densities during late _ spring and early summer are thus due at least _ in part to the lag between emergence and hatch of the next generation. Scouring of the streambed also appears to contribute to the _ population declines. Summer is the only season when nighttime air temperatures consistently remain above | freezing; stream discharge declines progres- ) sively during this period. Later successional _ species of periphyton are relatively more _ abundant in summer than in late spring, and LELAND ETAL.: SIERRA NEVADA AQUATIC INSECTS 609 primary productivity is at its annual maxi- mum. Some bivoltine and multivoltine ben- thic insects have summer cohorts and grow more rapidly than do individuals of the au- tumn cohorts. Autumn is a season of declining water tem- perature and stream discharge, and baseflow is reached in October. Leaf-fall occurs from mid-September through October, with leaf litter in the stream most abundant in October. Between-year differences in periphyton com- position were large, apparently due to envi- ronmental factors influencing the relative abundances of taxa at earlier successional stages (see also Leland and Carter 1986). Most slow-seasonal and multivoltine insects hatch during autumn, so this is a period of high abundance of early instars. Some of these are detritivorous (for example Paraleptophlebia pallipes and Lepidostoma spp.) and may have life histories timed to the autumn abundance of leaf litter. ACKNOWLEDGMENTS The contributions of Charles B. Doherty, Thomas L. Dudley, and William H. Peeler in fieldwork and in the identification and enu- meration of benthic insects are gratefully ac- knowledged. The identifications were con- firmed whenever possible using museum specimens of the California Academy of Sci- ences, San Jose State University, and Oregon State University. Rearing of specimens was done in some instances to aid in the identifica- tions. Larry J. Tilley and Suzanne F. Muzzio verified identifications of the Chironomidae. All the above individuals are or were em- ployees of the U.S. Geological Survey at the time the work was done. Sam L. VanLanding- ham, consulting biologist, did early identifica- tions of the green algae and blue-green algae. The program for computing secondary pro- duction was obtained from Charles C. Krueger (Department of Entomology, Fish- eries and Wildlife, University of Minnesota). Permission to conduct studies in Convict Creek was granted by the governing board of the Sierra Nevada Aquatic Research Labora- tory, University of California at Santa Bar- bara. Scott D. Cooper, Frank F. Hooper, Allen W. Knight, and Keith V. 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Marshall, eds., Advances in Ephemeroptera biol- ogy. Plenum Press, New York. DIATOM FLORA OF COWBOY HOT SPRING, MONO COUNTY, CALIFORNIA Laura Ekins’ and Samuel R. Rushforth! ABSTRACT. —The diatom flora of Cowboy Hot Spring, Mono County, California, was studied. Two habitats, one at 37C and one at 41C, were examined. Fifty-six taxa were identified from our samples. These taxa were mostly broadly distributed forms, and no endemic species were encountered. The dominant taxon was Nitzschia frustulum, followed by Achnanthes gibberula, Achnanthes exigua, Nitzschia hantzschiana, and Navicula cincta. Interest in natural thermal waters has in- creased substantially during the past few years. Early studies of these environments were generally concerned with Cyanophyta or the faunas of such habitats (Edwards 1868, Davis 1897, Tilden 1897, 1898). These au- thors often expressed surprise that organisms could exist in thermal environments and were interested in the upper temperature toler- ance of thermophilic species. Diatoms were not studied systematically until some time later (Lacsny 1912, Oestrup 1918, Strom 1921, Famin 1933, Springer 1930), when they were found to be common in hot springs. More extensive studies were be- gun in the 1940s (Negoro 1940, Emoto and Hirose 1940, Emoto and Yoneda 1941, Yoneda 1942a, 1942b) and have continued to the present time (Whitford 1956, Yoneda 1962, Thomas and Gonzalves 1965a, 1965b, 1966a-e, Biebl and Kusel-Fetzmann 1966, and others). Diatoms also have recently been studied in several western North American springs (Kaczmarska and Rushforth 1983, St. Clair and Rushforth 1977, Stockner 1967a, 1967b, 1968, Rushforth et al. 1986). Cowboy Hot Spring of the Mono Basin thermal area of eastern California is one of many thermal springs in the Basin and Range Geological Province (Great Basin). This province occurs east of the Sierra Nevada Mountains, between the Snake River and the Mojave Desert, and extends across Nevada into western Utah. The Great Basin is characterized by a thin- ning of the earth's crust, more than 200 north/ south-oriented mountain ranges with associ- ated valleys (Nelson 1981), an abundance of thermal springs, and a cold desert climate. These features appear to be a result of several geological phenomena, including the contin- ued spreading of the American Plate through the center of the Great Basin, past subduction of the Pacific Plate under the American Plate (Miller 1983), and the uplift of the Sierra Ne- vada. Thermal springs have been characterized as waters with a temperature 6—9C greater than the mean annual air temperature of the adja- cent area (Tarbuck and Lutgens 1984). A more detailed classification of thermal springs is that of Elenkin (Kol 1932), where waters be- low 15C were termed hypothermal, between 15C and 30C as mesothermal, and above 30C as euthermal. Water temperature in thermal springs generally remains quite constant be- cause the source water is continuously heated by tectonic events. We have studied the diatom flora of Cow- boy Hot Spring to compare the flora with that of other thermal waters of western North America. This euthermal spring is of particu- lar interest since its temperature of 41C is near the upper temperature limit for eukary- otic organisms (Ruttner 1963). The present paper lists and illustrates all known diatom taxa in Cowboy Hot Spring. SITE DESCRIPTION AND COLLECTIONS Cowboy Hot Spring is at 37° 38.6’ N lati- tude and 118° 45.45’ W longitude in Mono County, California. A concrete tub has been built around the source water, which is 41C. A runoff stream from the source contains water at 37C 3 m from the tub. Water flow is 150 ‘Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602 612 eee October 1986 l/min, and the total dissolved solids are 150 ppm (California Geol. Map 1980). All sampling was done 26 August 1981. Samples were taken from both the concrete tub and the runoff stream. Composite samples were collected by scraping the sides of the concrete tub and obtaining visible algae and placing these collections into vials. Stones and submerged wood in the stream were scraped, and these scrapings, together with visible al- gae and small amounts of sediment, were placed into vials. Composite samples of this type were obtained to insure that as many taxa as possible would be collected, since the pri- mary thrust of this study was a floristic survey. All samples were stored at air temperature and returned to our laboratory at Brigham Young University. METHODS Diatoms were cleared with nitric acid fol- lowing standard methods using boiling nitric acid (St. Clair and Rushforth 1977), and strewn mounts using Naphrax high resolution mount- ing medium were prepared. Slides were stud- ied using Zeiss RA microscopes with Nomarski and bright field illumination. Pho- tographs of each taxon were obtained using Nikon AFM photomicrographic equipment. A minimum of 500 frustules was counted on each slide to calculate the relative density for each taxon in each sample. Species diversity of each sample was measured by calculating the Shannon-Weaver index (Shannon and Weaver 1963, Margalef 1958, Patten 1962). Permanent diatom slides are in the collec- tions at Brigham Young University. RESULTS AND DISCUSSION Fifty-six diatom taxa in 20 genera were identified in samples from Cowboy Hot Spring. Most of the taxa encountered are cos- mopolitan, eurythermal forms and none were endemics. Kaczmarska and Rushforth (1983) found 136 diatom taxa in Blue Lake Warm Spring, in the Great Basin, a somewhat cooler spring at 29C than Cowboy Hot Spring, which might account for much of the difference in species diversity between the two sites. Twelve taxa in Cowboy Hot Spring had an _ average relative density above 1.0. Nitzschia | frustulum (Kuetz.) Grun. was the most abun- | EXKINS, RUSHFORTH: CALIFORNIA DIATOMS 613 dant, with an average density of 31%, fol- lowed by Achnanthes gibberula Grun. (10.3%), Achnanthes exigua Grun. (10.1%), Nitzschia hantzschiana Rabh. (9.3%), Navic- ula cincta Ralfs (8.6%), Anomoeoneis sphaerophora (Ehr.) Pftiz. (6.8%), Nitzschia communis Rabh. (4.1%), Achnanthes exigua var. 1 (3.1%), Nitzschia valdecostata Lange- Bert. and Simon. (2.2%), Navicula crypto- cephala var. veneta (Kuetz.) Rabh. (1.6%), Rhopalodia operculata (C.A.Ag.) Hakan. (1.6%), and Denticula elegans Kuetz. (1.2%). Kaczmarska and Rushforth (1983) found 17 taxa with importance values above 1.0 in Blue Lake Spring. Three of these also occurred in Cowboy Hot Spring, although none of them had average densities above 1.0%. Stockner (1967a) found 12 species common in waters above 35C in Yellowstone National Park. Fight of these taxa were also found in Cowboy Hot Spring. These included Achnanthes gib- berula Grun., Achnanthes lanceolata Breb.., Amphora coffeaeformis (Ag.) Kuetz., Dentic- ula elegans Kuetz., Gomphonema parvulum Kuetz., Navicula cincta Ralfs, Pinnularia mi- crostauron (Ehr.) Cl., and Rhopalodia gib- berula (Ehr.) O. Muell. None of the important taxa in Cowboy Hot Spring was confined to the stream or tub. However, several taxa did show a trend to- ward being restricted to one habitat or the other. Those reaching maximum develop- ment in the concrete tub were Cocconeis pla- centula var. lineata (Ehr.) V. H., Denticula elegans Kuetz., Epithemia argus (Ehr.) Kuetz., Fragilaria construens var. venter (Ehr.) Grun., and Tabellaria quadrisepta Knuds. Those with maximum development in the stream were Achnanthes linearis Grun.., Navicula confervacea var. peregrina (W.Sm.) Grun., Nitzschia microcephala Grun., and Pinnularia appendiculata (Ag.) Cl. The Shan- non-Weaver index value for the concrete tub was 3.67, and the stream value was 3.30. A taxonomic section follows with a descrip- tion of each taxon. A short discussion is in- cluded where appropriate. TAXONOMIC SECTION Achnanthes exigua Grun., Figs. 1-7. Valves 12.5-21.5 wm long by 5-6 pm wide; raphe valve striae 25-28 in 10 wm; rapheless valve striae 24-26 in 10 pm. This taxon oc- 614 GREAT BASIN NATURALIST Vol. 46, No. 4 Figs. 1-34. Diatom species: 1-7, Achnanthes exigua ; 8-11, Achnanthes exigua var. 1; 12-17, Achnanthes gibberula; 18, Cocconeis placeniula var. lineata; 19-20, Achnanthes linearis; 21-24, Achnanthes lanceolata; 25, Cyclotella meneghiniana; 26-27, Amphora veneta; 28, Diatoma hiemale var. mesodon; 29-30, Denticula cf. parva; 31, Caloneis ventricosa var. truncatula ; 32-34, Anomoeoneis sphaerophora. All figures are 2000X. October 1986 curred at both sites but was most common in the stream, where it reached 14.0% relative density. Achnanthes exigua var. 1, Figs. 8-11. Valves orbicular to elliptical with rounded to rostrate ends, 5-10 wm long by 3.5-5 pm wide; raphe valve with narrow, linear axial area and narrow, rectangular central area; rapheless valve with less distinct central area, often formed from one or two shortened striae; striae 22-24 in 10 pm on both valves. We considered several taxa for the placement of these specimens but were not satisfied with their fit. It seems probable that they belong with A. exigua except that valve shape differs somewhat and striae are slightly coarser. This taxon was present at about 3% relative density in samples from both collecting sites. Achnanthes gibberula Grun., Figs. 12-17. Valves 5-26.5 wm long by 3-5.5 wm wide; striae 20—22 in 10 um on both valves. Several of our specimens were shorter than ordinarily observed for this taxon. Even so, an unbroken series from very small to the largest speci- mens was observed. A. gibberula was abun- dant in the Cowboy Hot Tub system, with a relative density of 14.8% in the concrete tub and 5.8% relative density in the stream. Achnanthes lanceolata Breb., Figs. 21—24. Valves 8-15 pm long by 4.5—5 pm wide; striae 13-14 in 10 pm on both valves. Several of our specimens were smaller than usual for this taxon. It was most abundant in the concrete tub at 1.6% relative density. Achnanthes linearis Grun., Figs. 19-20. Valves 8.5-15.5 pm long by 2-3 pm wide; striae 24—28 in 10 pm on both valves. Several of our specimens were smaller than typically observed for the nominate variety. A continu- ous range in length was observed so that we | did not place our smaller specimens in A. . linearis £. curta. This taxon showed prefer- ence for the stream, where it had a relative | density of 1.3%. Very few specimens were | observed in samples from the concrete tub. _ Amphora cf. coffeaeformis (Ag.) Kuetz., _ Fig. 119. Valve 15 pm long by 3 pm wide; striae indistinctly punctate, 24 in 10 pm. This _ Amphora corresponds to the description of A. coffeaeformis sensu Patrick and Reimer | (1975). However, it does not fit the taxon as | reinterpreted by Archibald and Schoeman | (1984). We saw only a single valve of this _ Amphora, which is common in some marshes EKINS, RUSHFORTH: CALIFORNIA DIATOMS 615 surrounding the Great Salt Lake of Utah (Fe- lix and Rushforth 1979, Squires et al. in press). Amphora veneta Kuetz., Figs. 26-27. Valves 15-29 wm long 3-5 wm wide; striae 21—24 at midvalve, becoming 28-32 at the ends. This Amphora was somewhat more common in the tub, where it reached 1% rela- tive density. _Anomoeoneis sphaerophora (Ehr.) Pfitz., Figs. 32-34. Valves 36.5-45 wm long by 10-14 wm wide; striae 18—20 in 10 pm. This taxon was present at both localities at about 7% relative density. Caloneis ventricosa var. truncatula (Grun.) Meist., Fig. 31. Valve 36 wm long by 7ym wide; striae 18 in 10 um. A single valve of this taxon was observed during our study. Cocconeis placentula var. lineata (Ehr.) V.H., Fig. 18. Valves 17.5-23 pm long by 9-12 wm wide; striae 16-21 in 10 pm. This taxon was rare in our study and was found only in the concrete tub. Cyclotella atomus Hust., Figs. 117-118. Valves 4—5 wm in diameter; striae 16—20 in 10 wm. This taxon was collected infrequently from both habitats. Cyclotella comta (Ehr.) Kuetz., Fig. 35. Valve 28 wm in diameter; striae 13-14 in 10 wm. A single valve of C. comta was observed. Cyclotella meneghiniana Kuetz., Fig. 25. Valves 7.5—10 wm in diameter; striae 8—10 in 10 pm. Only two valves of this taxon were observed during our study. Denticula elegans Kuetz., Figs. 39-41. Valves 10—31 jm long by 4-7 wm wide; costae 3-4 in 10 wm; striae 17—21 in 10 pm. Dentic- ula elegans was less frequent in Cowboy Hot Tub than in many other thermal springs in western North America. It had a relative den- sity of 2.4% in the concrete tub. It was present in the stream in lower numbers. Denticula cf. parva Hust., Figs. 29-30. Valves 9-18.5 wm long by 3-3.5 wm wide; costae 6-10 in 10 wm; striae not resolved. We have seen this taxon in several spring systems throughout western North America. It was present in Cowboy Hot Spring in low num- bers, always less than 1% relative density. Diatoma hiemale var. mesodon (Ehr.) Grun., Fig. 28. Valve 17.5 pm long by 8.5 pm wide; costae 3 in 10 wm; striae 24 in 10 pm. Only a single valve of this taxon was observed. 616 GREAT BASIN NATURALIST Vol. 46, No. 4 Figs. 35-44. Diatom species: 35, Cyclotella comta; 36, Epithemia sorex; 37, Diploneis oblongella; 38, F ragilaria construens var. binodis: 39-41, Denticula elegans; 42, Diploneis oblongella; 43, Epithemia argus, 44, Epithemia adnata var. porcellus. All figures are 2000X. | Gain, _ wide; striae 16 in 10 pm. A single valve of this _ taxon was observed during our study. October 1986 Diploneis oblongella (Naeg. ex Kuetz.) Ross, Figs. 37, 42. Valves 31-52 pm long by 15-18 ym wide; striae 12-13 in 10 wm. Sev- eral valves were collected from both habitats during our study. Epithemia adnata var. porcellus (Kuetz.) Patr., Fig. 44. Valve 90.5 pm long by 14 pm wide; costae 3 in 10 wm; striae 12 in 10 pm, 46 between costae. A single specimen of this taxon was observed. Epithemia argus (Ehr.) Kuetz., Fig. 43. Valves 60—90 pm long by 12-16.5 wm wide; costae 2 in 10 pm; striae 10-11 in 10 wm, 4-6 between costae. This Epithemia was observed infrequently in samples from the concrete tub. Epithemia sorex Kuetz., Fig. 36. Valve about 25 wm long by 9 wm wide; costae 12 in 10 pm; striae 2 between costae. Only a single frustule of this taxon was observed. Fragilaria construens Grun., Fig. 45. Valves 10-15 ym long by 5.5 wm wide; striae 14 in 10 pm. Only two valves of this taxon - were observed. Fragilaria construens var. binodis (Ehr.) Fig. 38. Valve 22 pm long by 5 pm Fragilaria construens var. venter (Ehr.) Grun., Figs. 46-48. Valves 5.5-10 wm long by 4—4.5 wm wide; striae 12-16 in 10 pm. This taxon was quite rare in our study, always less than 1% relative density. Fragilaria lapponica Grun., Fig. 53. Valve | 32 pm long by 6 wm wide; striae 10-12 in 10 | pm. A single valve of this taxon was observed. Fragilaria pinnata var. lancettula (Schum.) Hust., Fig. 52. Valve 20 wm long by 6 wm | wide; striae 10 in 10 pm. A single valve of this diatom was observed. Fragilaria similis Krasske, Fig. 51. Valve 15 » wm long by 5 pm wide; striae 10 in 10 wm. A single valve of F. similis was observed. Gomphonema gracile Ehr., Figs. 58-60. . | ; Valves 2432.5 wm long by 6.5—7.5 pm wide; striae 12-16 in 10 pm. It was present in low _numbers in Cowboy Hot Spring, always less than 1% relative density. Gomphonema parvulum Kuetz., Figs. 49— 50. Valves 16-17.5 wm long by 4.5-5 wm ; wide; striae 14 in 10 tm. Two frustules of this S@omphonema were observed during our study. Navicula confervacea (Kuetz.) Grun., Figs. | 73-74. Valves 18-26 pm long by 7-8 pm | | EKINS, RUSHFORTH: CALIFORNIA DIATOMS 617 wide; striae 19-24 in 10 wm. This taxon was present in samples from both sites in low numbers, always less than 1% relative den- sity. Navicula confervacea var. peregrina (W. Sm.) Grun., Figs. 69-71. Valves 6-13 wm long by 3.5- “4 um wide; striae 24 in 10 wm. Specimens of this variety were somewhat smaller than generally observed. It was present at about the same density as the nomi- nate in samples from both sites. Navicula cincta (Ehr.) Ralfs, Figs. 54-57. Valves 14—24.5 pm long by 4—5.5 wm wide; striae 14-16 in 10 wm. Our specimens of this taxon differ from the typical by being smaller with somewhat coarser striae. This diatom was present in rather high numbers, reaching 11.0% relative density in the thermal stream. Navicula cryptocephala var. veneta (Kuetz.) Rabh., Figs. 65-68. Valves 18-26 wm long by 5.5-6.5 wm wide; striae 14—16 in 10 wm. This Navicula was present in the con- crete pool at 2.2% relative density and 1% relative density in the thermal stream. Navicula halophila (Grun.) Cl., Figs. 63-64. Valves 34-53 ym long by 10-12.5 pm wide; striae 16-20 in 10 wm. This Navicula was more common in the stream, where it reached nearly 2% relative density. Navicula mutica Kuetz., Fig. 72. Valve 21 wm long by 7.5 wm wide; striae 18 in 10 pm. A single valve of this taxon was observed. Nitzschia clausii Hantz., Figs. 61-62. Valves 25.5—-36.5 um long by 3-4.5 wm wide; striae not resolved; fibulae 13-16 in 10 wm. Fibulae of our specimens were somewhat finer than previously reported for this taxon. It was present at less than 1% relative density in both sampling localities. Nitzschia communis Rabh., Figs. 77—79. Valves 20-38.5 wm long by 4-5 pm wide; striae approximately 32-36 in 10 pm, often unresolved; fibulae 10-16 in 10 pm. It was present at 4% relative density in both the concrete tub and the spring stream. Nitzschia frustulum (Kuetz.) Grun., Figs. 82-85. Valves 8-22 wm long by 3-4 wm wide; striae 21-22 in 10 wm; fibulae 10-12 in 10 pm. This Nitzschia was one of the most common diatoms in our study. It occurred at 32% rela- tive density in the concrete tub and as high as 30% relative density in the runoff stream. Nitzschia frustulum var. subsalina Grun., Figs. 89-90. Valves 13-20 wm long by 2.5-3 618 GREAT BASIN NATURALIST Vol. 46, No. 4 63 construens; 46—48, Fragilaria construens var. venter, 49-50, Gom- phonema parvuium,; 51, Fragilaria similis; 52, F ragilaria pinnata var. lancettula; 53, Fragilaria lapponica; 54-57, Navicula cincta; 58-60, Gomphonema gracile ; 61-62, Nitzschia clausii; 63-64, Navicula halophila; 65-68, Navicula cryptocephala var. veneta; 69-71, Navicula confervacea var. peregrina; 72, Navicula mutica; 73-74, Navicula confervacea. All figures are 2000X. Figs. 45-74. Diatom species: 45, Fragilaria t i icephala; 82-85, Nitzschia frustulum: 86-87, Nitzschia hantzschiana; 88, Nitzschia valdecostata:; 89-90, Nitzschia frustulum var. subsalina; 91-95, Nitzschia hantzschiana ; 96, Nitzschia species; 97-98, Nitzschia gracilis ; 99, Nitzschia heufleriana. All figures are 2000X. October 1986 EKINS, RUSHFORTH: CALIFORNIA DIATOMS 619 Figs. 75-99. Diatom species: 75—76, Nitzschia valdecostata; 77-79, Nitzschia communis ; 80-81, Nitzschia micro- 620 wm wide; striae 32 in 10 wm; fibulae 14-15 in 10 wm. Several valves of this taxon were ob- served in both the concrete tub and the ther- mal stream. Nitzschia gracilis Hantz., Figs. 97-98. Valves 40.5—62.5 wm long by 3-3.5 wm wide; striae 32—33 in 10 wm, often unresolved; fibu- lae 11-14 in 10 pm. This taxon was present in samples from both collecting sites at a relative density of less than 1%. Nitzschia hantzschiana Rabh., Figs. 86—87, 91-95. Valves 10.5-60 wm long by 2-3.5 wm wide; striae 22—24 in 10 wm; fibulae 10-13 in 10 pm. Our collections of this taxon contain specimens that are both shorter and longer than generally reported. However, a good size gradient was observed. This taxon oc- curred between 9% and 10% relative density in both the concrete tub and the stream. Nitzschia heufleriana Grun., Fig. 99. Valve 85 wm long by 7 wm wide; striae 20 in 10 pm; fibulae 10 in 10 wm. We saw a single specimen of this Nitzschia that had ends less capitate than usual. Nitzschia microcephala Grun., Figs. 80- 81. Valves 8.5-11.5 wm long by 2.5-3 pm wide; striae unresolved; fibulae 10—14 in 10 wm. This taxon reached 1% relative density in the thermal stream but was absent in the con- crete tub. Nitzschia valdecostata Lange-Bert. and Si- mon., Figs. 75-76, 88. Valves 13-37.5 wm long by 3.5—4.5 um wide; striae 16-18 in 10 wm; fibulae 7-9 in 10 wm. We used the epi- thet N. valdecostata rather than N. valdestri- ata since our specimens appeared to lack a nodulus. Petersen (1930) has also collected this taxon (as N. valdestriata) from thermal waters. This Nitzschia occurred in the con- crete tub at about 2% relative density and about 3% relative density in the stream. Nitzschia species, Figs. 96, 114. Valves lin- ear, greater than 100 pm long by 8 wm wide; striae 17—20 in 10 pm; fibulae 3 in 10 wm. This Nitzschia has been seen in several California thermal springs but has never been abundant in any of our samples. At Cowboy Hot Spring it was present in both the concrete tub and the spring stream in low numbers. Pinnularia appendiculata (Ag.) Cl., Figs. 102-104. Valves 18.5-25 pm long by 4.5-6 wm wide; striae 20-22 in 10 wm. Some speci- mens demonstrated more strongly radiate striae near midvalve (Figs. 103-104) than oth- GREAT BASIN NATURALIST Vol. 46, No. 4 ers. However, it appeared that this feature intergraded in the population. This taxon was more common in the stream but occurred at less than 1% relative density. Pinnularia intermedia (Lagerst.) Cl., Figs. 107-109. Valves 11.5-17 pm long by 3-3.5 wm wide; striae 10-12 in 10 pm. Our speci- mens were shorter than usual for this taxon. Some of our specimens were similar to P. obscura since they had up to 12 striae in 10 ym. This diatom was rare in our samples. Pinnularia microstauron (Ehr.) Cl., Figs. 111-113. Valves 24-52 pm long by 7-13 pm wide; striae 10-12 in 10 wm. Some of our specimens are very similar to P. brebissonii. However, since they seemed to be at one end of a morphological gradient from the more common and more typical P. microstauron specimens, we opted to use the latter specific name. This taxon was not particularly com- mon, although a number of frustules were collected. Pinnularia stauroptera var. recta (May.) Cleve-Euler, Fig. 110. Valve 43 wm long by 7.5 jm wide; striae 9 in 10 wm. Hustedt (1930) used Pinnularia gibba var. linearis for specimens similar to ours. A single specimen of this taxon was observed in a sample from the thermal stream. Rhopalodia gibberula (Ehr.) O. Muell., Fig. 115. Valve 80-127.5 pm long by 12-12.5 wm wide; striae 18 in 10 wm; costae 3-5 in 10 wm. It was rare in the concrete tub. Rhopalodia operculata (C. A. Ag.) Hakans- son, Fig. 100-101, 120. Valves 25-49 pm long; 6-12 wm wide; striae 18-20 in 10 wm; costae 3-6 in 10 wm. This taxon occurred in samples from the concrete tub at 2% relative density and was somewhat rarer in the stream. Stauroneis wislouchii Poretz. et Anisi- mowa, Figs. 105-106. Valves 22-34 wm long by 5-8 wm wide; striae 22-26 in 10 wm. This taxon was present in samples from both col- lecting localities at about 0.5% relative den- sity. Stephanodiscus carconensis var. pusilla Grun., Fig. 124. Valves 26-28 wm in diame- ter; large striae 4 in 10 tm, composed of dis- tinct rows of punctae; rows of punctae 17-18 in 10 wm. Only two valves of this taxon were found in our samples. Surirella ovalis Breb., Fig. 116. Valve 113 wm long by 53 mm wide; striae 14 in 10 wm; October 1986 EXKINS, RUSHFORTH: CALIFORNIA DIATOMS 621 Figs. 100-114. Diatom species: 100-101, Rhopalodia operculata; 102-104, Pinnularia seas re Siamoncis wislouchii; 107-109, Pinnularia intermedia: 110, Pinnularia stauroptera var. recta; -113, microstauron, 114, Nitzschia species. All figures are 2000X. 622 GREAT BASIN NATURALIST Vol. 46, No. 4 Figs. 115-124. Diatom species: 115, Rhopalodia gibberula; 116, Surirella ovalis; 117-118, Cyclotella atomus; 119, Amphora cf. coffeaeformis; 120, Rhopalodia operculata; 121, Tabellaria quadrisepta; 122, Synedra ulna; 123, Surirella ovalis var. brightwellii; 124, Stephanodiscus carconensis var. pusilla. Figures 115, 116 and 122 are 1000X. All others are 2000X. October 1986 wing canals 3 in 10 pm. Only a single speci- men of this taxon was observed in our sam- ples. Surirella ovalis var. brightwellii (W. Sm.) Cl., Fig. 123. Valves 39 wm long by 21-23 wm wide; striae 19-20 in 10 wm; wing canals 5-6 in 10 wm. Only two specimens of this taxon were observed in our samples. Synedra ulna (Nitz.) Ehr., Fig. 122. Valve 115 pm long by 6.5 wm wide; striae 10 in 10 wm. This taxon occurred as a single specimen from the concrete tub. Tabellaria quadrisepta Knuds., Fig. 121. Valves 37-65 wm long by 6 wm wide at mid- valve; striae 15-16 in 10 pm. Only three frus- tules of this taxon were observed in our sam- ples. LITERATURE CITED ARCHIBALD, R. E. M., AND F. R. SCHOEMAN. 1984. Am- phora coffeaefromis (Agardh) Kuetzing: a revision of the species under light and electron micro- scopy. South African Jour. Bot. 3: 83-102. BIEBL, R., AND KUSEL-FETZMANN. 1966. Beobachtungen ueber das Vorkommen von Algen an Thermalstan- dorten auf Island. Osterreich. Bot. Zeitschr. 113: 408—423. CALIFORNIA GEOLOGIC Map SERIES. 1980. Map 4. Geothermal Resources of California. Nat. Geophys. and Solar-Terrestrial Data Center, Nat. Ocean. and Atmos. Admin. Davis, D. M. 1897. The vegetation of the hot springs of Yellowstone Park. Science 4(134): 145-157. Epwarps, M. E. 1868. On the occurrence of living forms in the hot waters of California. Amer. Jour. Sci. and Arts 2(65): 239-241. Emoto, Y., AND H. Hirose. 1940. Studien ueber die Ther- malflora von Japan (III). Thermale Bacterien und Algen aus den themalen Quellen von Hakone (2). Shokobutsu Kenkyn Zasshi 16: 405-420. Emoto, Y., ANDY. YONEDA. 1941. Bacteria and algae of the thermal springs in Simane Prefecture (1). Shoko- butsu Kenkyn Zasshi 17: 654-715. FaMIN, M. A. 1933. Action de la temperature sur les vegetaux. Part 4: les vegetaux vivant dans les eaux naturellement a temperatures elevees. Rev. Gen- erale Bot. 45: 574—595, 655-682. FELIX, E. A., AND S. R. RUSHFORTH. 1979. The algal flora of - the Great Salt Lake, Utah, USA. Nova Hedwigia 31(12): 163-195. . HusTEDT, F. 1930. Die Kieselalagen Deutschlands, Os- terreichs und der Schweiz mit Berucksichtigung der ubrigen Lander Europas sowie der angren- zenden Meeresgebiete. Vol. 7 in L. Rabenhorst, Kryptogamen-flora von Deutschland, Osterreich und der Schweiz. Akad. Verlagsgesell., Leipzig. Reprint 1971, Johnson Reprint Corp., London. KACZMARSKA, L., AND S. R. RUSHFORTH. 1983. The diatom flora of Blue Lake Warm Spring, Utah, USA Bib- lio. Diatomologica 2(1): 1-123. EKINS, RUSHFORTH: CALIFORNIA DIATOMS 623 KOL, E. 1932. Thermalvegetation von Hajduszoboszolo in Ungarn. Pt. 4, Algen. Archiv Protistenk. 76: 309— 324. Lacsny, I. L. 1912. Beitraege zur algen Flora der Ther- malwasser bei Nadyvard. Botanikai Kozlemenyek 11(5-6): 167-185. MarGa EF, R. 1958. “Trophic” typology versus biotic ty- pology, as exemplified in the regional limnology of northern Spain. Verh. intern. Ver. Limno. 13: 339-349. MILLER, R. 1985. Continents in Collision. Time Life Books, Alexandria, Virginia. 176 pp. NEGorO, K. 1940. The diatom flora of the Nasu Hot Spring (preliminary report). Bot. Mag. 54(638): 63-65. NELson, C. A. 1981. Basin and range province. Pages 203-216 in W. G. Ernst, The geotectonic devel- opment of California. Prentice-Hall, Inc., Engle- wood Cliffs, New Jersey. OestRup, E. 1918 (1920). Fresh-water diatoms of Iceland. Pages 1-90 in Rosenvinge and Warming, Botany of Iceland, vol. 2, 1(5): 1-96. Arbeider Botaniske Have Kobenhavn. PATRICK, R., AND C. REIMER. 1975. The diatoms of the United States exclusive of Alaska and Hawaii. Mon. Acad. Nat. Sci. Philadelphia 13, vol. 2, pt. 1. 213 pp. PATTEN, B. C. 1962. Species diversity in net phytoplank- ton of Raritan Bay. J. Marine Res. 20: 57-75. PETERSEN, J. B. 1930. Algae from O. Olufsens second Danish Pamir Expedition 1898-1899. Dansk Botanisk Arkiv 6 No. 6, Copenhagen. RUSHFORTH, S. R., L. E. SQUIRES, AND J. E. JOHANSEN. 1986. Three new records for diatoms from the Great Basin, USA. Great Basin Nat. 46(3): 398-403. RUTINER, R. 1963. Fundamentals of limnology. Univer- sity of Toronto Press. Toronto, Ontario, Canada. 295 pp. SHANNON, C. E., AND W. WEAVER. 1963. The mathemati- cal theory of communication. University of Illinois Press, Urbana. 117 pp. SPRINGER, E. 1930. Bacillariales aus den Thermen und der Umgebung von Karlsbad. Archiv Prot- istenkunde 71: 502-542. Sourres, L. E., S. R. RUSHFORTH, AND C. C. NEWBERRY. The diatom flora of newly inundated land at the south end of the Great Salt Lake, Utah, USA. Great Basin Nat. In press. St. Car, L. L., AND S. R. RUSHFORTH. 1977. The diatom flora of the Goshen Warm Spring ponds and wet meadows, Goshen, Utah, USA. Nova Hedwigia 28: 353-425. STOCKNER, J. G. 1967a. Observations of thermophilic algal communities in Mount Rainier and Yellowstone National Parks. Limnol. Oceanogr. 12(1): 13-17. .1967b. The ecology of the Ohanoapecosh Hot Springs, Mt. Rainier National Park, Washington. Unpublished dissertation, University of Washing- ton, Seattle. 232 pp. 1968. The ecology of a diatom community in a thermal stream. Brit. Phycol. Bull. 3(3): 501-514. Strom, K. M. 1921. Some algae from hot springs in Spitzbergen. Botaniska Notiser 1921: 17—21. TARBUCK, E. J., AND F. K. Lutcens. 1984. The earth: an introduction to physical geology. Charles E. Mer- rill Pub. Co., Columbus, Ohio. 594 pp. 624 GREAT BASIN NATURALIST Vol. 46, No. 4 THOMAS, J., AND E. A. GONZALVES. 1965a. Thermal algae of western India, I. Algae of the hot springs at Akloi and Ganeshpuri. Hydrobiologia 25: 330-340. . 1965b. Thermal algae of western India, II Algae of the hot springs at Palli. Hydrobiologia 25: 340-351. . 1966a. Thermal algae of western India, III. Algae of the hot springs at Sav. Hydrobiologia 26: 21-28. . 1966b. Thermal algae of western India, IV. Algae of the host springs at Aravali, Tooral and Rajewadi. Hydrobiologia 26: 29—40. . 1966c. Thermal algae of western India, V. Algae of the hot springs at Tuwa. Hydrobiologia 26: 41-54. . 1966d. Thermal algae of western India, VI. Algae of the hot springs at Unai, Lasundra and Unapdeo. Hydrobiologia 26: 55-65. . 1966e. Thermal algae of western India, VII. Algae of the hot springs at Ragapur. Hydrobiologia 26: 66-71. TILDEN, J. E. 1897. On some algal stalactites of Yellow- stone National Park. Bot. Gazette 24: 144-199. _____. 1898. Observations on some west American ther- mal algae. Bot. Gazette 25: 89-105. WHitTForD, L. A. 1956. The communities of algae in the springs and spring systems of Florida. Ecology 37(3): 432-442. YONEDA, Y. 1942a. Bacteria and algae of hot springs in Gihu Prefecture. Shokobutsu Bunrui Chiri 11: 83-100. —____. 1942b. Bacteria and algae of hot springs in Wakayama Prefecture. Shokobutsu Bunrui Chiri 11: 194-210. —____.. 1962. Study of the thermal algae of Hokkaido. Shokobutsu Bunrui Chiri 20: 308-313. INVENTORY OF UTAH CRAYFISH WITH NOTES ON CURRENT DISTRIBUTION James E. Johnson! ABSTRACT.—Crayfish distribution and composition in Utah are poorly documented. Based upon limited collections the native Pacifastacus gambelii is widespread and often abundant in the Bear and Weber river drainages and 6 occasionally represented in smaller tributaries to the Great Sait Lake and to the Raft River. Pacifastacus leniusculus was collected from Utah County and Procambarus clarkii from Tooele County; the origin of these populations is not known. The nonnative crayfish Orconectes virilis is currently abundant in the Virgin, Price, and Duchesne river basins and the Glen Canyon, Flaming Gorge, Deer Creek, and Starvation reservoirs; it is expanding in Huntington North, Scofield, and Willard reservoirs. Further work is required to develop a more complete inventory and monitor the impacts on aquatic ecosystems of expanding nonnative crayfish populations. Surreptitious stockings can be dealt with only if the public is made aware of the adverse consequences of ill-conceived introductions. Crayfish, as listed in Hobbs (1976), are rep- resented in Utah by only one native species, Pacifastacus (Hobbsastacus ) gambelii (Girard 1852), and that only north of approximately Salt Lake county (personal observation). Ap- parently crayfish are not native to the Green—Colorado River system (Dean 1969) or to that portion of the Bonneville Basin, in- cluding the Sevier River drainage, south of approximately Utah County (personal obser- vation). Two other species of Pacifastacus are native to waters adjoining Utah, Pacifastacus (Pacifastacus ) leniusculus (Dana 1852) in Ne- vada and Pacifastacus (Hobbsastacus) con- nectens (Faxon 1914) in Idaho (Pennak 1978) and may be native to Utah waters; however, this possibility has yet to be confirmed. Pacifastacus leniusculus is present in Utah County but may have been introduced there. The introduction of Orconectes virilis (Hagen 1870) has resulted in burgeoning populations of this nonnative in several of Utah’s major drainages in recent years. One isolated popu- lation of Procambarus (Scapulicambarus ) clarkii (Girard 1852) is found in Tooele County, probably also representing an intro- duction. Crayfish feed on vegetation, and certain species have been shown to control nuisance aquatic plants (Dean 1969). They also feed on detritous and are considered important com- ponents of food webs supplying certain fish- eries (Jones and Momot 1981). Their food habits and importance as food for man and as prey for various species of sportfish have led to the widespread introduction of certain cray- fish species. Since approximately 1950 Or- conectes virilis” has been stocked in the Colo- rado River watershed of western New Mexico and northeastern Arizona, primarily for vege- tation control (Dean 1969). During 1967, 1968, and 1970, O. virilis were collected by Utah Division of Wildlife Resources (UDWR) personnel from Nogal Lake in south central New Mexico, Red Lake on New Mexico's Navajo Indian Reservation, and the Little Colorado River of Arizona. These crayfish were stocked in the Sand Cove reservoirs (up- per Santa Clara River drainage of the Virgin River) and Pelican Lake, Walls, Vernal Golf Course, Rasmussen and Stringham ponds (in the Vernal, Utah, area, Duchesne River drainage), and in a golf course pond adjacent to the Price River near Price, Utah. Prior to 1968 the U.S. Fish and Wildlife Service (US- EWS) had planted O. virilis in Towave and Midway reservoirs, Uinta Indian Reservation (also Duchesne drainage). Crayfish were reported to be a prominent food of largemouth bass at Glen Canyon Reservoir, Utah-Arizona, shortly after its im- poundment (May et al. 1975), and the pres- ence of crayfish in Flaming Gorge Reservoir, Utah-Wyoming, was confirmed during the 1970s. In neither case was the species of cray- fish identified. lUtah Division of Wildlife Resources, 1596 West North Temple, Salt Lake City, Utah 84116. mer ae 7In the opinion of Horton H. Hobbs, Jr., Smithsonian Institution, Washington, D.C., Orconectes causeyi (Jester 1967) is a synonym of O. virilis (personal communication, 1984). Orconectes virilis and O. causeyi are, therefore, collectively referred to as O. virilis in this paper. 625 626 VU 7 TOOELE GREAT BASIN NATURALIST Vol. 46, No. 4 W/s/// Orconectes virilis NO \y Pacifastacus gambelii mee Pacifastacus leniusculus feaaysy;s Procambarus clarkii UTAH 0 10 20 © 0 © a) SCALE OF MILES | 1a.pring Ly, Gerge es. Re emma A.| ei py Jen W/Canyon \ Reservoir 0 io Fig. 1. Approximate current distribution of crayfish in Utah, based upon UDWR fishery collections made from 1978 to 1984. The distribution of crayfish has spread rapidly since 1975, although the UDWR made no successful introductions after July 1977, when the Fisheries Section imposed a moratorium on further stockings. Anglers, however, observed the importance of crayfish in the diets of game fish at Lake Powell and other waters, and it is suspected that this knowledge led to a rash of surreptitious plant- ings of crayfish by the public in additional drainages. (It is unlawful in Utah for anglers to transport live fish for bait; therefore, the bait- bucket is probably not a significant vector of crayfish in Utah.) Furthermore, previous in- troductions had resulted in some dense popu- lations that were spontaneously spreading within their drainages. The purpose of this paper is to contribute to October 1986 JOHNSON: UTAH CRAYFISH 627 TaBLE 1. Crayfish collection sites by UDWR fishery personnel, from which preserved specimens were made available to the author, 1978-1984. eee Collection Drainage Site date Collector Species COLORADO Virgin River Lower Sand Cove 1982 D. Hepworth and Orconectes virilis J. Leppink Gunlock 1983 D. Hepworth Orconectes virilis Beaver Dam Wash 3 July 1984 W. Gustaveson Orconectes virilis Glen Canyon Wahweap Bay Composite of W. Gustaveson Orconectes virilis Reservoir 1982 and 1983 San Juan Arm Composite of W. Gustaveson Orconectes virilis 1982 and 1983 Bullfrog Bay Composite of W. Gustaveson and = Orconectes virilis 1982 and 1983 S. Scott Flaming Gorge Near Dam October 1982 S. Brayton Orconectes virilis Reservoir Starvation Boat Ramp September 1983 M. Ottenbacher and Orconectes virilis Reservoir S. Scott Price River Scofield Reservoir September 1984 W. Donaldson Orconectes virilis Drainage GREAT SALT LAKE Provo River Deer Creek Reservoir 21 August 1981 D. Sakaguchi Orconectes virilis Weber River Willard Reservoir 7 December 1982 J. Leppink Orconectes virilis Morgan-Peterson 16 September 1982 J. Leppink Pacifastacus gambelii Lost Creek Lost Creek Reservoir 12 August 1984 J. Johnson Pacifastacus gambelii Bear River Wellsville and Hyrum September 1982 T. St. John Pacifastacus gambelii reservoirs Bear River Bear Lake Several B. Nielson ° Pacifastacus gambelii Bear River Big Creek 1982 J. Leppink Pacifastacus gambelii Great Salt Lake Salt Creek 5 September 1984 K. Summers Pacifastacus gambelii Utah Lake Salem Pond 1981 D. Sakaguchi Pacifastacus leniusculus Utah Lake Spring Pond 1981 D. Sakaguchi Pacifastacus leniusculus COLUMBIA RIVER Raft River Cotton-Thomas Basin _17 August 1984 J. Leppink Pacifastacus gambelii WESTERN BasINS __ Rush Valley near 1978 and D. Sakaguchi Procambarus clarkii St. John 17 August 1983 the current inventory of the distribution of crayfish in Utah and to serve as a baseline for future studies. The information presented is not comprehensive; sampling was largely by convenience rather than design, and large ar- eas, especially Utah’s western basins, were not sampled. METHODS All UDWR fishery biologists were asked to preserve in formalin all crayfish collected inci- dental to scheduled fish sampling. Thus, spec- imens were collected primarily with gill nets and electrofishing gear. In addition, some specimens were collected by hand from the substrate rubble and using baited lift nets and cage traps. Specimens were labeled as to cap- ture site, date, and method and sent to the author for identification. Collection sites, therefore, largely represent waters under public sportfishery management programs. Very few collection efforts were made on fish- less waters. Tentative identification was made using keys of Hobbs (1976) and Pennak (1978). Sam- ples of each species identified, with the excep- tion of Pacifastacus leniusculus, were sent to 628 H. H. Hobbs, Jr., Department of Inverte- brate Zoology, U.S. National Museum of Nat- ural History, Smithsonian Institution, Wash- ington, D. C., for confirmation of my tentative identification. RESULTS Colorado—Green River Drainage Although not endemic, crayfish are now widespread in the Utah portion of the Colo- rado—Green River drainage. All specimens collected to date have been Orconectes virilis (Fig. 1, Table 1). The species is abundant in Flaming Gorge and Glen Canyon reservoirs and in the Virgin and Duchesne drainages but has not appeared in fish collections from the San Juan, White, Yampa, Green, or Colorado rivers within Utah. It has, however, been col- lected from the Colorado River upstream of approximately Grand Junction, Colorado (Unger 1978). Crayfish were apparently absent until very recently in the Strawberry River above Sol- dier Creek Dam, based upon their absence from extensive UDWR fish sampling of Sol- dier Creek and Strawberry reservoirs and their tributaries. Anglers have reported, how- ever, that crayfish are now present in Soldier Creek Reservoir. Reports have been received of the presence of O. virilis from Scofield and Huntington North reservoirs and the Price River. A speci- men from Scofield Reservoir, sent to me for verification by W. Donaldson (UDWR South- eastern Regional office, Price, Utah, Novem- ber 1984), proved to be O. virilis. Great Salt Lake Drainage If crayfish were native to the Provo River drainage, the species should have been Pacifastacus gambelii, the documented na- tive of the Bonneville basin (Hobbs 1972). No specimens of P. gambelii from the Provo River have come to my attention; however, O. virilis began appearing in fish sampling gear in 1981 at Deer Creek Reservoir. By 1984 they were abundant at Deer Creek Reservoir and were reported to have been seen in the Provo River downstream of Deer Creek Dam (Sakaguchi 1984). Orconectes virilis was collected from the inlet of Willard Reservoir, a freshwater im- poundment on the Bear River arm of the GREAT BASIN NATURALIST Vol. 46, No. 4 Great Salt Lake, in November 1982. The col- lection site was below a large drop structure that may serve as a barrier to upstream migra- tion of crayfish to the Weber River. I have also identified this species in Stansbury Park Lake in Tooele County near the south shore of the Great Salt Lake. These populations appar- ently initiated with surreptitious stockings. Pacifastacus gambelii is native and wide- spread in the Ogden/Weber drainage. They were collected in abundance from the Weber River at Morgan in 1982 and from Lost Creek Reservoir in 1984. Crayfish have been ob- served by UDWR personnel in Rockport, Echo, and East Canyon reservoirs. With the exception of Willard Reservoir, only P. gam- belii has been identified from the Weber drainage to date. All collections to date from the Bear River have been identified as Pacifastacus gambelii (Table 1). The species is widespread and occa- sionally very abundant. No UDWR collec- tions of crayfish have been made from the Bear River upstream of approximately the Woodruff Narrows. Wyoming Game and Fish Department personnel (W. Wengert, Green River, Wyoming, personal communication, 1984) have observed crayfish in Woodruff Narrows, Huff Creek, and Salt Creek, of the Bear River drainage. These specimens were not identified but were very likely P. gambe- lit. Pacifastacus gambelii is abundant in Salt Creek, a tributary to the North Arm of the Great Salt Lake. Other tributaries to the North Arm contain crayfish but have not been inventoried. Specimens from Salem and Spring ponds near Payson, Utah County, Utah, collected by D. Sakaguchi, were tentatively identified as Pacifastacus leniusculus , native to California, Oregon, Idaho, Washington, and Nevada (Pennak 1978). The author has been informed of observa- tions of crayfish and shallow burrows in wet- lands surrounding Utah Lake, but no speci- mens have been collected. Raft River (Columbia) Drainage Although comparatively little effort has been expended searching for crayfishes in the Raft River basin of the Columbia Drainage, Pacifastacus gambelii has been collected from two small tributaries (Table 1) and is thought October 1986 to be present elsewhere in the Raft River drainage. Western Basins and Sevier River Drainages Crayfish have not been observed during extensive samplings of the Pilot or Deep Creek Mountain drainages; nor have crayfish been collected from western basin natural lakes and wetlands. Orconectes virilis has appeared in New- castle Reservoir in southern Iron County, ap- parently from sources in the adjacent Santa Clara River drainage (D. Hepworth, UDWR, personal communication, 1984). Specimens collected by D. Sakaguchi, UDWR, from a warm spring in Rush Valley, Tooele County, in 1978 and 1983 were identi- fied by H. Hobbs, Jr. (U.S. National Museum of Natural History, Smithsonian Institution, Washington, D.C.) to be Procambarus clarkii, a species common in the south central United States (Pennak 1978). No crayfish have been reported from any- where within the Sevier drainage. Crayfish may not be endemic to this area and, appar- ently, have not yet been introduced. DISCUSSION Colorado River Drainage The presence of Orconectes virilis in the Colorado River drainage is probably the result of a number of introductions. The Santa Clara River populations originated with the UDWR _ introduction in 1970 at Sand Cove Reservoir in the upper Santa Clara drainage. This intro- duction was followed by observations of cray- fish in Gunlock (1978) and Ivans (1980) reser- _voirs and the lower Santa Clara River and _ upstream through Baker Reservoir (1978). In _ 1983 O. virilis was collected from ponds along the East Fork of Beaver Dam Wash. In the Virgin River and Ash Creek, O. virilis is cur- rently distributed upstream to approximately La Verkin and Toquerville, respectively. Pro- cambarus clarkii is common in Lake Mead and therefore has access to the Virgin River. No. P. clarkii have been sampled to date from the Virgin drainage of Utah, however. The origin of Orconectes virilis in Glen Canyon Reservoir is less clear. One surrepti- tious plant of O. virilis is believed to have occurred in 1965; but crayfish were known to JOHNSON: UTAH CRAYFISH 629 be present in the reservoir as early as 1964 (May et al. 1975). Evidently O. virilis was present in the drainage at the time of im- poundment or was introduced accidentally with game fish, many of which were obtained from midwestern hatcheries. The apparent absence of crayfishes in Utah waters of the mainstem Colorado and Green rivers and the San Juan River upstream of Lake Powell deserves attention. Orconectes virilis now has access from both upstream and downstream and may colonize these reaches and eventually populate the White, Yampa, and other tributaries, all with unique native fish populations. Orconectes virilis generally occurs in relatively clear waters with stony bottoms (Pennak 1978) and may therefore prove intolerant of the Colorado system’s silt- laden reaches. The UDWR and USFWS introductions in the Uinta Basin during the period 1965-1968 resulted, by approximately 1975, in dense populations of Orconectes virilis in Bottle Hollow, Midview (Boreham), and numerous smaller reservoirs, as well as in the Duchesne River from its mouth upstream at least through the City of Duchesne. Although abundant in the water supply of Pelican Lake, the species has not become established in Pel- ican. Between 1980 and 1984 O. virilis ap- peared in Steinaker and Starvation reservoirs. The upstream extent of the current distribu- tion of O. virilis in Duchesne River tributaries is presently poorly documented. The UDWR introduction in a golf course pond near Price was probably the source of the lower Price River and Huntington North Reservoir popu- lations, but the crayfish in Scofield Reservoir probably resulted from surreptitious stock- ings. Crayfish were probably absent from Flam- ing Gorge Reservoir and its drainage at the time of impoundment. Wyoming Game and Fish Department planted Pacifastacus gam- belii in the Green River, between Flaming Gorge and Fontenelle reservoirs, in 1965 and 1966. The source of these crayfish was the Teton Valley Ranch near Jackson, in the Snake River drainage. In 1974 Wyoming planted largemouth bass into Flaming Gorge from Springer Pond, south of Buffalo, Wyo- ming. A few small crayfish were captured and stocked incidentally with the bass. The cray- 630 fish of Springer Pond have been identified by Wyoming Game and Fish Department (W. Wengert, Green River, Wyoming, personal communication 1984) as Orconectes virilis. Fish hatcheries with dirt ponds often harbor crayfish, and a variety of such hatcheries, some located within the native range of O. virilis, have contributed to the stocking of Flaming Gorge Reservoir over the years. This species is now abundant in Flaming Gorge Reservoir. It is the primary source of prey for the reservoirs smallmouth bass population and contributes to the diets of lake trout, brown trout, and rainbow trout (Pettengill et al. 1984). Great Salt Lake Drainage Pacifastacus gambelii has not been col- lected from Willard Reservoir; the Weber River downstream of approximately Peterson, Utah; or the Bear River from Cutler Reservoir downstream to the Great Salt Lake. Their apparent absence from these warmer waters suggests this species may be intolerant of warmer waters or of warm water fish popula- tions. It has not been collected from the Og- den River drainage, possibly because fish toxi- cants were used in the reclamation of the fisheries of Pineview and Causey reservoirs. The population of Orconectes virilis in Willard Reservoir is at the lower extreme of the Weber drainage. Confined by salt water downstream and a drop structure in the inlet canal, this species may not have access to the lower Weber River. If O. virilis succeeds in reaching the river, it can be expected to spread rapidly upstream, in a manner similar to its rapid colonization of the Duchesne and Price rivers. Furthermore, it is possible the native species will become extirpated or re- duced in number in much of the Salt Lake drainage, such as occurred to native species following O. virilis introductions in Maryland (Schwartz et al. 1963), Tennessee, West Vir- ginia, Mississippi (Bouchard 1976), and possi- bly California (Bouchard 1977, Eng and Daniels 1982). Orconectes virilis is a very suc- cessful and aggressive species (Bouchard 1977) and could well displace Utah’s native species (H. Hobbs, Jr., personal communica- tion, 1984). Because of the tendency of an- glers to transplant crayfish, and the availabil- ity of O. virilis in several popular fishing waters, including Deer Creek, Flaming GREAT BASIN NATURALIST Vol. 46, No. 4 Gorge, and Glen Canyon reservoirs, further appearances of O. virilis can be expected in waters attractive to anglers. Only one specimen of Pacifastacus lenius- culus each was collected from Salem and Spring ponds, Utah County. This species is clearly not as abundant or widespread as P. gambelii, and its origin in Utah is uncertain. If native, discovery of further populations in the Payson—Spanish Fork area, and perhaps in the western basins, can be expected. Raft River (Columbia) Drainage Pacifastacus connectens is reported to be native to Idaho and northern Utah (Eng and Daniels 1982). The specimens collected in the Raft River drainage proved to be P. gambelii, however, and no specimens of P. connectens have as yet come to my attention. If P. con- nectens is indeed represented in Utah, it might be expected to be present in the Co- lumbia drainage. All future specimens from this drainage should be closely inspected; both P. connectens and P. gambelii have dor- sal patches of setae on the palm of the chela, and a cursory inspection could therefore re- sult in misidentification. Western Basins and Sevier River Drainages The waters of western Utah, within the Bonneville basin, are largely uninventoried with respect to crayfish. The only crayfish populations recorded in this area would ap- pear to be Orconectes virilis in Newcastle Reservoir at the southern extreme of the basin, first observed in 1980 (the result of surreptitious stocking), and the population of Procambarus clarkii in a small warm spring near St. John in Rush Valley. It seems very unlikely P. clarkii is native, so far removed is Utah from its documented range. Introduc- tions have resulted in established populations in California and Nevada (Pennak 1978), but any introductions of the species in this remote St. John site are undocumented and unex- plained. Previous to its discovery, there was no reason to believe P. clarkii was present north of Lake Mead, Arizona-Nevada, and its tributaries. Extensive fishery collections have been made in most of the Sevier River drainage and, based upon their absence from these collections, it seems reasonable to conclude crayfish are currently not in the drainage. \ SSeS a eet enone sige ae AY October 1986 NEED FOR FURTHER STUDY Because of the rapid expansion of Or- conectes virilis in much of Utah, expansion of this species must be closely monitored and its impacts upon sportfisheries and native fauna should be documented. In addition to its po- tential impact on native crayfish, there are indications of negative effects of dense popu- lations with rainbow trout recruitment and early growth (Hepworth and Duffield, in press), and near elimination of aquatic vegeta- tion (Dean 1969). The effects of extensive re- moval of vegetation on invertebrate produc- tion and availability and diversity of littoral zone fishery habitats also require study. The rash of surreptitious stockings of cray- fish in recent years demonstrates an obvious need for a thorough public information pro- gram regarding the possible consequences of indiscriminate introductions. Such a_pro- gram, to be most effective, requires basis in fact and would alone justify well-conceived research into crayfish population dynamics and ecosystem interactions. Crayfish distribution is especially poorly documented in the western basins of Utah. Furthermore, the collection of Pacifastacus leniusculus in Utah County raises the ques- tion of whether that species is more widely distributed around Utah Lake or in the west- ern basins. Any crayfish specimens from Utah County or the western basins would therefore be of particular interest. ACKNOWLEDGMENTS I am grateful to Dr. Horton H. Hobbs, Jr., U.S. National Museum of Natural History, Smithsonian Institution, Washington, D.C., for confirming the identity of specimens from my reference collection, for identifying Pro- cambarus clarkii, and for his helpful com- ments. Specimens were collected by virtually every regional or research UDWR biologist. In addition, specimens and collection records from the Colorado River drainage were pro- vided by V. Lamarra and R. Valdez of Ecosys- tem Research Institute, Logan, Utah; R. Radant, Nongame Section, UDWR, and H. Tyus, USFWS, Vernal, Utah. JOHNSON: UTAH CRAYFISH 631 This work was accomplished in conjunction with fishery studies funded by Federal Aid to Fish Restoration Projects F-43-R and F-44-R. LITERATURE CITED BOUCHARD, R. W. 1976. Geography and ecology of cray- fishes of the Cumberland Plateau and Cumber- iand Mountains, Kentucky, Virginia, Tennessee, Georgia, and Alabama. Part I. The genera of Pro- cambarus and Orconectes . Freshwater Crayfish 2: 563-584. . 1977. Distribution, systematic status and ecologi- cal notes on five poorly known species of crayfishes in western North America (Decapoda: Astacidae and Cambaridae). Freshwater Crayfish 3: 409-423. J. L. 1969. Biology of the crayfish Orconectes causeyi and its use for control of aquatic weeds in trout lakes. U.S. Bureau of Sportfisheries and Wildlife, Technical Paper 24. 15 pp. ENG, L. L., AND R. A. DANIELS. 1982. Life history, distri- bution, and status of Pacifastacus fortis (Deca- poda: Astacidae). California Fish and Game 68(4): 197-212. HEPwortH, D. AND D. DUFFIELD. In press. Impact of an exotic crayfish on stocked rainbow trout in New- castle Reservoir, Utah. North American Journal of Fisheries Management. Hoss, H. H., Jr. 1976. Crayfishes (Astacidae) of North and Middle America. U.S. Environmental Protec- tion Agency Water Pollution Control Research Series, 18050 E1D05/72. Cincinnati, Ohio. 173 pp: | Jones, P. D., AND W. T. Momot. 1981. Crayfish produc- tion, allochthony, and basin morphometry. Cana- dian Jour. Fish. and Aqu. Sci. 38: 175-183. May, B., C. THOMPSON, AND S. GLoss. 1975. Impact of threadfin shad (Dorsoma petenense ) introduction on food habits of four centrarchids. Utah Division of Wildlife Resources, Publ. 75—4. 22 pp. PENNAK, R. W. 1978. Freshwater invertebrates of the United States. John Wiley and Sons, New York. 803 pp. PETTENGILL, T., S. BRAYTON, AND J. JOHNSON. 1984 annual report. Flaming Gorge fishery investigations. Utah Division of Wildlife Resources, Publ. 84—07. 50 pp. SAKAGUCHI, D. K. 1984. Considerations for managing with crayfish in the waters of the Central Region. Utah Division of Wildlife Resources, Central Regional Office, Springville, Utah. Memo report. 6 pp. SCHWARTZ, F. J., R. RUBELMANN, AND J. ALLISON. 1963. Ecological population expansion of the introduced crayfish, Orconectes virilis. Ohio Jour. of Sci. 63(6): 266-273. UnceEr, P. A. 1978. The crayfishes (Crustacae: Cam- baridae) of Colorado. Natural history inventory of Colorado No. 3., University of Colorado Museum. 20 pp. DEAN, CRYPTOGAMIC SOIL CRUSTS: RECOVERY FROM GRAZING NEAR CAMP FLOYD STATE PARK, UTAH, USA Jeffrey R. Johansen’ and Larry L. St. Clair® ABSTRACT. —The effects of grazing on the cryptogamic and vascular plant communities at two sites near Camp Floyd State Park, Utah County, Utah, were studied. The grazed site was subject to heavy grazing up until seven years prior to the study. The ungrazed site within the park boundaries had been protected from grazing for 20 years prior to the study and had a well-developed algal-lichen-moss crust. We found that the algae of the grazed site had recovered in terms of degree of crusting. There were no significant differences in the algal communities of the two sites when prevalent species were used as blocks in the ANOVAR analysis. However, when major algal groups were used as blocks, the analysis was significant, with the more recently grazed site having lower algal frequency. This difference, together with a few compositional differences, indicates that, although the algal community seven years following grazing is very similar to the algal community free of grazing for 20 years, the seven-year site is still in the process of recovery and community development. The diatom collections had a higher density in the grazed site, though the difference was not significant. Recovery of the lichen and moss community was not complete. The lichen Collema tenax and the mosses Bryum pallescens and Tortula ruralis were all significantly more abundant in the ungrazed area. Total cover of the lichen and moss components of the soil crusts was significantly lower in the more recently grazed area. Vascular cover was also lower. Cryptogamic soil crusts are an important component of many arid rangeland ecosys- tems in the western United States. Such crusts have been found to be important in nitrogen fixation (Snyder and Wullstein 1973, Rychert and Skujins 1974) and enhancement of seedling establishment (St. Clair et al. 1984). The greatest benefit of cryptogamic crusts, however, is probably reduction of soil erosion. Soil aggregation, particularly by blue-green algae, reduces detachment of soil particles by wind and rain (Bailey et al. 1973, Anantani and Marathe 1974). Improved water penetration in crusted soils reduces runoff and subsequent erosion (Brotherson and Rushforth 1983). Sedimentation is also re- duced by the increased tortuosity of surface water pathways due to the characteristic hum- mocking of desert crusts. The effects of both burning and grazing on soil cryptogams have recently been investi- gated. Range fires can severely damage all components of the soil crust (Johansen, Javakul, and Rushforth 1982, Johansen et al. 1984, Callison et al. 1985). Several workers have noted that moderate to heavy grazing can seriously damage soil cryptogams because the crusts are trampled by livestock. Rogers and ‘SER, 1617 Cole Blvd., Golden, Colorado 80401. Lange (1971) noted that stocking pressure was negatively correlated with lichen cover and that soil mobility and erosion was increased in areas of reduced lichen cover. Kleiner and Harper (1972) found that the effects of grazing were more notable on the soil cryptogams than on the vascular plant communities in Canyonlands National Park. They also noted that soil erosion was higher in areas with lower cryptogam cover and observed changes in or- ganic matter, available phosphorus, and cal- cium in eroded soils (Kleiner and Harper 1977). The effects of long-term moderate to heavy grazing near Navajo National Monu- ment also showed that grazing has a more pronounced effect on the cryptogamic cover and diversity than on vascular plant cover and diversity (Brotherson et al. 1983). Burros in the Grand Canyon are currently causing ero- sion through destruction of Tortula (bryo- phyte) and lichen crusts (Phillips et al. 1977). Several factors influence the development of cryptogamic crusts. Cryptogamic growth is best in soils of high electrical conductivity, high phosphorus, and high silt content (An- derson et al. Factors, 1982). Crust buildup is also positively correlated with soil alkalinity. The crusts of gypsiferous soils of southern “Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 632 October 1986 Utah are particularly well developed, having high lichen and algal diversity (Anderson and Rushforth 1976). Recovery of crusts from dis- turbance follows several trends. Algae are the most resistant component to disturbance (An- derson et al. Recovery, 1982) and are also the quickest to recover (Johansen et al. 1984). Lichens and mosses are slower to recover. Anderson et al. Recovery, (1982) found that lichens and mosses had become fairly well established after a 14~17-year period of pro- tection from grazing. Exclosures protected for 37-38 years showed little change in the moss flora, though lichen diversity was greater than in the 14~-17-year-old exclosures. This paper also indicated a need for studies examining recovery in the first 15 years following protec- tion from disturbance. The primary purpose of our study was to determine the degree of recovery of the cryp- togamic crust community in a soil that has been protected from grazing for seven years near Camp Floyd State Park. This study dif- fers from past studies in that the grazed area is compared with an adjacent area that has been protected for 20 years rather than with an area still under grazing pressure. SITE DESCRIPTION The study was conducted near and within the boundaries of Camp Floyd State Park, Utah County, Utah. The park is located 0.4 km southwest of Fairfield, Utah, along State Highway 73 in central Utah and is relatively small (16.2 hectares). It was established in 1962 with the principal point of interest being an inactive military cemetery which occupies only a small portion of the total area of the park. The balance of the park has not been developed and is dominated by an Atriplex confertifolia—Sarcobatus vermiculatus desert shrub community. A well-developed cryp- togamic soil crust flora consisting of various species of lichens, mosses, and algae is present. The soils at Camp Floyd belong to the Woodrow silt loam with an average water- holding capacity of 28-30 cm of water for the 1.5 m (5 ft) profile. This soil type has a slow permeability and is classified as a mixed (cal- careous) mesic, xeric torifluvent. The mean annual rainfail in this area is 35 cm. From 1935 until the establishment and sub- sequent fencing of the park area, the 1,600 JOHANSEN, ST. CLaiR: CRYPTOGAMIC SOIL CRUSTS 633 hectares of shrubland immediately around the cemetery were heavily grazed by sheep and cattle during winter months (October to May). Fencing of the park property has effec- tively eliminated grazing in the park since 1962. This has permitted the establishment of a diverse and well-developed algal-lichen- moss community. The area outside the park continued to be heavily grazed until 1975, at which time livestock were removed from this range. METHODS Two permanent transects with points placed every 2 m were established in the vicinity of Camp Floyd State Park. The un- grazed area transect consisted of 6 points within the park boundaries. Data from this transect were compared with similar data col- lected from burned sites, and that comparison is the basis of a previous paper (Johansen et al., 1984). The grazed area transect consisted of 8 points one mile south of the park. Field- work was conducted from September to Novemeber, 1982. Two crust samples were collected from each transect point at opposite compass points 0.25 m from the center of the point on 23 September 1982. Each sample was prepared for culturing by gently crushing soil clods to a maximum diameter of 5 mm. Twenty cm” of soil from each sample were placed in a steril- ized petri dish and saturated with 20 ml of distilled water. Samples were then incubated under continous cool-white fluorescent light at room temperature (24 C) for 10 days. Per- cent visible algal cover in each petri dish was estimated at the end of the culture period. Frequency and relative abundance of living algal species were also estimated at that time by subsampling the center of each petri dish and examining this subsample under the light microscope. A total of 25 microscope fields were examined for each subsample, and pres- ence and absence of each species in each field were noted. Permanent diatom mounts were prepared using standard acid oxidation proce- dures and Hyrax diatom mountant. Slides were prepared using quantitative dilutions for quantitative comparisons of soil diatoms (Jo- hansen et al. 1982). Visual estimates of total cryptogamic and vascular plant cover were made in the field 634 GREAT BASIN NATURALIST Vol. 46, No. 4 TABLE 1. Average percent frequencies of living algal species in grazed and ungrazed areas. Each species was tested using the Mann-Whitney test to determine if significant differences existed between treatments (* indicates p< .05, ** indicates p<.01). Estimated visible algal cover in culture dishes is also given. Species CYANOPHYTA Anabaena variabilis Kuetzing Aphanothece castagnei (Breb.) Rabh. Chroococcus minor (Kuetz.) Naegeli Chroococcus turgidus (Kuetz.) Naegeli Gloeothece linearis var. composita G. M. Smith Microcoleus vaginatus (Vauch.) Gomont Nostoc commune Vaucher Nostoc muscorum C. A. Agardh Nostoc cf. paludosum Kuetzing Nostoc species Phormidium minnesotense (Tild.) Drouet Synechococcus aeruginosus Naegeli Tolypothrix tenuis (Kuetz.) Schmidt Unknown Chroococcaceae CHLOROPHYTA Ulothrix tenerrima Kuetzing Unknown coccoids BACILLARIOPHYTA Hantzschia amphioxys (Ehr.) Grunow Navicula mutica Kuetzing Navicula paramutica Bock Pinnularia borealis Ehr. FLAGELLATES TOTAL SUM FREQUENCY VISIBLE ALGAL COVER SHANNON-WIENER DIVERSITY INDEX using a 1/4 m’ circular quadrat placed at each permanent transect point. In addition to the 1/4 m* circular quadrats, a smaller 2 x 50 cm rectangular quadrat consisting of ten 2 X 5cm subquadrats was placed perpendicular to the transect at each point and percent cover of algal crust and lichen and moss species was estimated. Cover was measured in November in the ungrazed area and in September in the more recently grazed area. Soils were analyzed by the Soil Analysis Laboratory, Department of Agronomy and Horticulture, Brigham Young University, for pH, phosphorus, nitrate, Kjeldahl nitrogen, potassium, calcium, Magnesium, zinc, iron, manganese, sodium, copper, organic matter, electrical conductivity, and sodium adsorp- tion ratio. All samples were taken from the top 17.5 cm of soil for these analyses. Mann-Whitney tests (Ryan et al. 1976) were performed for each species to determine significance levels of differences between grazed and ungrazed areas. This test was used in preference to the Students t-test (Snedecor Grazed Ungrazed 2x8) 4.0 1.0 3 1.0 3 2.0 ao 55.0 57.0 5) 10.0 * 3 8 ied 5.5 Toll 27.8 15.3 ate) 1.8 3a 1.5 4.0 3 3.0 16.3 23.0 18.5 26.0 5.0 12.3 1.0 3 8 7 3 Lod 137.0 173.3 - 16.6 BO0) Pd Bt5) Dor and Cochran 1980) because some of our data were not normal. Shannon-Wiener diversity indices were calculated for the living algae and diatom communities for each transect point (Patten 1962, Shannon and Weaver 1949). Similarity indices for all 14 stands were calculated following the methods of Ruzicka (1958). Four different sets of similarity indices were calculated using lichen and moss cover, living algal density, diatom density, and vas- cular plant cover. These four sets of indices were then clustered following Sneath and Sokal (1973) to illustrate the degree of similar- ity between grazed and ungrazed stands in the four different plant communities. Importance values for species of living algae and diatoms were determined by multiplying average per- cent relative density times presence (Warner and Harper 1972). Species with importance values above 1.00 were used in the subse- quent ANOVAR analyses. Multivariate analysis of variance adapted in preference to the Students t-test (Snedecor for a fixed effects, unbalanced design follow- October 1986 JOHANSEN, ST. CLAIR: CRYPTOGAMIC SOIL CRUSTS 635 ing the methods of Bryce et al. (1980) was used to analyze differences between blocks (lichens mosses, algae, vascular plants) and treatments (time since grazing). Several separate ANOVAR tests were run, including tests us- ing the vascular plant cover data, lichen and moss cover data, living algal data, and subfos- sil diatom data. In some data sets the variance of each species and group was related to the mean. To satisfy the homogeneity of variance assumption of analysis of variance for these data sets, a log (x+ 1) transformation was used (Bartlett 1947). With each analysis of vari- ance, the standardized residuals were plotted against the normal scores to give a measure of normality. In all cases the probability plot thus generated was subjectively judged as be- ing normal or close to normal. Our use of transects satisfied the requirements of sys- tematic sampling as described by Cochran (1977). The Duncan multiple range test was used to determine significance of differences between species means when analysis of vari- ance showed significance for this factor (Dun- can 1955). Unless otherwise stated, the alpha level used for this test was 0.01. RESULTS Living Algae The seven most important living algal spe- cies were Microcoleus vaginatus (importance value=55.86), Phormidium minnesotense (20.83), unknown green coccoid (18.46), Hantzschia amphioxys (17.06), Nostoc species (4.59), Navicula mutica (4.07), and Nostoc commune (1.42). Average percent frequencies for all algal species are given in Table 1. Multi- variate analysis of variance based on the above seven prevalent species showed that the algal communities of the two areas were not signifi- cantly different. The Mann-Whitney test on the visual algal cover estimated in the field supported this conclusion (Table 3). The mul- tivariate analysis did show that the species had significantly different frequencies. Duncan's test showed that Microcoleus vaginatus was significantly more abundant than all other al- gal species. Phormidium minnesotense, un- known coccoid green algae, and Hantzschia _ amphioxys were significantly more abundant _ than all less common algal species. The inter- action between treatment and species was sig- nificant (p=.023). This was likely due to the Fig. 1 60 ; ____—— Microcoleus vaginatus 50 40 30 LIVING ALGAE Hantzchia amphioxys Ba Coccoid green algae ————— - Phormidium minnesotense Navicula mutica 10 Nostoc commune Nostoc species PERCENT FREQUENCY OF 7 years 20 years TIME SINCE GRAZING Fig. 1. Interaction between living algal species and treatment, significant at p=.023, using multivariate anal- ysis of variance. fact that Hantzschia amphioxys, Navicula mu- tica, Nostoc commune, and coccoid green al- gae were all more abundant in the park area, whereas Phormidium minnesotense was more numerous in the more recently grazed stands (Fig. 1). Microcoleus vaginatus and Nostoc species showed only minor differences in fre- quency between the two sites. A separate analysis of variance was con- ducted to compare major algal groups. When using this data set, the difference between treatments was significant (p=.041), the more recently grazed plots having lower values. The groups were also significantly different (p<.001). Duncan’s test showed _ that Cyanophyta were significantly more abun- dant than the other three algal groups. Bacil- lariophyceae and Chlorophyta were signifi- cantly more abundant than unidentified flagellates. Nostoc commune had a significantly greater population in the 20-years-since-grazing area (p=.024) according to the Mann-Whitney test. No other living algal species or groups were significantly different at the two sites according to this test. Subfossil Diatoms The nine most important subfossil diatom species, including chrysophyte cysts ob- served in diatom mounts, were Hantzschia amphioxys (importance value 69.93), Navic- ula mutica (54.96), chrysophyte cysts (46.43), Pinnularia borealis (18.91), Navicula mutica var. cohnii (6.06), Navicula paramutica (3.65), Navicula mutica var. nivalis (2,42), Navicula contenta f. parallela (2.39), and Cy- 636 GREAT BASIN NATURALIST Vol. 46, No. 4 TABLE 2. Average densities (1,000 cells/cm) of diatom species in grazed and ungrazed areas. Each species was tested using the Mann-Whitney test to determine if significant differences existed between treatments (* indicates p<.05, ** indicates p<.01). Density of chrysophyte cysts is also given. Species Grazed Ungrazed Achanthes lanceolata Breb. 1 Anomoeneis species 1 Cocconeis placentula var. lineata (Ehr.) Cleve 5 1 Cyclotella kutzingiana Thwaites 39 11* Denticula elegana f. valida Pedic. 2 1 Diploneis oblongella (Naeg. ex Kuetz.) Ross 1 Epithemia adnata var. minor (P. & H.) Patr. 2} Epithemia turgida (Ehr.) Kuetzing 10 Js Fragilaria brevistriata Grunow 3 Fragilaria construens var. venter (Ehr.) Grunow 9 4 Fragilaria pinnata Ehr. 3 Hantzschia amphioxys (Ehr.) Grunow 1,250 562** Melosira dendroteres (Ehr.) Ross 2 Melosira granulata (Ehr.) Ralfs 3 9 Melosira species 2 Navicula contenta f. parallela Petersen 43 24 Navicula cuspidata (Kuetz.) Kuetzing 1 Navicula elginensis var. rostrata (Mayer) Patr. 1 Navicula excelsa Krasske 3 2) Navicula mutica Kuetzing 698 728 Navicula mutica var. cohnii (Hilse) Grunow 85 78 Navicula mutica var. nivalis (Ehr.) Hust. 46 21 Navicula paramutica Bock 82 19** Nitzschia paleacea 1 Pinnularia appendiculata (Ag.) Cleve : 1 Pinnularia borealis Ehr. 404 105** Pinnularia species : 1 Rhopalodia gibba (Ehr.) Mueller 1 Rhopalodia gibberula (Ehr.) Mueller 3 Stephanodiscus carconensis (Eul.) Grunow 1 Stephanodiscus hantzschii Grunow 1 Stephanodiscus species 2 TOTAL DIATOMS 2,690 1,570 TOTAL CHRYSOPHYTE CYSTS 675 530 SHANNON-WIENER DIVERSITY INDEX 1.94 2.14* Fig. 2 2000 Bryum pallescens Igoe Navicula mutica Bee ee amphioxys Collemaltenax Chrysophyte cysts Tortula ruralis We Pinnularia borealis Navicula mutica var. cohnii ESE A SE desi Ss ee contenta f. parallela Navicula mutica var. nivalis yO ff oO (eo) (1000 CELLS/CM) oO (@) Navicula paramutica Cyclotella kuetzingiana Dermatocarpon lachneum Caloplaca tominii DENSITY OF SUBFOSSIL DIATOMS CRYPTOGAMS-PERCENT COVER 7 years 20 years 7 years 20 years TIME SINCE GRAZING TIME SINCE GRAZING Fig. 2. Interaction between subfossil diatoms and Fig. 3. Interaction between cryptogam species and treatment, significant at p<.001, using multivariate anal- treatment, significant at p<.001, using multivariate ysis of variance. analysis of variance. October 1986 JOHANSEN, ST. CLatr: CryPTOGAMIC SOIL CRUSTS 637 TABLE 3. Average percent cover values in grazed and ungrazed areas. Each species the Mann-Whitney test to determine if significant differences existed between tre indicates p<.01). and cover class was tested using atments (* indicates p<.05, ** Species Grazed Ungrazed Cover based on 100 cm’ rectangular quadrats LICHENS Caloplaca tominii Sav. A p* Collema tenax (Sw.) Ach. 3.0 6.2** Dermatocarpon lachneum (Ach.) A. L. Sm. Le l** Lecidea decipiens (Hedw.) Ach. P TOTAL LICHEN COVER Sl! 6.4 MOSSES Bryum pallescens Schwaeg. 2.0 oo Pterygoneurum lamellatum (Lindb.) Jur. P Tortula ruralis (Hedw.) Gaertn., Meyer & Scherb. 3 6.1** TOTAL MOSS COVER 28 13.4** TOTAL ALGAL COVER 21.0 22.6 TOTAL CRYPTOGAMIC CRUST COVER 28.4 42.4 Sarcobatus vermiculatus I Fig. 4 = 30 WwW 2 25 a | A. XO Seedlings eam SS 15 te) z 10 fs) = Sitanion hystrix o 5 Chrysothamnus nauseosus 2 Atriplex confertifolia > 7 years 20 years TIME SINCE GRAZING Fig. 4. Interaction between vascular plant species and treatment, significant at p=.001, using multivariate analysis of variance. clotella kuetzingiana (1.52). Average densities of all diatom species are given in Table 2. Multivariate analysis based on the data for these nine taxa showed that diatoms were sig- nificantly more abundant in the recently grazed area (p<.001). Species were also sig- nificantly (p<.001) different. According to Duncan's test, Hantzschia amphioxys, Navic- ula mutica, and chrysophyte cysts were all significantly higher than the other diatom taxa. Pinnularia borealis was significantly more abundant than all less abundant species. Navicula mutica var. cohnii was more abun- dant than the other three prevalent species used in the analysis. The interaction between species and treatment (p<.001) is illustrated in Figure 2. Lichens and Mosses Multivariate analyses demonstrated several important differences among treatments, blocks, and interactions in the lichen and moss communities. Total lichen and moss cover was substantially greater in the park area (p<.001). Species of cryptogams had sig- nificantly (p<.001) different cover. Duncan’s test showed that Collema tenax and Bryum pallescens were significantly more abundant than the other taxa. Tortula ruralis cover was significantly greater than Caloplaca tominii cover. The interaction between species and treatment was significant (p<.001). This was due to the fact the Bryum pallescens, Tortula ruralis, and Collema tenax had higher cover in the park area, whereas Caloplaca tominii and Dermatocarpon lachneum were most abundant in the more recently grazed area (Fig. 3). A separate analysis of variance was run us- ing total lichen and moss cover data. This test also showed a significant difference between treatment means (p<.001). The interaction between cryptogam class and treatment was significant (p<.001) because lichen cover is only slightly greater in the park area and moss cover is markedly greater in the park area. The Mann-Whitney test supports this conclu- sion in that the difference in total moss cover between treatments is significant, but the dif- ference between mean lichen cover is not (Table 3). 638 GREAT BASIN NATURALIST Vol. 46, No. 4 TABLE 4. Average percent cover values in grazed and ungrazed areas. Each species and cover type was tested using the Mann-Whitney test to determine if significant differences existed between treatments (* indicates p<.05, ** indicates p<.01). Cover type Grazed Ungrazed Cover based on 1/4 m* circular quadrats Bare ground Pla 4.3* Litter 26.9 23.3 Cryptogamic crust 28.1 38.3 Total vascular plant cover BLES 132 Atriplex confertifolia (Torr. & Frem.) S. Wats. 2s 4.3 Chrysothamnus nauseosus (Pall.) Britt. 6.0 Sarcobatus vermiculatus (Hook.) Torrey 12.8 34.2 Sitanion hystrix (Nutt.) J. G. Smith el 7.8 Vascular seedlings 3 ZOE es TABLE 5. Average nutrient levels and soil characteris- tics of soil samples taken from grazed and ungrazed areas. Each factor was tested using a two-tailed t-test to deter- mine if significant differences existed between treatments (* indicates p<.05). Soil factor Grazed Ungrazed | pH 7.98 C1 Phosphorus (ppm) 20.8 20.2 Nitrate-N (ppm) 6.03 6.30 Potassium (ppm) 944 743* Calcium (ppm) 56.2 72.8 Magnesium (ppm) 8.4 SK) Copper (ppm) 1.14 1.15 Zinc (ppm) 0.68 0.79* Iron (ppm) 2.03 2.94* Manganese (ppm) 2.34 OESSs Sodium (ppm) 66 209* Total organic matter (%) 1.30 ell Total nitrogen (%) 0.100 0.107 Electrical conductivity x 1000 0.91 1.81 Sodium adsorption ratio 0.41 1.54* Multivariate analysis of the vascular plant community data showed that cover was signif- icantly greater in the area protected from grazing for 20 years (p<.001). Species were also significantly different (p=.006), though only the means for Sarcobatus vermiculatus and Chrysothamnus nauseosus were signifi- cantly different according to Duncan’s test. The interaction between treatment and spe- cies was also significant (p=.001) and is illus- trated in Figure 4. Average percent cover values for all vascular species are given in Table 4. The soils of the two sites were similar to each other (Table 5) though cations were gen- erally higher in the park area. Students t-tests showed that the sites were significantly differ- ent in ppm K, Zn, Fe, Mn, Na, and sodium adsorption ratio. Except for potassium, the ungrazed site had greater values for all the above. DISCUSSION It is apparent from the data that the algal community, in terms of both the living algae and subfossil diatoms, has nearly recovered from the influence of grazing in the more re- cently grazed area. Visual estimates in the field as well as microscopic examinations in the laboratory support this conclusion. When the similarity indices for the transect points were clustered, all points were similar in re- gard to both living algae and subfossil di- atoms. The greater density of subfossil di- atoms in the more recently grazed area is difficult to explain. It may be due to the lower vascular plant density in the grazed area, which in turn results in increased light inten- sity and lower litter cover. These factors in combination could favor diatom growth. The dominant lichen and moss species of this area, Collema tenax, Bryum pallescens, and Tortula ruralis , have not fully recovered in the grazed area. An unusual observation was the significantly greater amount of the lichen Dermatocarpon lachneum in the more recently grazed area. We hypothesize that this may indicate an intermediate succes- sional stage in the recovery process. Factors contributing to this phenomenon may include compositional differences in both the vascular and nonvascular plant communities as well as biological modifications of local abiotic factors such as light and moisture. This hypothesis will be tested by future monitoring of the lichen and moss community at these two sites. The greater density of the three dominant lichen and moss taxa in the ungrazed area ; October 1986 JOHANSEN, ST. CLAIR: CRYPTOGAMIC Som CRruSTS 639 Percentage Similarity 100 90 80 Ungrazed Sites Ugr Ugr Ugr Ugr Ugr U 70 60 50 40 30 20 | 59% gr Gr Grazed Sites Gr Gr Gr Gr 26% Gr Gr or 31% Fig. 5. Fourteen transect points clustered on the basis of Ruzicka’s similarity index as computed from lichen and moss data. All stands were similar, yet still clustered into two main groups, i.e., grazed and ungrazed sites. combined with the significantly greater cover of D. lachneum in the more recently grazed area resulted in the formation of discrete site groupings when similarity indices of transect points were clustered (Fig. 5). However, the two clusters based on algal data did not form such discrete clusters, probably because of the high level of similarity between all points. This demonstrates that the algal community has essentially recovered in the seven years following removal of livestock from the range, whereas the lichen and moss communities are still in the process of recovery. . Anderson et al. Recovery, (1982) indicate that reestablishment of cryptogamic soil crusts is substantial after 14-18 years. Our study indicates that the recovery rate of the algae is much more rapid than estimated in | their study, occurring in fewer than 7 years. The lichens and mosses, on the other hand, fit the predicted recovery patterns and apparently require more than 7 years to fully recover. An important dimension in the develop- ment of soil crusts is frequency and abun- dance of moisture. The annual precipitation in Utah County has been above normal for the past three years and has undoubtedly played a role in the reestablishment of the soil crusts at the disturbed sites near Camp Floyd. Subjec- tive observations of the grazed site in 1981 indicated that differences between visible cryptogamic cover were evident. Noticeable recovery of the crust in the grazed area oc- curred during the ensuing moist year before the present study was undertaken. It is possi- ble that in drier areas or drier years develop- ment of cryptogamic crusts following grazing disturbance might take longer than the seven- year period observed in the present study. The greater cover of vascular seedlings ob- served in the ungrazed area was likely due to temporal differences in sampling. The more recently grazed area was examined in Sep- tember, whereas the ungrazed area was sam- 640 pled in November after two months of mild, moist weather. A noteworthy difference in vascular plant community structure was ob- served. Atriplex confertifolia populations were substantially greater in the grazed area (Fig. 4). On the other hand, Sarcobatus ver- miculatus was most dense in the ungrazed area. Sheep have been known to browse both Sarcobatus and Atriplex. Thus, over a period of 40 years the abundance of both taxa could have been severely reduced in the grazed area. With the end of grazing pressure, Sarco- batus vermiculatus , a vigorous root sprouter, has begun reestablishing itself. In the area protected for 20 years, the development of the Sarcobatus population has possibly pro- ceeded to the point that Atriplex is being crowded out. In the more recently grazed area the vascular cover is less abundant, and Atriplex has not been excluded. ACKNOWLEDGMENTS This research was supported by the Shrub Sciences Laboratory, Forest Service, USDA, Provo, Utah 84601. Dr. Brent Mishler of Duke University kindly assisted in identifying our bryophyte material. LITERATURE CITED ANANTANL, Y. S., AND D. V. MARATHE. 1974. Soil aggregat- ing effects of some algae occurring in the soils of Kutch and Rajasthan. J. Univ. Bombay 41(68): 94-100. ANDERSON, D. C., AND S. R. RUSHFORTH. 1976. The cryp- togam flora of desert soil crusts in southern Utah, USA. Nova Hedwigia 28: 691-729. ANDERSON, D. C., K. T. HARPER, AND R. C. HOLMGREN. 1982. Factors influencing development of cryp- togamic soil crusts in Utah deserts. J. Range Man- age. 35: 180-185. ANDERSON, D. C., K. T. HARPER, AND S. R. RUSHFORTH. 1982. Recovery of cryptogamic soil crusts from grazing on Utah winter ranges. J. Range Manage. 35: 355-359. BaILey, D., A. P. MAZURAK, AND J. R. Rosowski. 1973. Aggregation of soil particles by algae. J. Phycol. 9: 99-101. BARTLETT, M. S. 1947. The use of tranformations. Biomet- rics 3: 39-52. BROTHERSON, J. D., AND S. R. RUSHFORTH. 1983. Influence of cryptogamic crusts on moisture relationships of soils in Navajo National Monument, Arizona. Great Basin Nat. 43: 73-78. BROTHERSON, J. D., S. R. RUSHFORTH, AND J. R. JOHANSEN. 1983. Effects of long-term grazing on cryptogam crust cover in Navajo National Monument. J. Range Manage. 36: 579-581. GREAT BASIN NATURALIST Vol. 46, No. 4 Bryce, G. R., D. T. Scott, AND M. W. Carter. 1980. Estima- tion and hypothesis testing in linear models—a reparameterization approach to the cell means model. Comm. Stat.-Theor. Meth. A9(2): 131-150. CALLISON, J., J. D. BROTHERSON, AND J. E. Bowns. 1985. The effects of fire on the blackbrush (Coleogyne ramosis- sima) community of southwestern Utah. J. Range Manage. 38: 535-538. Cocuran, W. G. 1977. Sampling techniques. 3d ed. John Wiley and Sons, Inc., New York. 428 pp. Duncan, D. B. 1955. Multiple range and multiple F tests. Biometrics 11: 1-42. JOHANSEN, J. R., A. JAVAKUL, AND S. R. RUSHFORTH. 1982. The effects of burning on the algal communities of a high desert soil near Wallsburg, Utah, USA. J. Range Manage. 35: 598-600. JOHANSEN, J. R, L. L. St. Crate, B. L. Wess, AND G. T. NEBEKER. 1984. Recovery patterns of cryptogamic soil crusts in desert rangelands following fire disturbance. Bryologist 87: 238-243. KLEINER, E. F., AND K. T. HarPeR. 1972. Environment and community organization in grasslands of Canyonlands National Park. Ecology 53: 229-309. KLEINER, E. F., AND K. T. Harper. 1977. Soil properties in relation to cryptogamic ground cover in Canyonlands National Park. J. Range Manage. 30: 202-205. PATTEN, B. C. 1962. Species diversity in net phytoplankton of Raritan Bay. J. Marine Res. 20: 57-75. PuILuips, A. M., IIL, L. T. GREEN, AND G. A. RUFFNER. 1977. Investigations of feral burro impact on plant commu- nities, Grand Canyon, Arizona. Pages 20-99 in G. A. Ruffner, A. M. Phillips, II, and N. H. Goldberg, eds., Biology and ecology of feral burros (Equus asi- nus ) at Grand Canyon National Park, Arizona. Report submitted to National Park Service, Grand Canyon National Park, Arizona 86023. ROGERS, R. W., AND R. T. LANGE. 1971. Lichen populations on arid soil crusts around sheep watering places in South Australia. Oikos 22: 93-100. Ruzicka, M. 1958. Anwendung mathematisch-statistischer Methoden in der Geobotank (synthetische Bearbei- tung von Aufnahmen). Biologia Bratisl. 13: 647-661. Ryan, T. A., B. L. JOINER, AND B. F. Ryan. 1976. Minitab stu- dent handbook. Duxbury Press, North Scituate, Mas- sachusetts. 341 pp. RYCHERT, R. C., AND J. SkujINs. 1974. Nitrogen fixation by blue-green algae-lichen crusts in the Great Basin desert. Soil Sci. Soc. Amer. Proc. 38: 768-771. SHANNON, G. E., AND W. WEAVER. 1949. The mathematical theory of communication. University of Illinois Press, Urbana. 117 pp. SNEATH, P. H., AND R. R. SOKAL. 1973. Numerical taxonomy. W. H. Freeman and Co., San Francisco. 573 pp. SNEDECOR, G. W., AND W. G. CocHRAN. 1980. Statistical meth- ods. 7th ed. Iowa State University Press, Ames. 507 PP. SNYDER, J. M., AND L. H. WULLSTEIN. 1973. The role of desert cryptogams in nitrogen fixation. Amer. Nat. 90: 257-265. St. Ciair, L. L., B. L. Wess, J. R. JOHANSEN, AND G. T. NEBEKER. 1984. Cryptogamic soil crusts: enhance- ment of seedling establishment in disturbed and undisturbed areas. Reclam. Reveg. Res. 3: 129-136. WaRNER, J. H., AND K. T. HARPER. 1972. Understory character- istics related to site quality for aspen in Utah. Brigham Young University Sci. Bull., Biol. Ser. 16(2): 1-20. NEW SPECIES AND NEW RECORDS OF NORTH AMERICAN PITYOPHTHORUS (COLEOPTERA: SCOLYTIDAE), PART VI. THE LAUTUS GROUP Donald E. Bright! ABSTRACT. —Four new species of Mexican Pityophthorus in the Lautus group are described: P. indefessus (Jalisco), P. inhabilis (Guerrero), P. tutulus (Veracruz), and P. vegrandis (Quintana Roo) and a new locality record is given for P. corruptus Wood. This is the third paper describing the previ- ously unnamed species of Pityophthorus col- lected by Dr. T. H. Atkinson and his col- leagues (Centro de Entomologia y Acaro- logia, Colegio de Postgradados, Chapingo, Mexico). The present contribution describes four species in the Lautus group and gives new locality records for one species in the group. As in the previous papers (Great Basin Nat. 45: 467-482), the key in my 1981 mono- graph (Mem. Ent. Soc. Canada 118, pp. 54, 50) is modified to accommodate the newly named species. I thank Dr. Atkinson for sending the speci- mens to me and also thank him and his stu- dents for their diligent searching for Scolyti- dae in previously unrecognized host plants. I also thank my colleagues Dr. Y. Bousquet and Dr. L. LeSage for reviewing the manuscript. Pityophthorus indefessus , n.sp. Length 1.3-1.4 mm, 2.7 times longer than wide. Frons transversely impressed from epis- tomal margin to upper eye level, impression moderately deep, with obscure, weakly ele- vated, impunctate median carina extending from epistoma halfway or less across impres- sion; surface shining, finely, densely punc- tured, setae short, inconspicuous. Antennal club 1.1 times longer than wide, widest through third segment; first two sutures weakly arcuate, almost invisible except where heavily sclerotized at lateral margins; first two segments occupy about one-third of total club length. Pronotum slightly more than 1.1 times longer than wide, widest at level of summit; sides very weakly arcuate, weakly converging on posterior half, broadly rounded anteriorally, anterior margin with about nine distinct serrations; anterior slope with three irregular rows of asperities, these rows some- what broken, several obscure additional rows around summit; summit weakly elevated; pos- terior area of disc with numerous fine, shallow punctures, these separated by distance equal to or less than their diameters, surface be- tween punctures shining, smooth with nu- merous, very fine impressed points; median line obscure, narrow, impunctate. Elytra 1.6 times longer than wide, apex very broadly rounded, almost truncate; discal striae punc- tured in regular rows, punctures fine, shal- low, slightly larger than those on posterior portion of pronotum; interstriae about twice as wide as striae, moderately shining, with numerous, very fine, minute points, 3, 5, 7 with three or four erect, flattened scales on posterior half, 1 with row of four or five scales extending to base. Declivity convex, flat- tened; interstriae 1 elevated, with median row of fine granules and short, fine setae; interstriae 2 weakly impressed, with median row of very fine setae and granules, these sometimes evident only on upper half of inter- stria; interstriae 3 very weakly elevated, with median row of distinct, fine granules and erect, spatulate setae, these longer than setae on | and 2; remaining interstriae (3, 5, 7) with a median row of erect spatulate setae; punc- tures in striae | and 2 distinct. TYPE MATERIAL.—The holotype is labeled: “Estacién Bioldgica, Chamela, Edo. Jalisco, 7.111.82, S-390, 80 msnm, Col. Armando Equihua’/“HOLOTYPE Pityophthorus inde- fessus D. E. Bright, 1986, CNC 18747.” Two 1Biosystematics Research Institute, Agriculture Canada, Ottawa, Canada K1A O0C6. 641 642 paratypes bear the same locality data plus paratype labels. The holotype is in the Cana- dian National Collection; the two paratypes were returned to T. H. Atkinson. COMMENTS. —The sexes of the three speci- mens in the type series could not be deter- mined since, in this group, sex can only be estabilished by examining the abdominal ter- gites. This was not done on the three speci- mens at hand. The specimens in the type series were found in a sample of P. molestus Wood and are very similar to that species except that declivi- tal interstria 2 of P. indefessus bears a row of fine granules and very fine setae, these present or most obvious on upper half of the interstria. This species will key to near P. nemoralis Wood and P. concentralis but may be distinguished by the characters mentioned in the key. Pityophthorus inhabilis , n.sp. Length 1.8—2.0 mm, 3.0 times longer than wide. FEMALE.—Frons flattened on semicircular area extending laterally from eye to eye and longitudinally from epistoma to well above upper eye level; surface densely, closely punctured in flattened area with brush of dense, erect setae all of equal or nearly equal length, surface above and lateral to flattened area shining, glabrous, with much larger, deeper, sparser punctures. Antennal club oval, about 1.4 times longer than wide, widest through second segment; suture 1 transverse, heavily sclerotized except for short space in middle, 2 transverse, lightly sclerotized only at lateral margin; segments 1 and 2 occupy more than two-thirds of total club length, su- ture 1 located just below middle of club, 2 located just below apex of club. Pronotum 1.1 times longer than wide, widest at posterior angles; sides slightly converging to broadly rounded anterior margin; asperities on ante- rior slope arranged into 4—6 or more irregular concentric rows, first row more regular, re- maining rows slightly broken; summit dis- tinct; posterior area of disc moderately shin- ing, punctures large, deep, close, inter- puncture space with numerous, fine, im- pressed points; median line narrow, not ele- vated, impunctate. Elytra 1.8 times longer than wide; apex narrowly rounded, elevated interstriae | extending slightly beyond elytral GREAT BASIN NATURALIST Vol. 46, No. 4 outline; discal striae punctured in regular rows, punctures large, larger than those on posterior portion of pronotum, deeply im- pressed; discal interstriae about as wide as or narrower than striae, moderately shining, glabrous, with numerous, fine, impressed points. Declivity almost evenly convex, weakly bisulcate; interstriae 1 wide, distinctly elevated, with median row of fine, shallow punctures and short, fine setae; interstriae 2 weakly impressed, flat, glabrous, slightly wider than strial width; interstriae 3 very weakly elevated, with several, large punc- tures and short, fine setae; striae 1 narrowly impressed, punctures obscure, 2 distinctly punctured, slightly curved in middle; vesti- ture in remaining interstriae consisting of fine setae near declivity. MALE.—Frons weakly transversely im- pressed to upper level of eyes; surface of im- pressed area densely, coarsely punctured, se- tae absent except along epistomal margin, surface above and iateral to impression more deeply, less closely punctured. Antennal club with first suture slightly closer to base than on female, second suture very weakly indicated near apex. Pronotum and elytra essentially as described for female. Declivity slightly more deeply bisulcate, otherwise as described for female. TYPE MATERIAL.—The holotype (@) is la- beled: “Chilapa, Guerrero, 23-II-82, 1800 m, S-337, Col. Atkinson y Equihua’/“HOLO- TYPE Pityophthorus inhabilis D. E. Bright, 1986, CNC No. 18447.” The allotype and six paratypes bear the same locality label plus the appropriate type label. The holotype, allotype, and two paratypes are in the Canadian National Collection; four paratypes were returned to Dr. Atkinson. COMMENTS.—Compared to adults of other species in the Lautus group, those of this spe- cies are unique by having a more broadly sloping elytral declivity on which the first and second striae are distinct, by having distinct sexual dimorphism on the frons, and by the unique antennal club on which the first two segments occupy almost the entire face of the club. The first antennal suture is located near the middle of the club and is distinctly scler- otized; the second suture is located just before the apex of the club and is weakly sclerotized and obscure. Other characters of species in the Lautus group, such as the concentric rows October 1986 of pronotal asperities and the distinctly punc- tured first and second declivital striae, are all present on adults of this species. Pityophthorus tutulus, n.sp. Length 1.5-1.8 mm, 2.8 times longer than wide. FEMALE.—Frons_ broadly flattened to weakly concave from eye to eye and from epistoma to well above eyes; surface on lower half smooth, moderately shining, sometimes with few, minute, impressed points and few, scattered setae, upper half with dense cover- ing of very short, stout, recumbent scales, periphery of flattened area with row extend- ing from eye to eye of long, incurved setae. Antennal club large, elongate-oval, 1.5 times longer than wide, widest through segment 3; suture | weakly arcuate, 2 transverse, both sclerotized, 2 more so than 1; segments 1 and 2 together occupy about one-third of total club length. Pronotum 1.1 times longer than wide, widest at middle; sides weakly arcuate, feebly constricted before broadly rounded anterior margin; asperities on anterior slope arranged into three distinct and one or two indistinct concentric rows; these rows may be broken, especially in median area; summit distinct; posterior area of disc moderately shining, punctures of moderate size, deep, distinct, close, interpuncture space with nu- merous, distinct, minute, impressed points; median line broad, not elevated, with numer- ous impressed points. Elytra 1.6 times longer than wide; apex broadly rounded; discal striae punctured in regular rows, punctures large, larger than those on posterior portion of pronotum, deeply impressed, close; discal in- terstriae about as wide or slightly narrower than striae, surface moderately shining, glabrous, with numerous fine points and lines. Declivity steep, convex; interstriae 1 very slightly impressed below level of 3 on upper half, with median row of fine granules; interstriae 2 flat, as wide as on disc, weakly but distinctly impressed below 1, surface smooth, glabrous; interstriae 3 weakly ele- vated, with median row of fine granules; striae 1 narrow, distinctly impressed, 2 slightly less deeply impressed, both straight and distinct; vestiture consisting of fine setae on lateral interstriae and in all interstriae, except 2, near declivity. MALE.—Frons weakly concave, upper mar- BRIGHT: AMERICAN PITYOPHTHORUS 643 gin of concavity arcuate, extending above up- per level of eyes; surface minutely punctate, with few, scattered, fine setae. Otherwise es- sentially as in female. TYPE MATERIAL.—The holotype (@) is la- beled: “Jalapa, Veracruz, 28-XI-83, FANM 100, col. Felipe A. Noguera’/“Hosp. Rhus radicans (Anacardiaceae)’/“HOLOTYPE Pityophthorus tutulus D. E. Bright, 1986, CNC No. 18448.” The allotype and_ six paratypes bear the same locality and host data plus the appropriate type labels. The holotype, allotype, and two paratypes are in the Canadian National Collection; four paratypes were returned to Dr. Atkinson. COMMENTS.—This species and P. crinalis are unique among North American species of the genus in that the upper half of the female frons has a dense brush of numerous short, recumbent, plumose scales. This brush ex- tends from eye to eye and has a fringe of much longer, incurved plumose setae on the upper margin (see figure 37 in my 1981 monograph). The lower half of the female frons is smooth, shining, and glabrous. The males of these two species differ from those of other species in the group only in minor details. Both species occur in Rhus spp. Adults of P. tutulus differ from those of P. crinalis by the slightly larger body size, by the larger antennal club, by the slightly larger granules on declivital interstriae 1 and 3, by the slightly more deeply impressed elytral declivity, and by the distribution. Pityophthorus vegrandis, n.sp. Length 1.0-1.1 mm, 2.7 times longer than wide. FEMALE.—Frons evenly convex, very weakly transversely, narrowly flattened just above epistoma; surface dull, densely micro- reticulate, with very faint, shallow, scattered punctures, setae absent except along epis- tomal margin. Antennal club oval, 1.4 times longer than wide, widest through segment 3; suture 1 moderately arcuate, sclerotized through entire length, suture 2 transverse to weakly arcuate, sclerotized at lateral margins; segments | and 2 together occupy about one- half of total club length. Pronotum as long as wide, widest at level of summit; asperities on anterior slope arranged into three even con- centric rows, one very faint additional row may be detected around summit; summit dis- 644 tinctly elevated; posterior area of disc smooth, dull, densely microreticulate, with large, shallow, widely separated punctures; median line broad, impunctate, reticulate. Elytra about 1.6 times longer than wide; apex broadly rounded; discal striae punctured in even rows, punctures fine, shallow, smaller than those on posterior portion of pronotum; interstriae about 1.5 times wider than striae, surface smooth or weakly reticulate, shining, without setae. Declivity convex, steep; inter- striae 1 and 3 equal in height, both with me- dian row of very fine granules; interstriae 2 flat, equal to discal width, weakly impressed below level of 1 and 3; striae 1 and 2 weakly impressed, 1 more strongly so; scattered setae present in all interstriae except 2. MALE.—Virtually identical to female ex- cept frons very weakly flattened, with dis- tinct, large, deeply impressed punctures. TYPE MATERIAL.—The holotype (@) is la- beled: “Chetumal, Quintana Roo, 10-Julio- 1982, 20 m, SM-020, E. Martinez’ /“HOLO- TYPE Pityophthorus vegrandis D. E. Bright, 1986, CNC No. 18449.” The allotype and one paratype bear the same locality label plus the appropriate type label. One damaged speci- men, not designated as a paratype, is labeled: “Laguna de Bacalar, Quintana Roo, 10-Julio- 1982, 20 m, SM-020, E. Martinez.” The holotype and allotype are in the Cana- dian National Collection; the two paratypes were returned to Dr. Atkinson. COMMENTS.—The relationships of this spe- cies are unclear. Although it keys to near P. sambuci, the two are not closely related. Adults are most easily distinguished by the small size, by the dull, densely, minutely reticulate frons of both sexes, by the very weakly impressed elytral declivity, and by the weak development of sexual dimorphism. Revised key to species in the Lautus group il. Male and female frons similar, pubescence SParse} caetasrtaie Ae Ey ear ee een eee 2) — Male and female frons sexually dimorphic, female frons distinctly pubescent, male frons only sparsely pubescent ................. 11 2(1). +Declivital interstriae 2 bearing median row of fine punctures or fine setiferous granules and fine setae; antennal club narrowly oval, about 1.5 times longer than wide ................ 3 — Declivital interstriae 2 never bearing gran- ules or setae; antennal club broadly oval, less than 1.5 times longer than wide ............ 5 GREAT BASIN NATURALIST 3(2). Vol. 46, No. 4 Declivital interstriae 2 bearing a median row of fine setae, these as long as those on inter- striae | and 3; surface between punctures on pronotum strongly reticulate; Honduras to CostaiRicay aes. de eee nemoralis Wood Declivital interstriae 2 bearing a median row of fine granules or punctures and fine setae, setae much shorter than those on interstriae 3 (and sometimes 1); surface between punc- tures on pronotum smooth or with fine points, brightly/shining(- =. eee 4 Declivital interstriae 2 bearing a median row of fine punctures and extremely fine hairlike setae; setae on declivital interstriae 3-9 all hairlike; surface between punctures on pro- notum smooth; frons without longitudinal carina above epistoma; Florida and Cuba ... concentralis Eichhoft Declivital interstriae 2 bearing a median row, at least on upper half, of fine granules and fine, flattened setae; setae on declivital inter- striae 3-9 spatulate; surface between punc- tures on pronotum with numerous fine points; frons with a weak longitudinal carina extend- ing from epistoma halfway to upper eye level; Jalisco. (222). eae indefessus Bright Frons bearing weak but distinct, longitudinal carina or elevation; punctures on posterior portion of pronotum numerous, small, and shallow (except borrichiae) ............... 6 Frons without indication of carina, sometimes bearing very small tooth on epistomal margin; punctures on posterior portion of pronotum large, deep, and widely spaced ........... 10 Frons flattened or transversely concave to up- per level of eyes, divided by weak, longitudi- nal, narrow elevation; declivity sloping; as- perities on anterior pronotal slope arranged into broken concentric rows ............-.- 7 Frons convex, usually with distinct, narrow elevation extending from epistoma to vertex, elevation interrupted in center by weak, transverse impression; if elevation absent, then frons rugose, elevation frequently indi- cated by small, elongate callus at upper level of eyes; declivity steep; asperities on anterior pronotal slope arranged in even, concentric TOWS sad St aqeishs dbo ues Syn eee 8 Occurs in eastern United States; setae on de- clivital interstriae about 1.5 times longer than interstrial width; median elevation on frons only weakly indicated ......... lautus Eichhoff Occurs in eastern Mexico; setae on declivital interstriae longer, more than 2.0 times longer than interstrial width; median elevation on frons sharply elevated ......... molestus Wood Body length 1.0-1.3 mm; declivital setae stout, about equal in length to interstrial width wil onidaaen esa borrichiae Wood Body length 1.4-1.7 mm; declivital setae fine, hairlike, nearly 2.0 times longer than interstrial width; Mexico and Central America .......... 9 October 1986 9(8). 10(5). 11(1). 12(11). 13(12). ’ Frons shining, deeply punctured, frontal ele- vation not evident but frequently indicated by elongate callus at upper level of eyes; discal interstriae smooth, with sparse, minute points; Chiapas to Honduras ... morosus Wood Frons dull, reticulate, sparsely punctured, el- evation usually distinct but frequently inter- rupted in middle by weak, transverse, densely punctured impression; discal inters- triae with numerous fine lines, surface irregu- lar wMlexicomenat csc ee patulus Wood Frons evenly convex or weakly flattened, sur- face dull, microreticulate; surface between punctures on posterior portion of pronotum densely reticulate; length 1.0 mm; Quintana Roo vegrandis Bright Frons flattened, usually with small tooth or weak elevation on epistomal margin, surface shining, smooth; surface between punctures on posterior portion of pronotum shining, smooth or faintly reticulate; length 1.4-1.7 avons |AIICO Sy 00b0000n0600 sambuchi Blackman Female frons densely pubescent only on up- per margin above upper level of eyes, shining and glabrous below; male frons flattened, densely punctured 12 Female frons pubescent over entire area be- tween eyes, setae may be longer, more abun- dant on periphery of flattened area; male frons weakly transversely impressed 15 First two segments of antennal club occupy more than two-thirds of total club length; de- clivity very weakly bisulcate, interstriae 2 widened, flat; male frons weakly transversely impressed, setae sparse; female frons flat- tened with erect setae, all of equal length; Guerrero inhabilis Bright First two segments of antennal club occupy less than two-thirds of total club length; de- clivity variable, not as above; male frons vari- able; female frons variable, with setae in vari- ous patterns but not as above 13 First two segments of antennal club occupy more than half of total club length, club 1.4 times or less longer than wide; lower half of female frons weakly but distinctly punctured, punctures rather large, upper margin with fringe of plumose setae; southeastern USA. . liquidambaris Blackman BRIGHT: AMERICAN PITYOPHTHORUS 16(15). 645 First two segments of antennal club occupy about one-third of total club length, club 1.5 times longer than wide; lower half of female frons smooth, brightly shining, sometimes with minute punctures, upper area with dense recumbent scales in addition to setae . 14 . Occurs in eastern USA; length 1.3-1.6 mm . crinalis Blackman Occurs in southern Mexico; length 1.5-1.8 TAM sce acme eee em eeaeee tutulus Bright Occurs in Central America; setae on declivital interstriae scalelike in male, hairlike in fe- male; pubescence on female frons abundant on periphery, sparse in central area ....... perexiguus Wood Occurs in Mexico; setae on declivital inters- triae as above or hairlike in both sexes; pubescence on female frons variable 16 Body size 0.8-1.6 mm; female frons pubes- cent on narrowly oval, median area, all setae of equal length; granules on declivital inters- triae 3 large, setae on declivity scalelike in male, hairlike in female; Mexico Pea clonontoknn Caen Ine attenuatus Blackman Body size 1.5-1.8 mm; female frons pubescent on broad area extending from eye to eye and to above upper eye level; central portion of female frons less densely pubes- cent, setae on periphery longer; granules on declivital interstriae 3 small; setae on decliv- ity hairlike in both sexes corruptus Wood NEW RECORD Pityophthorus corruputus Wood Pityophthorus corruptus Wood, 1976, Great Basin Nat. 36, p. 363; Bright, 1981, Mem. Ent. Soc. Canada 118, p. 68; Wood, 1982, Mem. Great Basin Nat. 6, p. 1133. This species was previously known only from the type series from Puebla. A series of 29 specimens were seen bearing the labels: “San Rafael, Mex., 4.1X.81, S-242, 2400 m, Atkinson - Equihua’/“Hosp.: Rhus sp.” The specimens are identical to the two paratypes in the Canadian National Collec- tion. INITIAL SURVEY OF ACETYLENE REDUCTION AND SELECTED MICROORGANISMS IN THE FECES OF 19 SPECIES OF MAMMALS C. Y. Li’, Chris Maser”, and Harlan Fay’ ABSTRACT. —Nitrogen-fixing bacteria, as demonstrated by the acetylene reduction method; yeasts, and actino- mycetes were found in feces of mammals collected from St. Lawrence Island, Alaska, to the North Carolina—Tennessee border. The mammals, representing four orders and 19 species, occupy a wide variety of habitats and may play an important role in dispersing microorganisms vital to the ecosystem. The California red-backed vole (Clethri- onomys californicus), the northern flying squirrel (Glaucomys sabrinus), and the deer mouse (Peromyscus maniculatus) are forest- dwelling rodents that may play an important role in maintaining forest productivity. These mammals consume hypogeous mycorrhizal fungi and disperse fecal pellets containing fungal spores, which germinate and form my- corrhizae with roots of forest trees (Hunt and Maser 1985, Maser et al. 1978; Food habits , 1985; Northern flying squirrel, 1985; Ure and Maser 1982). The feces of these animals also contain nitrogen-fixing bacteria and yeast. The nutrient in the feces is as effective as yeast extract in promoting bacterial growth and ni- trogenase activity (Li et al. 1986). When these animals dig at the bases of trees, the organ- isms in their feces could inoculate rootlets with nitrogen-fixing bacteria, yeast, and spores of mycorrhizal fungi. At times, actino- mycetes are also present; they produce sub- stances important in formation of soil humus (Krassilnikov 1981). Having worked out the basic links in the abili- ties of small mammals in western Oregon to pass viable nitrogen-fixing bacteria and yeast through their digestive tracts (Li et al. 1986), the next question was, How widespread is this phe- nomenon? We conducted a survey of feces of 51 mammals of 19 additional species, collected from St. Lawrence Island, Alaska, to the North Carolina-Tennessee border, for acetylene-re- ducing (nitrogen-fixing) bacteria (Postgate 1982), yeasts, and actinomycetes. MATERIALS AND METHODS Fresh fecal pellets from 51 mammals of 19 species, representing four orders, were col- lected in sterile vials (see Li and Maser 1986 for collecting techniques). Acetylene Reduction Activity One fecal pellet each from 48 small mam- mals was placed in 20 ml of Débereiner's N- free liquid medium (Débereiner and Day 1976) and Burk’s (1930) liquid medium in a 60-ml serum bottle. We used only the central portions of the pellets from three large mam- mals: black-tailed jackrabbit (Lepus californi- cus), eastern cottontail (Sylvilagus flori- danus), and elk (Cervus elaphus). Bottles were capped and flushed for 5 min with nitro- gen gas containing less than 10 ppm oxygen. The liquid medium became turbid after incu- bation for two days at 30 C. Acetylene was then injected into each bottle to 10 percent (v/v); the bottles were gently swirled immedi- ately after acetylene was added and were left to stand at 30 C. Bottles without acetylene injection served as controls. After 18 hr, 0.1 ml gaseous samples were removed from each bottle and analyzed for ethylene and acetylene with a Hewlett-Packard 5830A gas chromatograph’ fitted with a 2 m X 2.1 mm, 80-100 mesh Poropak R column. Oven tem- perature was adjusted to 70 C. Injection and flame ionization detector temperatures were each adjusted to 100 C. Nitrogen carrier gas flow rate was adjusted to 40 ml/min. 1ULS. Department of Agriculture, Forest Service, Pacific Northwest Research Station, Forestry Sciences Laboratory, 3200 Jefferson Way, Corvallis, Oregon 97331. US Department of the Interior, Bureau of Land Management, Forestry Sciences Laboratory, 3200 Jefferson Way, Corvallis, Oregon 97331. Use of trade names does not imply endorsement or approval of any product by the USDA Forest Service to the exclusion of others that may be suitable. 646 October 1986 LI ET AL.: ECOLOGY OF MAMMAL FECES 647 TABLE 1. Nitrogenase activity and microorganisms in feces of 51 mammals of 19 species from Alaska to North Carolina—Tennessee. eee Nitrogenase’ Animal Geographic location activity Yeasts” Actinomycetes” INSECTIVORA Soricidae (shrews) Sorex cinereus (1)° Morocco, IN oF not determined not determined LAGOMORPHA Leporidae (hares and rabbits) _Lepus californicus (1) Summer Lake, OR + + 4 Sylvilagus floridanus (1) Morocco, IN + 0 0 RODENTIA Sciuridae (squirrels) Cynomys leucurus (7) Meeteetse, WY 0) + 0 Cynomys leucurus (3) Laramie, WY 0 At 0 Eutamias townsendi (1) Umpqua, OR + = + Glaucomys volans (2) Morocco, IN + 0 Spermophilus parryi (1) St. Lawrence Island, AK 0 + 0 S. tridecemlineatus (2) Cedar Falls, IA + + + Tamias striatus (1) Cedar Falls, IA =f a + Geomyidae (pocket gophers) Geomys busarius (1) Morocco, IN + 0 + Cricetidae (native mice) Peromyscus leucopus (6) Morocco, IN + + 0 P. maniculatus (4) Morocco, IN + 0 0 Arvicolidae (voles) Clethrionomys gapperi(10) | Roon Mountain, NC-TN + + + Lagurus curtatus (1) Silver Lake, OR o + + Microtus ochrogaster (2) Morocco, IN + + 0 M. oeconomus (4) St. Lawrence Island, AK + + 0 M. oregoni (1) Marys Peak, OR + + + Muridae (Old World mice) Mus musculus (1) Morocco, IN oo + + ARTIODACTYLA Cervidae (deer) Cervus elaphus (1) Post, OR 0 + 0 1Four replicates (all positive or all negative). ? Average of three replications of one individual from each species. 4 g Pp p Number of individuals tested for nitrogenase activity. Bacterial cultures that reduced acetylene were all isolated but were not all purified. Yeast and Actinomycetes Sodium albumenate agar (Waksman and Fred 1922) was used to test for yeast and actinomycete populations. One fecal pellet (or central portion of a pellet from the three large mammals) per vial was removed with sterile forceps under an isolation hood. Each pellet was crushed and thoroughly mixed in 30 ml of sterile distilled water. One ml, 0.5 ml, and 0.1 ml of this fecal suspension were each plated with 20 ml of sodium albumenate agar. Colonies developed on the surface of the agar. The presence of yeasts and actinomycetes was confirmed under a light microscope, and colonies were counted after three days’ incu- bation at 30 C. Colonies were sometimes so numerous, even at high dilutions, that their numbers had to be estimated. RESULTS AND DISCUSSION Results of our study are given in Table 1. Acetylene Reduction Activity Feces of mammals of the 19 species were tested for acetylene reduction, which is a uni- versal and specific property of nitrogenase of nitrogen-fixing bacteria (Postgate 1982). Six- teen samples were positive (Table 1). Thir- teen of the 19 species are known to eat the fruiting bodies of hypogeous, mycorrhizal fungi from which they could ingest nitrogen- fixing bacteria (Li and Castellano 1985, 1986). These 13 mycophagists are: masked shrew (Sorex cinereus ) (Hamilton 1941); Lepus cali- 648 fornicus (Ponder 1980); Townsend chipmunk (Eutamias townsendi) (Maser et al. 1978); southern flying squirrel (Glaucomys volans) (Maser and Maser, unpublished data); eastern chipmunk (Tamias_ striatus) (Maser and Maser, unpublished data); white-footed mouse (Peromyscus leucopus) (Fogel and Trappe 1978, Maser et al. 1978, Whitaker 1966); P. maniculatus (Hunt and Maser 1985, Maser et al. 1978); southern red-backed vole (Clethrionomys gapperi) (Fogel and Trappe 1978, Maser et al. 1978, Ure and Maser 1982); sage vole (Lagurus curtatus ) (Dowding 1955, Maser et al. 1978); prairie vole (Microtus ochrogastar ) (Fogel and Trappe 1978); tundra vole (M. eoconomus) (Fogel and Trappe 1978); creeping vole (M. oregoni) (Maser et al. 1978); and house mouse (Mus musculus) (Whitaker 1966). Yeast Mammals that feed on fungi can also ingest yeasts (Anderson and Skinner 1947, Kockova- Kratochvilova et al. 1984). Yeasts were virtu- ally ubiquitous in our samples and passed through the digestive tracts of 15 of the 18 species checked (Table 1). Yeast propagules ranged from zero to an estimated 1,800,000 per pellet. Actinomycetes Actinomycetes, often called “ray fungi,” are actually higher bacteria. They may occur both in soil and on the surfaces of plant leaves (Dickinson et al. 1975, Lechevalier 1981). Of the 18 species examined for actinomycetes, the 9 that were positive (Table 1) are known to eat substantial amounts of green vegetation (Bailey 1936, Bee and Hall 1956, Hamilton and Whitaker 1979, Hansen and Flinders 1969, Lechleitner 1969, Whitaker 1966). Acti- nomycetes ranged from zero to an estimated 600,000 per fecal pellet. Potential Interrelations Mammals generally are abundant and mo- bile and form functional links with all areas of the terrestrial habitat, from below the ground into the tree tops. They deposit fecal pellets throughout their habitats. Fecal pellets of some species contain viable nitrogen-fixing bacteria, yeast propagules, and actinomycetes (Table 1). Yeast and actinomycetes apparently can be obtained by mammals on plant mate- GREAT BASIN NATURALIST Vol. 46, No. 4 rial (Dickinson et al. 1975, Lechevalier 1981); however, nitrogen-fixing bacteria in pellets seem to be associated more with soil and food obtained below the ground. For example, fe- ces of white-tailed prairie dog (Cynomys leu- curus), arctic ground squirrel (Spermophilus parryi), and Cervus elaphus showed no acetylene reduction activity (Table 1); their sole diet in early summer when the pellets were collected might have been only the aboveground portions of green vegetation. At other times of the year, their droppings may include nitrogen-fixing bacteria because of shifts in food habits. Spermophilus parryi, for instance, may eat mushrooms at certain times (Bee and Hall 1956) and thus may ingest nitro- gen-fixing bacteria. Cervus elaphus eat hypo- geous fungi part of the year (Trappe et al., unpublished data) and may ingest nitrogen- fixing bacteria. Other species, such as the plains pocket gopher (Geomys bursarius), feed on subterranean portions of plants and also ingest some soil while digging (Hamilton and Whitaker 1979); this behavior could also account for nitrogen-fixing bacteria in the fe- ces. Thus far, we have been able to identify only three of the isolated nitrogen-fixing bacteria: Azospirillum sp., Clostridium butyricum, and C. beijerinckii. Azospirillum sp. has been isolated from feces of the California red- backed vole and the northern flying squirrel (Li et al. 1986), and the creeping vole (Micro- tus oregoni) (Li and Maser, unpublished data). Clostridium butyricum has been iso- lated from feces of the deer mouse (Li et al. 1986), and C. beijerinckii from feces of the thirteen-lined ground squirrel (Spermophilus tridecemlineatus) (this study). These three species of nitrogen-fixing bacteria occur freely in the soil (Buchanan and Gibbons 1974, Hammann and Ottow 1976, Jones and Bangs 1985, Lakshmi et al. 1977). Azospirillum sp. can penetrate plant roots (Lakshmi et al. 1977, Patriquin and Débereiner 1978), and Azospirillum sp. and Clostridium sp. have also been found associated with ectomycor- rhizae of Douglas-fir (Pseudotsuga menziesii) (Li and Hung, Plant and Soil, in press). Some species of yeast may increase nitro- gen fixation in the presence of mycorrhizal fungi and thereby improve site productivity (Li et al. 1986, Maser et al. 1984). Yeast in the feces of mycophagous mammals may also be October 1986 important because spore germination of some mycorrhizal-forming fungi is stimulated by extractives from other fungi, such as yeast (Fries 1966, 1982, Oort 1974). Actinomycetes produce substances that are important in the formation of soil humus (Krassilnikov 1981), and humus, in turn, is important to the formation of mycorrhizae (Harvey et al. 1976, Kumuda et al. 1961, Maser et al. 1984). As stated by Linderman (1985), however, microbial interactions are complex, and actinomycetes are but a fraction of the complexity. Mammals may play an important functional role in dispersing microorganisms vital to the ecosystem. Their potential importance is fur- ther suggested when these new data are cou- pled with the role of mammals in the dispersal of viable spores of mycorrhizal fungi, which are obligatory symbionts of most plants (Fogel and Trappe 1978, Kotter and Farentinos 1984, Maser et al. 1978, Rothwell and Holt 1978, Trappe and Maser 1977). CONCLUSION We reiterate that our survey was intended to ascertain the potential geographical scope of nitrogen-fixing bacteria (through acetylene reduction) in mammals. Our survey has some obvious deficiencies: for example, mammals were collected at different seasons and in dif- ferent habitats, so standardizing was impossi- ble. Isolating and identifying the nitrogen- fixing bacteria was extremely difficult; to our knowledge, no one has done this type of study before and laboratory techniques had to be mod- ified (Li and Maser 1986). Finally, other mi- croorganisms, such as yeasts and actinomycetes, are both poorly known and understood. To study and understand the vast array of ecosystem processes require a carefully planned, interdisciplinary approach. As we learn more about mammal-habitat interac- tions, research must be aimed at the mammals as complex, functional links in the ecosystem. Understanding these dynamic linkages will help us to manage wisely both the mammals and their habitats to maintain or improve the health of the ecosystem. ACKNOWLEDGMENTS We thank G. E. Menkens, Jr. (Wyoming), R. L. Rausch (St. Lawrence Island, Alaska), J. Li ET AL.: ECOLOGY OF MAMMAL FECES 649 O. Whitaker, Jr. (Indiana), P. D. Weigl (North Carolina—Tennessee border), and N. Wilson (Iowa), who sent us specimens; P. J. Bottomley (Department of Microbiology, Oregon State University, Corvallis), R. L. Rausch (Division of Animal Medicine, Uni- versity of Washington, Seattle), R. F. Tarrant (Department of Forest Science, Oregon State University, Corvallis), and W. Thies (USDA Forest Service, Pacific Northwest Research Station), who critically reviewed the paper; and G. Bissell, who typed the various drafts. 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Linn Benton Community College. Albany, Oregon. Ure, D. C., AND C. Maser. 1982. Mycophagy of red-backed voles in Oregon and Washington. Canadian J. Zool. 60: 3307-3315. : WAKSMAN, S. A., AND E. B. FRED. 1922. A tentative outline of the plate method for determining the number of mi- croorganisms in the soil. Soil Sci. 14: 27-28. WHITAKER, J. O., JR. 1966. Food of Mus musculus, Peromyscus maniculatus bairdi and Peromyscus leucopus in Vigro County, Indiana. J. Mammal. 47: 473-486. i NOTES ON THE BIRDS OF COLD SPRING MOUNTAIN, NORTHWESTERN COLORADO Peter O. Dunn’” and Ronald A. Ryder! ABSTRACT.—Observations are presented on 117 bird species seen in a 250-km” area of northwestern Colorado adjacent to Utah and Wyoming. Three previously unreported species and seven status changes are listed for the Rangely, Colorado, latilong block. The bird life of northwestern Colorado and adjacent Utah and Wyoming is among the least known in the lower 48 states. Previous descriptions of this area resulted from short- term visits between the early 1900s and the mid-1960s (Cary 1909, Hendee 1929, Behle and Ghiselin 1958, Hayward 1967). The need for more ornithological fieldwork in this area became apparent with the advent of bird map- ping schemes in Colorado and Utah based on latilong blocks (1 degree latitude and 1 degree longitude in size) (Chase et al. 1982, Walters and Sorensen 1983, respectively). These dis- tribution plotting systems are valuable for providing an environmental data base for land-use planning and management, and, with some additional effort, for testing hy- potheses about migration routes and distribu- tions (see Bock 1984 for a similar example using 5-degree latilong blocks). However, for latilong information to be useful, all blocks should have adequate and similar data bases. Unfortunately, these maps often show not the distribution of birds but the distribution of bird-watchers. For exam- ple, vesper sparrow (Pooecetes gramineus ) and white-crowned sparrow (Zonotrichia leu- cophrys) were two of the most common breeding species on our study area (Table 1), yet they had not been recorded previously as breeding in the Rangely latilong block be- cause of the paucity of bird-watchers. To sup- plement the inadequate ornithological records for northwestern Colorado and to in- _ crease the usefulness of the Colorado latilong data base, this report presents bird observa- tions during a study of sage grouse (Centro- cercus urophasianus ) conducted in the north- western corner of Colorado (northwest corner of the Rangely latilong block). The sage grouse study was centered on Cold Spring Mountain approximately 1 km east of Utah, 7 km south of Wyoming, and immediately north of Browns Park National Wildlife Refuge, Moffat County, Colorado. Cold Spring Mountain (2,622 m) is part of the eastern extension of the Uinta Mountains, the largest east-west range in the Western Hemi- sphere. The topography of the study area varies from mountainous to rolling hills and mesas_ (1,820-2,909 m). Big sagebrush (Artemisia tridentata)—dominated rangeland and pinyon pine (Pinus edulis )—Utah juniper (Juniperus osteosperma) cover most of the study area. Quaking aspen (Populus tremu- loides) woodland occurs on Cold Spring Mountain and in most canyons and mountain- sides. Lodgepole pine (Pinus contorta) and Douglas-fir (Pseudotsuga menziesii) occur above 2,620 m on Middle Mountain (2,904 m) and Diamond Peak (2,909 m) near the Wyo- ming border. Snowfall at lower elevations oc- curs from November to April, whereas moun- tains in the study area often remain snow covered from late October to mid-May. This late snowmelt delays spring arrivals and nest- ing of many passerines on Cold Spring Moun- tain. The annotated list (Table 1) is only of birds seen in the vicinity of Cold Spring Mountain, Diamond Peak, Middle Mountain, and Sugar- loaf Flats; this is an area of approximately 250 km? covering the northwest corner of Colo- rado. The list of waterfowl and shorebirds is 1Department of Fishery and Wildlife Biology, Colorado State University, Fort Collins, Colorado 80523. 2Present address: Department of Zoology, University of Alberta, Edmonton, Alberta, Canada T6G 2E9. 651 652 GREAT BASIN NATURALIST Vol. 46, No. 4 TABLE 1. List of bird species seen on Cold Spring Mountain and vicinity, northwestern Colorado. Species Horned Grebe (Podiceps auritus) Eared Grebe (Podiceps nigricollis) Green-winged Teal (Anas crecca) Mallard (Anas platrhynchos) Northern Pintail (Anas acuta) Blue-winged Teal (Anas discors) Northern Shoveler (Anas clypeata) American Wigeon (Anas americana) Turkey Vulture (Cathartes aura) Bald Eagle (Haliaeetus leucocephalus) Northern Harrier (Circus cyaneus) Sharp-shinned Hawk (Accipiter striatus ) Coopers Hawk (Accipiter cooperii) Northern Goshawk (Accipiter gentilis) Swainson’s Hawk (Buteo swainsoni) Red-tailed Hawk (Buteo jamaicensis ) Ferruginous Hawk (Buteo regalis) Rough-legged Hawk (Buteo lagopus) Golden Eagle (Aquila chrysaetos) American Kestrel (Falco sparverius ) Merlin (Falco columbarius) Prairie Falcon (Falco mexicanus ) Chukar (Alectoris chukar) Blue Grouse (Dendragapus obscurus ) Sage Grouse American Coot (Fulica americana) Sandhill Crane (Grus canadensis) Killdeer (Charadrius vociferus ) American Avocet (Recurvirostra americana) Greater Yellowlegs (Tringa melanoleuca) Lesser Yellowlegs (Tringa flavipes) Solitary Sandpiper (Tringa solitaria) Spotted Sandpiper (Actitis macularia) Western Sandpiper (Calidris mauri) Long-billed Dowitcher (Limnodromus scolopaceus) Common Snipe (Gallinago gallinago) Wilson’s Phalarope Franklin’s Gull (Larus pipixcan) Mourning Dove (Zenaida macroura) Great Horned Owl (Bubo virginianus ) Northern Pygmy-Owl Long-eared Owl (Asio otus) Common Nighthawk (Chordeiles minor) Common Poorwill White-throated Swift Broad-tailed Hummingbird (Selasphorus platycercus) Red-naped Sapsucker (Sphyrapicus nuchalis ) Downy Woodpecker Hairy Woodpecker (Picoides villosus) Status Latilong study! (Changes, Abun- this study) dance? M U B R FC R C R FC B FC R U R FC B FC W R R FC R U B U R(B) U B FC R FC B U W C R FC R FC M B FC R U R FC R C B M R R C B M M M B FC M FC M U B FC b FC M R C R C (W) R U B FC b(B, FC Hendee 1929) b U B C M R b FC M FC Dates seen? Earliest Latest 10 Aug? 20 May® 30 Apr 6 Oct 31 May 10 Aug 30 Apr 10 Aug 20 May 8 Jul 21 May 10 Aug 8 Jun 25 Aug 24 May 27 Sep 5 Apr 7 Nov Seen every month 4, 26 Aug? 30 Apr 13 Sep 30 Apr 19 Sep 16 Aug 27 Sep 5 Apr 19 Sep 5, 6 Apr? 1 Nov 7 May Seen every month 16 Apr 3 Oct 20 Aug? 4 Apr 27 Jul 20,24 Jul; 8 Sep® 20 Apr 19 Sep Seen every month 30 May” 4,5 Apr? 16 Apr 15 Aug 20 May? 23 Jul 4 Aug? 22 Jul 31 May 17 Aug 13, 15 Aug? 3 Aug? 26 Apr 23 Jul 30 Apr 28 Jul 7 May” 7 May 6 Oct Nesting dates' Comments Young:8 Jul Young:8 Jul Young:8, 15 Jul Nest:30 Apr Nest:30 Apr Nest:10 Apr 5 birds feeding on Mormon Crickets on 5 and 6 Aug Young:19 Jul Nests:9(Z nests), 19, 20(2), 21, 31(2) May; second nests: 21, 31, May; 5, 17 June; young:many 3- week and older chicks after 12 Jul Many juvenile birds: 26-31 Jul Seen or heard every month Seen in Irish Canyon on 4 Feb 1984 18 Jun 21 Sep 31 May 25 Aug 14 Jul 19 Sep 12 May 26 Sep 3 Jun 26 Aug 18 Apr 21 May 12 Apr 19 Sep 18 Jul 17 Aug Hendee (1929) found a nest 28 May near the Lit- tle Snake River Most common second week in Aug; 20—40 birds/ night October 1986 Table 1 continued. DuNN, RYDER: COLORADO BIRDS 653 Status Latilong study! (Chances hans Dates seen® Species this study) | dance* _— Earliest Latest = Nesting dates* Comments Three-toed Woodpecker (Picoides tridactylus ) (n or b) Seen on 24 July 1981 at Swede Spring on Cold Spring Mountain Northern Flicker (Colaptes auratus ) R C Seen every runth : Western Wood Pewee b FC 8 Jun ~10 Aug Dusky Flycatcher (Empidonax oberholseri) B FC 7 Jun 18 Aug Gray Flycatcher (Empidonax wrightii) B 3 Jun? Western Kingbird (Tyrannus verticalis ) b R 8 May 15 Aug Horned Lark (Eremophila alpestris ) R Cc Seen every month Nest: 5, 7 Jun Tree Swallow (Tachycineta bicolor) B G 11 Apr 5 Aug Violet-green Swallow (Tachycineta thalassina) b(B) C 21 May 3 Aug Adult carrying fecal sac:23 Jun Barn Swallow (Hirundo rustica) B FC 15 Apr 5 Aug Steller’s Jay (Cyanocitta stelleri) R FC 19 Jul, 13 Sep” Scrub Jay (Aphelocoma coerulescens ) R FC Seen every month Pinyon Jay (Gymnorhinus cyanocephalus ) R Cc Seen every month More than 50 birds feeding on Mormon Crickets 5 Aug Clark’s Nutcracker (Nucifraga columbiana) R Gc Seen every month Ten birds feeding on Mormon Crickets 5 Aug Black-billed Magpie (Pica pica) R C Seen every month More than 100 birds feeding on Mormon Crickets 5 and 17 Aug American Crow (Corvus brachyrhynchos ) R FC Seen every month Common Raven (Corvus corax) R FC Seen every month Black-capped Chickadee W(N) FC 20 Feb, 14 Apr, 3 May, 15, 17 Aug; 19 Sep° Mountain Chickadee (Parus gambeli) R FC Seen every month Plain Titmouse (Parus inornatus ) B FC 3, 4, 8 May” Bushtit b 5 May? Red-breasted Nuthatch (Sitta canadensis ) R U 29 Jun 26 Aug White-breasted Nuthatch (Sitta carolinensis ) R 21 May” Rock Wren (Salpinctes obsoletus ) B U Seen every month House Wren (Troglodytes aedon) B G 4May 24 Aug Fledglings:27 Jun Ruby-crowned Kinglet (Regulus calendula) M U 17, 21 May? Blue-gray Gnatcatcher (Polioptila caerulea) B 15 Jun? Mountain Bluebird (Sialia currucoides ) B FC 17 Mar 3 Oct Townsend’s Solitaire (Myadestes townsendi) M FC 4 May 3 Oct American Robin (Turdus migratorius ) R FC Seen every month 2 nests with 4 at lower elevations eggs:5 Jun Northern Mockingbird (Mimus polyglottos ) b UC 6 Jun 3 Oct Sage Thrasher (Oreoscoptes montanus ) B FC 6 Apr 25 Aug Water Pipit (Anthus spinoletta) (b or M) 3 Oct 1981° 4 birds near Arthur's Reservoir Bohemian Waxwing (Bombycilla garrulus ) Ww 3 Nov? : Northern Shrike (Lanius excubitor) WwW U 1,7 Nov? | Loggerhead Shrike (Lanius ludovicianus ) B U 31 May, 4 Jun? European Starling (Sturnus vulgaris ) R E Seen every month Warbling Vireo (Vireo gilvus ) b(B) FC 31 May 19Sep 2nests:19 Jun Yellow Warbler (Dendroica petechia) B FC 15 Jun 17 Aug Yellow-rumped Warbler (Dendroica coronata) B U 21 May 17 Aug Sa feeding young: u Black-throated Gray Warbler 2 (Dendroica nigrescens ) B U 17, 21 May? MacGillivray’s Warbler (Oporornis tolmiei) B FC 9 Jun 19 Sep Common Yellowthroat (Geothlypis trichas ) B 15 Jun? Wilson’s Warbler (Wilsonia pusilla) B FC 17, 18, 25 Aug Western Tanager b FC 10 Aug? Black-headed Grosbeak ; (Pheucticus melanocephalus ) B FC 1 Jun 17 Aug Female with food: 27 Jul 654 GREAT BASIN NATURALIST Vol. 46, No. 4 Table 1 continued. Status Latilong study! (Changes, Abun- Dates seen® Species this study) dance? Earliest Latest Nesting dates* Comments Lazuli Bunting (Passerina amoena) b 1] Aug? Green-tailed Towhee (Pipilo chlorurus ) B FC 4 May 13 Sep Nest with 4 eggs: Rufous-sided Towhee (Pipilo erythrophthalmus ) B American Tree Sparrow (Spizella arborea) Ww Chipping Sparrow (Spizella passerina) B Brewer s Sparrow (Spizella breweri) B Vesper Sparrow b(B) Lark Sparrow (Chondestes grammacus ) B Sage Sparrow (Amphispiza belli) B Savannah Sparrow (Passerculus sandwichensis ) b White-throated Sparrow (Zonotrichia albicollis ) M White-crowned Sparrow M Dark-eyed Junco (Junco hyemalis ) R Red-winged Blackbird (Agelaius phoeniceus ) Western Meadowlark (Sturnella neglecta) Yellow-headed Blackbird (Xanthocephalus xanthocephalus ) B Brewer's Blackbird (Euphagus cyanocephalus) B Brown-headed Cowbird (Molothrus ater) Rosy Finch (Leucosticte arctoa) Cassin’s Finch (Carpodacus cassinii ) Red Crossbill (Loxia curvirostra) Pine Siskin (Carduelis pinus ) American Goldfinch (Carduelis tristis ) DeTnmet 5 Jun; young:20 Jun U 1 May 7 Jun FC 3 Nov 4 Apr FC 16 Jun 25 Aug Cc 31 May 19 Sep Nest:8, 21 Jun; One of the most young:23 Jun, common breeding 24 Jul species € 25 Apr 26 Sep Eggs:21 Jun; One of the most fledglings:27 common breeding Jun, 23-31 Jul species FC 8 May 4 Jun R 3, 6, 30 Apr? 17 Aug? 31 May? € 10 May 26 Aug Nests:1 Jun Very common eggs:1,5 Jun breeding species young:15 Aug FC Seen every month Nest:16 Jun young:28 Jun C 20 Apr 19 Sep IKE; 13 Apr 3 Oct FC 2 May 19 Jul FC 1 May 19 Sep Nest:19 Jun More than 100 birds feeding on Mormon Crickets on 6, 18 Aug FC 2 May 4 Sep U 1 Nov 12 Apr FC 16, 18 May? 31 May? FC 1 Nov 30 Apr 30 May? 1Status given in Chase et al. (1982). R = Resident year-round (breeds): N = Nonbreeder present year-round, or a year-round resident whose breeding has not been documented; B = Breeding (documented); b = Likely breeder; W = Winter visitor; M = Migrant; blanks indicate no record. Abundance categories: C = Common; FC = Fairly common; U = Uncommon; R = Rare; Irr = Irregular. Abundance was estimated in this study, and categories are from Chase et al. (1982). A blank indicate too few data to evaluate. Dates observed represent our records and may not be indicative of actual arrival/departure dates for migrants and summer residents in the entire region. ‘Nesting dates are presented only where information is known. °Only date(s) seen. relatively short because most of these species stop at better habitat in Browns Park; Cold Spring Mountain is mostly xeric, with the ex- ception of six small reservoirs. A notable as- pect of the species list is that the breeding status of several relatively common species remains to be documented. Future field workers in this area should attempt to docu- ment breeding for Wilson's phalarope (Phalaropus tricolor), northern pygmy owl (Glaucidium gnoma), common poorwill (Pha- laenoptilus nuttallii), white-throated swift (Aeronautes saxatalis), downy woodpecker (Picoides pubescens), western wood pewee (Contopus sordidulus), western kingbird (Tyrannus verticalis), black-capped_chick- adee (Parus atricapillus), bushtit (Psaltri- parus minimus), and western tanager (Pi- ranga ludoviciana), among others. The northern pygmy owl, three-toed woodpecker, and water pipit were recorded for the first time from this area. ACKNOWLEDGMENTS Financial support for this project was pro- vided by the Colorado Division of Wildlife through Federal Aid to Wildlife Restoration Project W-37-R and the Rob and Bessie Welder Wildlife Foundation, Sinton, Texas. October 1986 The support of these agencies and organiza- tions is gratefully acknowledged. A draft of the manuscript was reviewed by C. M. Haynes and W. D. Snyder. C. E. Braun kindly re- viewed drafts of the manuscripts and provided access to his field notes. J. W. Hupp, T. E. Olsen, and D. Ward also provided records of their observations. This is Welder Wildlife Foundation publication number 303. LITERATURE CITED BEHLE, W. H., AND J. GHISELIN. 1958. Additional data on the birds of the Uinta Mountains and basin of northeastern Utah. Great Basin Nat. 18: 1-22. DUNN, RYDER: COLORADO BIRDS 655 Bock, C. E. 1984. Geographical correlates of abundance vs. rarity in some North American winter land- birds. Auk 101: 266-273. Cary, M. 1909. New records and important range exten- sions of Colorado birds. Auk 26: 180-185. Cuase, C. A., S. J. BissELL, H. E. KINGERY, AND W. D. GrauL. 1982. Colorado bird distribution latilong study. Colorado Field Ornithologists, Denver Mus. Nat. Hist., Denver, Colorado. 78 pp. HaywarbD, C. 1967. Birds of the upper Colorado River basin. Brigham Young University Sci. Bull., Biol.. Ser. 9(2): 1-64. HENDEE, R. 1929. Notes on birds observed in Moffat County, Colorado. Condor 31: 24-32. WALTERS, R. E., AND E. SORENSEN. 1983. Utah bird distri- bution: latilong study 1983. Utah Dept. Nat. Re- sour., Div. Wildl. Resour., Nongame Sec., Salt Lake City. 97 pp. LIFE STRATEGIES IN THE EVOLUTION OF THE COLORADO SQUAWFISH (PTYCHOCHEILUS LUCIUS) Harold M. Tyus’ ABSTRACT.—The Colorado squawfish, a large predaceous cyprinid, is a generalist species adapted to the large seasonal water fluctuations, low food base, and changing riverine subsystems of the Colorado River. Extant at least as early as the Miocene epoch, Ptychocheilus has survived by incorporating life strategies to deal with changing climates varying from arid to pluvial. Migration and long-term movement patterns appear to have evolved as tactics to perpetuate a grand reproductive strategy for exploiting the changing habitats and general environmental conditions of the late Cenozoic era. Accordingly, high mobility of a large fish would aid in selection of optimum spawning, nursery, and adult habitats in the dynamic lacustrine/riverine system that existed at that time. A spatial separation of life stages thus produced would aid in the reduction of intraspecific competition. Large size, long life, and late spawning of Ptychocheilus indicate that mortality of young must be disproportionately high compared to that of the adult form. Growth to a large size should reduce predation by other fishes and, once attained, would facilitate long distance movement for reproduction, feeding, and other purposes. Such a strategy, formerly highly adaptive, may now be implicated in the decline of this species in controlled riverine systems. The genus Ptychocheilus includes the largest cyprinids in North America. Repre- sented by four species today, the largest of these, the Colorado squawfish (Ptychocheilus lucius Girard) formerly grew to a size of about 1.8 mand 45 kg (Miller 1961). Endemic to the Colorado River Basin, this fish, once dis- tributed throughout the basin, has declined since the 1930s and is today restricted to the upper Colorado River Basin, where it is classi- fied as endangered by the U.S. Fish and Wildlife Service (1973, 1974). The loss of the Colorado squawfish from parts of the Colo- rado River is apparently related to major wa- ter developments that have ostensibly re- duced P. lucius to about 25% of its former range (Tyus 1984). Although many workers have postulated man-induced changes in riv- erine conditions as primary factors in the re- duction of the range and abundance of this species (Miller 1961, Holden and Wick 1982, Ono et al. 1983), a lack of basic knowledge about its life history, especially in locations where the fish has been lost (Minckley, 1973), has made these implications impossible to prove. Recent research in the Green River Basin (Fig. 1) by the U.S. Fish and Wildlife Service (Tyus and McAda 1984) resulted in the first discovery of a spawning grounds of this species in 1981 and identified migrations and movements as important factors in the 1U.S. Fish and Wildlife Service, 1680 West Highway 40, Vernal, Utah 84078. reproductive strategy of this species. These findings have been substantiated by the work of Haynes et al. (1984), Wick et al. (1983), Tyus (1985), and others. With the present knowledge of the life his- tory requirements of P. lucius, it is now possi- ble to relate its apparent life strategies with its evolution and adaptations to conditions in the Colorado River Basin. In so doing I have drawn heavily from the works of G. R. Smith (1981) and M. L. Smith (1981), who presented the background on late Cenozoic climates and adaptations of the southwestern fish fauna, particularly P. lucius , upon which this work is based. CLIMATE AND ADAPTATION OF PTYCHOCHEILUS The cyprinid fishes apparently arrived in the New World from Asia in the Miocene epoch, and fossil Ptychocheilus species similar to modern Ptychocheilus lucius have been re- ported from the middle Pliocene in the Colo- rado River system of northern Arizona (Miller 1961). Ptychocheilus had widespread distri- bution in the Pliocene, as evidenced by fossils in Lake Idaho (Smith 1975), the Great Basin (G. Smith 1981), and Arizona (Miller 1961). Furthermore, the similarity between the Pliocene fossils and modern forms suggests that the adaptation to swift water habitat had 656 October 1986 Tyus: COLORADO SQUAWFISH Stillwater COLORADO River GRAND JUNCTION e, Mn en Zi, \00 a Fig. 1. Upper Colorado River Basin and Green River study area (shaded). occurred in Ptychocheilus by the mid- Pliocene (Miller 1961). Nonetheless, the largest Ptychocheilus reported in the fossil record lived in Pliocene Lake Idaho (Smith 1975), indicating that Ptychocheilus success- fully utilized both riverine and lacustrine sys- tems. The Southwestern United States is more arid today than in the Late Cenozoic, and this increasing aridity no doubt resulted in the loss or reduction of lacustrine habitats and the extinction of lake dwelling salmonids and cen- trarchids from the Colorado River system. This change was progressive from the Pliocene, when a system of lakes covered the lower and upper Colorado River Basins, and persisted during pluvial periods until the Pleistocene. During this epoch the life history of fishes was remarkably impacted by such long pluvial periods interrupted by short peri- ods of desert conditions (G. Smith 1981). An evaluation of the fish fauna of the Colo- rado River in recent times (before introduc- tions by man) might lead one to conclude that 658 the isolated drainages and depauperate faunas of today reflect Cenozoic conditions. They do not. Instead, the fossil record shows that the large regional desert environs of the South- west are “geologically new” (M. Smith 1981) and not typical in the development of life history attributes of the fish fauna. This has led M. Smith (1981) to propose that the eco- logical history of the fishes suggests they should be considered generalists, not special- ist species. In this case Ptychocheilus would have developed the capability to utilize both riverine and lacustrine habitat depending upon the climatic conditions prevailing. During the late Cenozoic, estuarine condi- tions in the lower basin and widespread lacus- trine habitat during pluvial periods would provide eutrophic conditions that could be exploited by a top carnivore like Ptycho- cheilus. These same areas, however, might not have provided the best spawning and nursery conditions because of adverse envi- ronmental (e.g., oxygen, substrate) and bio- logical (e.g., predation) factors. If Ptycho- cheilus could move between preferred spawning and feeding areas, it might have the best of both. G. Smith (1981) proposed that migration would be a major adaptation to dry seasons for intermountain desert fishes like Ptychocheilus, and that emigration of young fish to unoccupied areas might be selected for in genotypes. If movement and/or migration is highly adaptive, this behavior would have evolved with modern Ptychocheilus. Another consideration in the evolution of Ptychocheilus is large adult size. The popular notion of a richer food supply in the recent past is interesting, but is probably not the factor driving the adaptation to large body size. In the intermountain desert G. Smith (1981) noted the tendency for large habitats to produce large fishes, and, in view of the low food ration available, suggested that life his- tory adaptations to the growing season and differential mortality are primary determi- nants. Species experiencing low adult mortal- ity that grow larger and live longer could be expected to produce a iarge number of off- spring in the desirable wet years and outcom- pete those species that sacrifice size and longevity for early reproduction. Since Ptychocheilus and various salmonids are the only large native predators throughout most of the Colorado River, survival to a moderate GREAT BASIN NATURALIST Vol. 46, No. 4 KILOMETERS MAY JUNE JULY AUG SEPT MONTH Fig. 2. Movement of radiotelemetered Colorado squawfish, Yampa and Green rivers, 1983 and 1984 (after Archer et al. 1985). Mouth of Yampa River = 0 km. size would insure low adult mortality. Thus, modern Ptychocheilus should display rapid growth and delayed reproduction to favor a large adult size if these attributes have selec- tive advantage. STRATEGIES OF PTYCHOCHEILUS LUCIUS As stated previously, modern Ptycho- cheilus exists today in conditions different from those in which it evolved. An examina- tion of the known life history attributes of P. lucius contrasted with the conditions and po- tential adaptations to late Cenozoic conditions may reveal life strategies in its evolution that would aid in its survival and potential recoy- ery. Migration, Movement, and Habitat Selection As predicted by G. Smith (1981), P. lucius makes extensive use of migration in its life October 1986 400 Tyus: COLORADO SQUAWFISH 659 GREEN WHITE YAMPA 300 a co 00 Se = Zz 100 Y J A Y J A Yeah Fig. 3. Catch of Colorado squawfish from the Green, White, and Yampa rivers (Tyus et al. 1982, Miller et al., White River, 1982; Yampa River, 1982. Y=young of year, J=juveniles, A=adult. strategy, and adults have been documented as homing to desirable spawning sites (Tyus 1985). Figure 2 illustrates the spectacular spawning migrations to the Yampa River spawning site in 1983 and 1984. Migrations of young are not so easily documented, but downstream transport of larvae have been noted by Haynes et al. (1984) and Tyus and McAda (1984). A net long-term movement of juveniles must occur to populate adult areas upstream, probably in the late young-adult stage, is indicated by collection data (Tyus et al. 1982). Figure 3 illustrates that, in the mainstem Green River, young P. lucius are relatively abundant and juveniles common; however, in the major tributaries (White and Yampa rivers) where adults predominate, ju- veniles are rare and young absent during most of the year. Habitat selection appears to be the driving force for migration. Hence, adults move up to 200 km to spawn in white-water canyons. Af- ter hatching, young larvae can drift down- stream and occupy warm shallow habitats where rapid growth is possible. These move- ments also aid in reducing intraspecific preda- tion since the adults and young tend to con- centrate in different river sections. Recent studies (Archer et al. 1985) also show that during flood periods adult P. lucius move out of the river banks and occupy flooded bot- toms, where they presumably feed on terres- trial wildlife such as small mammals (Beckman 1952). Potamodromous migrations of cyprinid fishes are not well documented for North American forms, at least not for migrations of 100 km or more. Such migrations are not un- common in flood plain rivers in other parts of the world (Welcomme 1979). Ptychocheilus lucius appears to take advantage of river trans- port at the end of the flood period for the dispersal of young from the spawning grounds downstream into productive nursery habitat (Tyus and McAda 1984). This behavior resem- bles some South American freshwater species in this regard, and it has been noted that in Africa potamodromy may protect the young from predation and secure dispersal over the river basin (Welcomme 1979). Reproductive Adaptations _ The spawning of P. lucius occurs in middle to late summer under a decreasing flow regi- men. This is unusual among most stream fishes, which spawn in the spring and early summer with rising water levels. As with other potamodromous riverine species, tim- ing of reproduction is very important, and studies of spawning P. lucius (Archer et al. 1984) indicate the fish apparently times its spawning to coincide with the descending limb of the hydrograph, a time when down- stream transport of young would distribute 660 them into the shallow nursery habitat that forms during this period in the Green River. Such a temporal adaptation fits in well with the life strategy of P. lucius, for the length of exposure of P. lucius young to predators is reduced. This reduced time for the young to feed is balanced by delivering them into ideal conditions for growth. This species selects highly oxygenated white-water rapids and riffles for spawning sites that may be 100 km or more from their preferred adult habitat at that time (Archer and Tyus 1984). Although the mechanism by which these fish congregate in spawning areas is unknown, a homing response (Tyus 1985) could result in sufficient breeding adults re- turning to a small area to insure good genetic recombination and, therefore, maintain a high degree of genetic diversity in the population. Natural Adaptations and Controlled Systems Ptychocheilus lucius evolved as a species adapted to conditions existing at the close of the Cenozoic era. These same adaptations en- abled it to compete and survive in the isolated and depauparate Colorado River Basin in the Holocene until the coming of man. Although cause-effect relationships between water de- velopment and the decline of the Colorado squawtish have not been proven de facto, it is generally agreed that such development neg- atively affects the fish (Ono et al. 1983). The life strategies developed from comparing life history attributes of P. lucius with late Ceno- zoic climatic, geologic, and fossil records sug- gest that evolving life strategies that adapted P. lucius to the natural system would ill befit the fish to a controlled system. Paramount in the life strategy of P. lucius is the need for unimpeded movement within the riverine system, and blockage of major stream sections where P. lucius occurs has resulted in the extirpation of the fish from these areas (Tyus 1984). In addition, the downstream transport of larva and establish- ment of shallow euphemeral embayments for nursery areas are needed, and a proper dis- charge regime must be maintained for spawn- ing and rearing of young. Life strategies proposed herein for P. lucius need refinement and further substantiation. Only by understanding these strategies, how- ever, can we place its evolution in proper context and provide for its future. GREAT BASIN NATURALIST Vol. 46, No. 4 ACKNOWLEDGMENTS The research upon which concepts in this paper are developed was supported, in part, by the Fish and Wildlife Service, Bureau of Reclamation, National Park Service, Bureau of Land Management and the States of Colo- rado and Utah. Principal field personnel in- cluded C. W. McAda, B. D. Burdick, K. C. Harper, R. M. McNatt, J. J. Krakker, Jr., W. B. Harned, E. J. Wick, and D. L. Skates. Administrative direction was furnished by W. H. Miller and D. L. Archer. My thanks are given to W. R. Hansen and R. L. Jones, who provided suggestions for the manuscript. LITERATURE CITED ARCHER, D. L., AND H. M. Tyus. 1984. Colorado squawfish spawning study, Yampa River. U.S. Fish Wildl. Serv., Colorado River Fish. Proj., Salt Lake City, Utah. 34 pp. : ARCHER, D. L., H. M. Tyus, AND L. R. KAEDING. 1985. Colorado River fish monitoring project. Final re- port. U.S. Fish Wildl. Serv., Colorado River Fish. Proj., Salt Lake City, Utah. BECKMAN, W. C. 1952. Guide to the fishes of Colorado. University of Colorado Museum, Boulder, Colo- rado. 110 pp. Haynes, C. M., T. A. LYTLE, E. J. WICK, AND R. T. MUTH. 1984. Larval Colorado squawfish (Ptychocheilus lucius Girard) in the upper Colorado River Basin, Colorado. 1979-1981. Southwest. Nat. 29: 21-34. HOLDEN, P. B., AND E. J. Wick. 1982. Life history and prospects for recovery of Colorado squawfish. Pages 98-108 in W. H. Miller, H. M. Tyus, and C. A. Carlson, eds., Fishes of the upper Colorado River system: present and future. West. Div., Amer. Fish. Soc., Bethesda, Maryland. 131 pp. MILLER, R. R. 1961. Man and the changing fish fauna of the American Southwest. Pap. Mich. Acad. Sci., Arts, Lett. 46: 365-404. MILLER, W. H., D. L. ARCHER, H. M. Tyus, AND K. C. HARPER. 1982. White River fishes study. Final report. U.S. Fish Wildl. Serv., Colorado River Fish. Proj., Salt Lake City, Utah. MILLER, W. H., D. L. ARCHER, H. M. Tyus, AND R. M. McNatr. 1982. Yampa River fishes study. Final report. U.S. Fish Wildl. Serv., Colorado River Fish. Proj., Salt Lake City, Utah. MINCKLEY, W. L. 1973. Fishes of Arizona. Arizona Game and Fish Department, Phoenix, Arizona. Ono, R. P., J. D. WILLIAMS, AND A. WAGNER. 1983. Vanish- ing fishes of North America. Stone Wall Press, Inc., Washington, D. C. 257 pp. SMITH, G. R. 1981. Effects of habitat size on species rich- ness and adult body sizes of desert fishes. Pages 125-171 in R. J. Naiman and D. L. Soltz, eds., Fishes in North American deserts. John Wiley and Sons, New York. October 1986 SmiTH, M. L. 1975. Fishes of the Pliocene Glenns Ferry formation, southwest Idaho. University of Michi- gan Papers on Paleontology 14: 1-68. _____.. Late Cenozoic fishes of the warm deserts of North America: a reinterpretation of desert adaptations. Pages 11-38 in R. J. Naiman and D. L. Soltz, eds., Fishes in North American deserts. John Wiley and Sons, New York. Tyus, H. M. 1984. Loss of stream passage as a factor in the decline of the endangered Colorado squawfish. Pages 138-144 in Issues and technology in the management of impacted western wildlife. Tech. Pub. 14, Thorne Ecol. Inst., Boulder, Colorado. ____. 1985. Homing behavior noted for Colorado squawfish. Copeia 1985: 213-215. Tyus, H. M., AND C. W. McADa. 1984. Migration, move- ments and habitat preferences of Colorado squaw- fish, Ptychocheilus lucius, in the Green, White, and Yampa rivers. Colorado and Utah. Southwest. Nat. 29: 289-299. Tyus: COLORADO SQUAWFISH 661 Tyus, H. M., C. W. MCADa, AND B. D. Burpick. 1982. Green River fishery investigations: 1979-1981. Pages 1-99 in Final report of the U.S. Fish and Wildlife Service and U.S. Bureau of Reclamation, Part 2. U.S. Fish Wildl. Serv., Colorado River Fish. Proj., Salt Lake City, Utah. U.S. FISH AND WILDLIFE SERVICE. 1973. Threatened wildlife of the U.S. Resource Publ. 114. Washing- ton, D.C. —_—.. 1974. Colerado squawfish: determination as an endangered species. Fed. Reg. 45(80): 27710- 27713. WELCOMME, R. L. 1979. Fisheries ecology of floodplain rivers. Longman Group Ltd. London. 296 pp. WICK, E. J., D. L. STONEBURNER AND J. A. HAWKINS. 1983. Observations on the ecology of Colorado squaw- fish (Ptychocheilus lucius) in the Yampa River, Colorado, 1982. Technical Report 83-7, Water Re- sources Laboratory, Natl. Park Serv., Fort Collins, Colorado. 55 pp. PARASITES OF THE WOUNDFIN MINNOW, PLAGOPTERUS ARGENTISSIMUS, AND OTHER ENDEMIC FISHES FROM THE VIRGIN RIVER, UTAH Richard A. Heckmann’, James E. Deacon’, and Paul D. Greger” ABSTRACT.—Iwo hundred woundfin minnows, Plagopterus argentissimus, from four sites along the Virgin River, Utah, were examined on two dates during summer 1985. The foreguts of 211 woundfin and variable numbers of other fishes from the Virgin River near Beaver Dam Wash, Arizona, and Mesquite, Nevada, were examined for cestodes on four dates throughout 1979. Seven parasites were found in P. argentissimus: Posthodiplostomum minimum (metacer- cariae), Diplostomum spathaceum (metacercariae), Bothriocephalus acheilognathi, Gyrodactylus sp., Lernaea cypri- nacea, Trichodina sp., and Ichthyophythirius multifiliis. Fungal infections were noted on two fish during the study. Seventeen Virgin River roundtail chub, Gila robusta seminuda, were examined from two of the four sites in 1985 and 64 specimens from Beaver Dam Wash were examined in 1979. Gila robusta seminuda was infected with Posthodiplosto- mum minimum (metacercariae) and Bothriocephalus acheilognathi, the Asian fish tapeworm. This cestode probably gained entrance into the ichthyofauna of the Virgin River from red shiners, Notropis lutrensis, and has the potential of being very detrimental to the endemic and endangered fishes of the Virgin River. Parasite loads were correlated with water quality and habitat disturbance, with highest number and frequency occurring in “disturbed” sites. Low river flows and increased total dissolved solids appear to be associated with a higher parasite frequency and mean number in fishes of the Virgin River. These data represent the first known published records for parasites of the woundfin minnow and Virgin River roundtail chub. There is a paucity of information on the parasitofauna of the woundfin minnow, Plagopterus argentissimus, and other species of fish from the Virgin River, Utah-Arizona- Nevada. Many of the fishes in the Virgin River are endemic and have been listed as endan- gered species, the woundfin included. Hoff- man (1967) lists no parasites for the woundfin. Other common fishes in the Virgin River drainage include the Virgin roundtail chub, Gila robusta seminuda; speckled dace, Rhinichthys osculus; Virgin spinedace, Lepi- domeda mollispinis; desert sucker, Cato- stomus clarki; flannelmouth sucker, Cato- stomus latipinnis; and the introduced red shiner, Notropis lutrensis. Parasites of these later species are also poorly known (Hoffman 1967). Parasites can have adverse effects on fish populations. Changes in incidence, inten- sity, or parasite species infecting or infesting a host can provide important clues to the health and status of fish host populations. To under- stand management options, it is essential to also know the life cycle of the parasite and its effects at various levels of infection. For exam- ple, parasitism can be responsible for reduced growth rates, reduced egg production, poor swimming performance, aberrant behavior, ‘Department of Zoology, Brigham Young University, Provo, Utah 84602. “Department of Biology, University of Nevada, Las Vegas, Nevada 89154. etc. (David 1947, Dogiel 1958, Hoffman 1967). The primary objectives of this study were as follows: 1. Identify the species of parasites inhabiting the woundfin minnow at selected sites in the Virgin River in Utah. 2. Determine the frequency of occurrence, abundance, and temporal variation of parasites for P. argentissimus in the summer during the normal period of low flow. 3. Determine the relationship between parasitism and the immediate habitat of the host. 4. Determine the pathogenicity of the parasite to the host. In a previous study conducted during 1979 (Greger 1983), woundfin minnows examined during February and June near Beaver Dam Wash, Arizona, were not parasitized by ces- todes. Cestodes were present, however, in the foreguts of P. argentissimus and increased in both number and frequency from Septem- ber to December. Frequency of infection of woundfin with cestodes was much less near Beaver Dam Wash, Arizona, than was true further downstream in Nevada. This sug- gested that woundfin may be more vulnerable to parasitism in an agriculturally disturbed habitat than in a more natural environment. Because the Virgin River near St. George, 662 October 1986 Utah, exhibits similar agricultural distur- bances, our hypothesis was that the parasite load in the fish population near St. George would be higher than in the less disturbed sections of Virgin River in Utah. The life cycles of cestodes and other para- sites of fishes from the Virgin River have not been determined in detail. In general, fishes can be both definitive and intermediate hosts for cestodes. Reproductively mature adult cestodes present in fish will release eggs from gravid proglottids while in the intestine of the host. The eggs passs through the host’s anus and may settle in the stream sediments. For many cestodes, increasing temperature causes the operculate eggs to hatch, releasing aciliated coracidium (motile oncosphere). Cy- clopoid copepods (first intermediate host) in- gest these larvae and become infected. The oncosphere sheds its ciliated coat in the gut of the copepod and burrows through the intesti- nal wall to the hemocoel, where it develops into a procercoid larvae (Cheng 1973). This mesacestode-type larva cannot become infec- tive to a definitive host for about two to three weeks, until cercomer formation (Cheng 1973). If a fish ingests an infected copepod, the adult cestode may commence develop- ment in the intestine of the host. Although larval (pleroceroid) development in a second intermediate host (such as a smaller fish) is possible, it is unlikely to occur in the Virgin River ichthyofauna, since no primarily pisciv- orous fishes are present. It seems more proba- ble that development of the adult cestode in Virgin River fishes occurs following ingestion of an infected copepod. Direct development of the adult pseudophyllidean cestode Eu- bothrium salvelini following ingestion of in- fected Cyclops sp. by Sockeye salmon, Oncorhynchus nerka, has been reported by Smith (1973) from a lake in British Columbia. The effects of adult cestodes upon fish hosts have not been studied in detail. Rees (1967), after an extensive review of the literature, reached no definitie conclusions regarding the lethal effects of adult cestodes. Other in- vestigators have suggested that adverse ef- fects are considerable. Smith (1973) reported that noninfected salmon smolts grew 5%-7% longer and 17%-24% heavier than infected smolts. Field observations and swimming performance tests suggested cases of reduced swimming abilities and earlier fatigue in HECKMANN ETAL.: WOUNDFIN MINNOW PARASITES 663 salmon and trout infected with helminths (Smith and Margolis 1970, Heckman 1983). These data also demonstrate some nutritional or growth impairments for infected fish. Other researchers (Wardle 1933, Dogiel 1958, Dombroski 1955) have reported that adult pseudophyllidean cestodes can affect host nutrition and survival. Severe occlusion (impaction) of the gut has been reported for infected salmon (Dogiel 1964) and appeared to affect the nutritional status in severe cases. Impaction by parasites would also affect re- productive potential. Williams and Halvorsen (1971) negated the belief that contents of the fish host intestine represent an unlimited source of food for cestodes. Impaction and reduced growth and vigor would also reduce reproductive potential. Other indirect effects of cestode adults on their hosts have been demonstrated in the laboratory. Boyce and Clarke (1983) deter- mined that Sockeye salmon yearlings infected with tapeworms had a reduced ability to adapt to seawater as evidenced by increased mortal- ity and elevated plasma sodium levels. Boyce and Behrens-Yamada (1977) also reported Sockeye salmon juveniles infected with the same cestode, Eubothrium salvelini, to be more sensitive to zinc toxicity. These effects could make the infected fish less vigorous and more susceptible to predation during its sea- ward migration. The Asian fish tapeworm, Bothriocephalus acheilognathi, introduced into this country by imports of grass carp, has been described as a dangerous parasite in Europe (Bauer et al. 1981). This cestode has apparently become established in the ichthyofauna of southern Utah. The most common fish parasite found in this survey was the metacercarial state of the digenetic trematode, Posthodiplostomum min- imum. “Black spot,” for example, is caused by metacercarie found in melanin-pigmented cysts in the skin. This larval stage of flukes may be found in all tissues of fish (Spall and Summerfelt 1969b). MATERIALS AND METHODS The 1979 collections were made in Febru- ary, June, September, and December near Beaver Dan Wash, Arizona, and Mesquite, Nevada (Fig. 1). At each location, the foregut 664 4, te, Oe BYA peed » a is Sy. mys % esl q a7. Mvke 99 LEGEND “= Diversion We ge Town / / SCALE (0) 10 20 ESO GREAT BASIN NATURALIST Vol. 46, No. 4 Fig. 1. Fish sampling sites along the Virgin River, Utah, Nevada, Arizona, for this study. TABLE 1. Rating scale to indicate approximate density of the metacercarial infection in Virgin River fishes. Assigned Average number of metacercariae scale number per microscopic field at 20X 0 None 1 1-9 2 10-19 3 20-29 4 30-39 5 40-TNTC of 22-34 woundfin and variable numbers of other species was examined. Because the 1979 study was conducted as a part of a study of food habits (Greger 1983), we looked only for cestodes in the foregut. The 1985 collections were taken from the Virgin River on 27 and 28 July and 23, 24, 25, and 31 August 1985. Collections were made about 1/4 mile downstream from the inflow of the Santa Clara River, at Twin Bridges, about two miles below Berry Springs, and at Hurri- cane Bridge in Utah, and downstream in Ne- vada near Mesquite (Fig. 1). At each location 23-25 P. argentissimus and variable numbers of other fish species were examined. At Mesquite we examined 88 red shiners. In 1985 external examination for parasitism was made immediately following death of the fish. Scrapings of mucus and epithelial cells were taken from the gill surface, the base of the fins, and the lateral surface of the body. Scrapings were mixed in physiological saline and examined at 100X and 430X. Blood sam- ples were obtained from peripheral circula- tion on microscope slides, air dried, and later stained (Giemsa-Wright combination) prior to examination at 430X and 1000X. Each blood slide was.examined for a minimum of 10 min- utes. The abdominal cavity was opened ventrally and internal organs were examined. Each or- gan was removed and placed in a saline solu- tion prior to examination with a dissecting microscope. Eye tissues, including the lens, were also examined. Cestodes were excised from the intestinal tract, enumerated and fixed in AFA. Some individual worms were prepared for examina- tion by scanning electron microscopy by fixa- tion in 3% gluteraldehyde with an acrolein buffer. Viscera and gills from two woundfin from each of the four sampling locations were fixed in buffered 10% formalin and prepared for tissue evaluation at the Brigham Young Uni- versity laboratory. These samples were pro- cessed by standard methods, stained, and ex- amined closely to evaluate parasite pathology. Two stains, trichrome, and hematoxylin and eosin were used for the tissue sections. During the first day of study it was noted that many fish were heavily parasitized by metacercariae of the trematode, Postho- October 1986 HECKMANN ET AL.: WOUNDFIN MINNOW PARASITES 665 TABLE 2. Summary of parasite data from woundfin captured at each of four locations 27-28 July (N = 25) and 23-25 August (N = 23). August data are in parentheses. —————— oor Posthodiplostomum Number with Sample Average total Average other disease location length Percent scale number problems Twin Bridges 76.4 (75.7) 100 (96) 5.0 (4.0) 3/25° (5/23°) Santa Clara inflow 69.0 (69.8) 100 (100) 3.6 (2.7) 1/25° (5/23°) Hurricane Bridge 74.1 (80.8) 44 (57) 1.0 (1.0) 0 (0) Berry Springs 70.7 (64.9) 8 (22. <1.9 (<1.0) 0 (1/23°) 93/25 Eye fluke (metacercariae), Diplostomum spathaceum, fungal growth, dorsal fin. 51/25 Fungal growth, dorsal fin, skin erosion and fungal growth, pectoral fin; Gyrodactylus, gills. °5/23 Eye fluke (metacercariae), Diplostomum spathaceum, Roundworm, intestine, 3 fish. Trichodina, ciliated protozoan (gills), Anchor worm, Laernea, 2 fish. 45/93 Eye fluke (metacercariae), D. spathaceum, Anchor worm, Laernea, 3 fish. Trichodina, ciliated protozoan (gills), 2 fish. 1/23 Trichodina, ciliated protozoan (gills). diplostomum minimum. A rating scale was de- veloped to represent different densities of in- fection (Table 1). RESULTS Results of the parasitological examination are presented in Tables 2 to 6. These data show that the “disturbed” segments of the river (Mesquite, Twin Bridges, and Santa Clara inflow) carried a heavier parasite load than in undisturbed segments (Beaver Dam Wash, Hurricane Bridge, and Berry Springs) (Deacon and Hardy 1984). There is further indication that the trematode parasite, Posthodiplostomum minimum, in woundfin minnows in the “undisturbed” segment of the Virgin River in Utah may have increased slightly from July to August 1985 (Table 2). Whereas the trematode, Posthodiplostomum minimum, was clearly the major parasite in- fecting woundfin, other parasites were de- tected on some individuals (Table 2). A limited number of other fish were also examined for parasites (Tables 3 to 6). Of greatest concern was the presence of the Asian tapeworm, Bothriocephalus acheilo- gnathi, in the roundtail chub population. Table 5 shows frequency of occurrence and mean numbers of a cestode parasite in the foregut of all cyprinids occurring in Virgin River near Beaver Dam Wash, Arizona, in 1979. This cestode was later identified as Bothriocephalus acheilognathi. It is evident that frequency of infections varies seasonally and that the roundtail chub is more frequently infected with more tapeworms per fish than are other native species. The introduced red shiner appears to be about as heavily infected as is the roundtail chub. Table 5 also demon- strates that frequency and density of infection tends to increase in all species in September and December. Table 6 shows that woundfin in the dis- turbed river segment near Mesquite tend to be more heavily parasitized throughout the year than at Beaver Dam Wash. The red shiner is heavily parasitized by cestodes at both Mesquite and Beaver Dam Wash, but until 1985 remained abundant throughout the year only near Mesquite. Thirty-seven adult Asian tapeworms were identified from 24 (27%) of 88 red shiners collected in Virgin River near Mesquite, Ne- vada, on 25 August 1985. Mesquite, like Twin Bridges and Santa Clara inflow, is in a dis- turbed segment of the Virgin River. COMMENTS FOR SELECTED PARASITES OBSERVED DURING STUDY Posthodiplostomum minimum This was the most common parasite ob- served in fishes from the Virgin River in Utah. An excellent review article of North American studies of this fish parasite is found in Spall and Summerfelt (1969b). Metacercariae of the strigeid fluke, Posthodiplostomum minimum (MacCallum 1921), the white grub, are re- ported in most American helminthological surveys of fishes. The metacercaria was first reported over a century ago. It is enzootic in the U.S. exclusive of alpine regions and oc- curs in abundance in many of the 100 species of North American fishes that have been stud- ied to date (Hoffman 1967). The trematode larvae are generally so numerous in the liver, kidney, heart, and other viscera that many 666 GREAT BASIN NATURALIST Vol. 46, No. 4 TABLE 3. Results of parasite inventory for other fish species examined from Twin Bridges, 27—28 July (1-12) and 23 August (13-15) 1985. Fish Measurements (mm) Species number TL SL Parasites Flannelmouth Sucker 1 115 90 No parasites (Catostomus latipinnis ) Virgin River Roundtail 1 160 145 Metacercariae: Chub (Gila robusta Posthodiplostomum seminuda ) 2 145 130 Metacercariae: Posthodiplostomum 3 152 139 Metacercariae: Posthodiplostomum Asian Tapeworm: Bothriocephalus 4 155 141 Metacercariae: Posthodiplostomum Asian Tapeworm: Bothriocephalus Anchor worm: Laernea 5 142 130 Metacercariae: Posthodiplostomum 6 136 123 Metacercariae: Posthodiplostomum i 134 124 Metacercariae: Posthodiplostomum 8 145 132 Metacercariae: Posthodiplostomum 9 135 126 Metacercariae: _ Posthodiplostomum 10 125 115 Metacercariae: Posthodiplostomum II 160 145 Metacercariae: Posthodiplostomum 12 145 130 Metacercariae: Posthodiplostomum 13 170 142 Metacercariae: Posthodiplostomum 14 170 138 Metacercariae: Posthodiplostomum 15 135 112 Metacercariae: Posthodiplostomum TABLE 4. Results of parasite inventory for other fish species examined from mainstream Virgin River about 1/4 mile below inflow of Santa Clara River, 27—28 July 1985. Fish Measurements (mm) Species number WL SL Parasites Flannelmouth Sucker 1 115 90 No parasites (Catostomus latipinnis ) 2 164 140 No parasites Largemouth Bass 1 58 50 Metacercariae: (Micropterus salmoides ) Posthodiplostomum 2 54 47 Metacercariae: Posthodiplostomum Virgin River Roundtail 1 155 145 Metacercariae: Chub (Gila robusta seminuda) Posthodiplostomum Asian Tapeworm: Bothriocephalus 2 165 146 Metacercariae: Posthodiplostomum | : | October 1986 HECKMANN ET AL.: WOUNDFIN MINNOW PARASITES 667 TABLE 5. Frequency of occurrence (%) and mean number of the cestode, Bothriocephalus acheilognathi, (Order: Pseudophyllidea) from the foreguts of fishes collected from the Virgin River near Beaver Dam Wash, Arizona, 1979. Abbreviations are as follows: R = Rhinichtnys osculus; G = Gila robusta; L = Lepidomeda; P = Plagopterus; N = Notropis lutrensis. Month R G N 28 20 February % 3.7 35.0 Mean 0.3 0.6 SE 0.1 0.2 N D7 4 June % 0.0 25.0 Mean 0.0 0.8 SE 0.0 0.8 N 29 31 September % 18.2 66.6 Mean 0.5 2.0 SE 0.2 0.5 N 23 9 December % 52.0 88.8 Mean 93,0 16.8 SE 1.2 6.0 Note: Asterisks denote a significant difference among species at p < 0.01. ND = No data. SE is the standard error of the mean. TABLE 6. Frequency of occurrence (%) and mean num- ber of cestode parasites (Order: Pseudophyllidea) from the foreguts of fishes from the Virgin River in Nevada, 1979. Anasterisk denotes a significant difference between species at p < 0.01. Plagopterus Notropis Month argentissimus _ lutrensis t N 24 48 February % 33 69 Mean 0.54 2.94 3.94* N 23 25 June % 13 36 Mean 0.13 3.76 2.01 N 34 29 September % 53 86 Mean 1.76 6.03 3.45* N 27 th December % 37 81 Mean 0.63 Cree) HW observers have implicated them as being histopathogenic. The pathogenicity of the lar- val stage is usually due to compression or occlusion of a vital organ. Early literature concerning the classifica- tion of Posthodiplostomum minimum is in- vested with synonymy and misinformation, partly because of inadequate description and erroneous identification and partly because some larval stages were described before their life histories, especially the adult, were F. ID N Ratio 24 22 4.1 0.0 ND 0.04 0.0 8.18* 0.1 26 ND 0.0 ND 0.0 31 28 26 12.0 10.7 84.6 0.2 0.1 3.9 28.04* 0.2 0.1 0.6 Mi 27 27 7.4 33.0 88.8 1.7 0.6 9.3 23.14* Loa 0.3 2.0 known. Nomenclatural history has been re- viewed by Miller (1954), Hoffman (1958), and Bedinger and Meade (1967). Studies by Hunter (1937) on the transfor- mation of Cercaria multicellulata to Neascus van cleavei and by Ferguson (1938) on trans- formation of metacercariae of N. van cleavei to adult Neodiplostomum culminated in the first description of the life cycle of Posthodiplosto- mum minimum. Metacercariae have been found in all vis- ceral organs but occur in abundance in the liver, spleen, kidneys, mesenteries, sinus venosus, heart, and ovaries (Figs. 2a, 2b, and 3a, 3b). Some strigeid larvae show positive histotropic effects toward specific fish tissue in vitro (Davis 1936). Avault and Allison (1965) found that the heart, liver, and kidneys con- tained approximately 79% of the total metac- ercariae in bluegill (Lepomis machrochirus). Metacercariae have not been reported in fish testis, apparently the only visceral organ alien to this parasite. The occurrence of numerous metacercariae in visceral organs suggests deleterious effects on the well-being of the host and implicates P. minimum as a cause of mortality or morbidity. Hunter (1937, 1940) stated that death resulted if sufficient liver or other visceral tissue were destroyed by the metacercariae. Hughes (1928) observed bluegill, which had a heavy 668 GREAT BASIN NATURALIST Vol. 46, No. 4 Fig. 2a, b. The metacercarial stage of Posthodiplostomum minimum (P) from the internal viscera of Plagopterus argentissmus. Note encapsulation (E) of the larval trematode and organ compression of the liver (L). Hemorrhaging (R) has occurred near the site of metacercarial encapsulation. Magnification, 2a—100X, 2b—430X. October 1986 HECKMANN ETAL.: WOUNDFIN MINNOW PARASITES 669 * . eZ Wed, Se ec | | al # _— its Fig. 3a, b. The larval stage of Posthodiplostomum minimum (P) from the internal viscera of Plagopterus argentissmus. 3a represents metacercariae encapsulated (E) in outer layer (F) (fibro serosa) of the intestinal tract. Note muscularis externa (M) (100X). 3b represents metacercariae encapsulated (E) in the mesentaries (M) of the viscera exhibiting compression on the intestine (N) (100X). 670 infection of P. minimum metacercariae, in Fife Lake, Michigan, dying in large numbers. This represents an example of a questionable cause-and-effect relationship because the par- asite is ubiquitous and usually numerically abundant. More than circumstantial evidence is required to substantiate an allegation of P. minimum being a cause for fish mortality. Wild fish, with several hundreds of encysted metacercariae in the liver, sinus venosus, heart, and kidneys, are often observed to suf- fer no obvious debilitating effects. Colley and Olsen (1963) found as many as 991 metacer- cariae per bluegil, with metacercariae so dense as to be clumped en masse. Spall and Summerfelt (1969a) have observed 2,041 metacercariae in a bluegill from an Oklahoma reservoir. Mortality due to stress and trauma from penetration of the cercariae has been ob- served in the laboratory following exposure of suitable host fish to high numbers of cercariae (Hunter 1937, Bedinger and Meade 1967). Host reactions following cercarial penetration include petechial hemorrhage at the site of invasion followed by congestion of surround- ing venules and local edema, and an aggrega- tion of leucocytes at the point of entry, partic- ularly the phagocytic elements. Pathological effects include increased rate of excretion, in- creased plasma globulin and albumin, in- creased liver respiration, and decreased he- matocrits (Smitherman 1964). Hemorrhage or a decrease in erythropoiesis would reduce he- matocrits. The increase in the plasma proteins may represent a homeostatic response to the nutritional demands of the parasite, altered liver function, or effects on capillary perme- ability. After encystment (19 days), mortality infre- quently occurs. There is no experimental evi- dence to indicate mortality or other detrimen- tal effects from the occurrence of encysted metacercariae. Bothriocephalus acheilognathi = (B. gowkongenesis = B. opsalichthydes) The Asian fish tapeworm, Bothriocephalus acheilognathi, represents a new introduction in North America, brought in through imports of grass carp to this country from China. Be- cause of the new introduction and size of the GREAT BASIN NATURALIST Vol. 46, No. 4 Fig. 4. A scanning electron micrograph of the Asian fish tapeworm, Bothriocephalus acheilognathi, from the in- testine of the Virgin River roundtail chub, Gila robusta seminuda. Note the pit-viper shaped scolex (SC) and numerous proglottids (P). Photographed at 100X magnifi- cation. adult worm, this parasite has become of major concern to fish and game officials throughout the country. Excellent reviews of the histo- pathology, biology, life history, control, and management of Bothriocephalus are found in a series of papers by Nakajima and Egusa (1947a, b, c, 1976a, b). The Asian fish tapeworm, characterized by its arrow or heart-shaped scolex (Fig. 4), has been a dangerous parasite for cultured grass carp and German carp fingerlings in Europe (Bauer et al. 1981). In Europe it has also been found in European catfish, guppies, mosquito fish, and other species (Hoffman 1983, Hoff- man and Shubert 1984). In the United States it has been found in golden shiners and fat- head minnows (Hoffman 1976), as well as in grass carp, Colorado squawfish, and mosquito fish. We add speckled dace, roundtail chub, Virgin spinedace, woundfin, and red shiner to that list. The best known carp parasite transported to the fish ponds of many countries with the October 1986 HECKMANN ET AL.: WOUNDFIN MINNOW PARASITES 671 Fig. 5a, b. Trichodina (T) infesting the gill lamellae (G) of Plagopterus argentissmus. Note the macronucleus (M) and cilia (arrow) characteristic of this ciliate. There is tissue granulation and hypertrophy (gh) of the host gill tissue near one parasite. 1000X magnification. 672 Chinese carp is Bothriocephalus acheilo- gnathi ( = B. gowkongensis, = B. op- salichthydis) (Bauer et al. 1981). All Eu- ropean countries that culture carp in large quantities now have this pathogen. The spread of this parasite to new localities usually results in heavy infection of young fishes dur- ing the first years after it appears. Bothrio- cephalus acheilognathi, athermophyllic para- site, can infect many fish species. Presumably it traveled to the United States by airplane in grass carp shipped from Asia. Trichodina: Trichodinosis This is a ciliated protozoan that was com- monly observed on the gills of the woundfin minnow in August (Table 2). This ciliate is ubiquitous among fish para- sites throughout the world and usually is of minor concern for fish health. Members of the genus Trichodina Ehrenbert (Family; Urce- olariidae Stein) are commonly seen. on ail kinds of aquatic animals. In fish they may settle on the skin in such numbers as to ob- scure the normal structure (Fig. 5a, 5b), and they are easily recognized by their similarity to a suction disk. Classification methods have been reviewed by Tripathi (1954). Trichodina parasitizes the skin, gills, and urinary bladder of the fish and is found both in freshwater and the sea. The species of Trichodina that occur on North American freshwater fishes have not received the attention that they have in other areas of the world even though they are one of the most important groups of ectoparasites of freshwater fishes. Frequent references to tri- chodinids parasitizing fish occur in fish cul- ture literature but the species are not named, as in the case for our study. Only Mueller (1937, 1938), David (1947), Lom (1963), Lom and Hoffman (1964), Hoffman (1967), and Wellborn (1967) have made major contribu- tions to the knowledge of the taxonomy and distribution of Trichodina of North American freshwater fishes. Because of their small size, supposed lack of specific characters, and diffi- culty of removal from their hosts, they have been largely ignored. More than 90 species of Trichodina have been described from the skin and gills of marine and freshwater fishes of the world (Hoffman 1967, Wellborn 1967). Many of GREAT BASIN NATURALIST Vol. 46, No. 4 these were described as new only because they were found on a different host or in an- other geographic location. In many cases the descriptions were inadequate since the uni- form body structure of these ciliates yields few characters for differentiation of the species (Lom 1961, 1970, Mueller 1937). The inexact and insufficient descriptions of most early au- thors make the identification of many species doubtful. But the recently employed silver- impregnation technique of Klein and Chat- ton-Lwoff (Corliss 1953) reveals details of the adhesive disk that are important features of trichodinid taxonomy. Padnos and Nigrelli (1942) used the silver-impregnation tech- nique to determine the ciliar patterns of tri- chodinids. But, according to Lom (1958), Raabe (1950) was the first to employ this tech- nique in the study of the structure of the adhesive disk. Trichodina rarely give rise to pathological manifestations of disease. It may be sporadi- cally found in living fish, but it will only multi- ply in moribund and weakened ones. A macronucleus, mironucleus, and numerous food vacuoles are to be seen in the cytoplasm. In our study we observed Trichodina on the gill surface (Fig 5a, 5b) of woundfin minnows in August. DISCUSSION Deacon and Hardy (1984) referred to seg- ments of the Virgin River above Washington Diversion and above Mesquite Diversion as relatively undisturbed. Segments of the river below these two diversions were referred to as disturbed largely by irrigation withdrawals. Table 7 demonstrates that mean and mini- mum flows in May-November were substan- tially reduced at Bloomington below the Washington Diversion in 1985. Table 8 demonstrates a similar reduction in flow at Riverside, below the Mesquite Diversion in 1979. In addition, water quality in disturbed segments of Virgin River is also reduced, largely as a consequence of its use for agricul- tural irrigation (Sandberg and Sultz 1985). In June 1985 discharge from a salt spring (Pah Tempe Spring) suddenly increased dramati- cally, resulting in degradation of water quality throughout the Utah portion of the Virgin River (Table 9). Therefore, our 1979 data clearly contrast parasite loads in fishes ex- October 1986 HECKMANN ETAL.: WOUNDFIN MINNOW PARASITES 673 TABLE 7. Provisional mean, maximum, and minimum daily discharge (cfs) of the Virgin River at Hurricane and Bloomington, Utah, in 1985. Month January February March April May June July August September October November December Mean 197 201 270 637 229 96.9 116 86.7 79.1 76.7 183 171 Hurricane Maximum 361 266 515 Minimum Mean 238 220 Bloomington Maximum Minimum 433 198 301 169 574 171 1,050 285 477 63 150 25 481 31 49 28 132 32 142 73 808 88 319 119 TABLE 8. Mean, maximum, and minimum daily discharge (cfs) of the Virgin River at Littlefield, Arizona, and Riverside, Nevada, in 1979. Month January February March April May June July August September October November December Mean 200 324 603 1,262 1,559 390 68.9 127 65.9 92.3 171 194 Littlefield Maximum Minimum 458 139 389 256 2,440 252 1,840 760 2,000 962 815 79 79 63 637 70 68 63 128 65 211 137 219 173 Riverside Maximum 591 447 Minimum 178 279 280 692 789 182 TABLE 9. Specific conductance, Virgin River, pre- and postdevelopment of a dramatic increase of flow from Pah Tempe Springs. Location of water sample Pah Tempe Springs Below Pah Tempe Below Ash Creek Above Quail Creek Hurricane Bridge or Berry Springs Above wash diversion or at inlet Above Twin Bridges or above Fort Pierce Wash Below Bloomington posed to nearly natural conditions near Beaver Dam Wash with fishes exposed to re- duced flow and water quality near Mesquite. Predevelopment Sandberg Hickman and Sultz 1984-85 1985 16,000-17,500 12,600-—13,000 5,900—6,600 850-4, 430 1,500-—1,800 905-3, 700 1,600—2, 100 825-3, 140 1,600—1,900 850-3, 120 1,600-—1,900 820-3, 450 1,700—2, 200 890-4, 390 2,000—2, 2000 870—4, 200 Postdevelopment Hickman 1985 9,400—10,900 6,500—7,900 5,000—8,000 3,900—5, 000 3, 900-6, 500 4,000—5,000 3,900—4, 800 Our 1985 data, though designed to contrast the same environmental conditions in a differ- ent segment of Virgin River, have the addi- 674 tional complication of a sudden reduction in water quality throughout the entire Utah seg- ment of the river one to two months prior to the time of our collections. Nevertheless, it is clear that the parasite burden in fishes of Vir- gin River living in more disturbed habitats is much higher than for the same species living in less disturbed habitats. Cestodes (Bothriocephalus acheilognathi) occurred in all Cyprinids inhabiting the lower mainstream of the Virgin River in Arizona and Nevada in 1979 (Tables 5, 6). The exotic red shiner, Notropis lutrensis, was infected more frequently and with a greater number of ces- todes per individual than were all native spe- cies except the Virgin roundtail chub, Gila robusta seminuda. Infection frequency and density were variable seasonally, with heavi- est parasite loads in general occurring in fall and winter and lightest parasite loads in sum- mer (Tables 5, 6). The woundfin in the “dis- turbed” segment of the lower mainstream (Table 6) was more heavily infected than was the population in the “undisturbed” segment (Table 5). The red shiner was equally heavily infected in both river segments (Tables 5, 6). In 1985 cestodes were not detected in woundfin minnows at any of the four locations sampled in Utah. Since we sampled only dur- ing late July and August, it is possible that the absence of cestodes reflects only seasonal vari- ation. Both frequency and density of trematode infection in woundfin, however, are much greater in the lower, more dis- turbed portion of the river near St. George (Table 2). The reduced flows near St. George may force fish into slow-flowing pools or ponded waters. The more ponded conditions proba- bly permit the development of dense popula- tions of cercariae that are released by snails, an intermediate host in the trematode life cycle. These same conditions make the fish more vulnerable to piscivorous birds, facilitat- ing completion of the trematode life cycle. The degraded water quality resulting from agricultural return flows may also increase stress on the fish population, which may mani- fest itself in an increased parasite load. If this is a significant factor, the increased total dis- solved solids from increased flows of Pah Tempe Springs beginning in summer 1985 (Table 9) may result in an increased parasite burden in fishes throughout the Utah portion GREAT BASIN NATURALIST Vol. 46, No. 4 of the Virgin River. The discovery of the Asian fish tapeworm, B. acheilognathi, in the fish population of the Virgin River is of major concern. The parasite was probably introduced into the U.S. by the grass carp. It is considered to be especially dangerous in Europe, where it was also intro- duced. W. L. Minckley (personal communi- cation) reports that it is especially damaging to Colorado squawfish at the Dexter National Endemic Fish Hatchery in Deming, New Mexico. The high incidence of infection with large numbers of tapeworms in the endan- gered roundtail chub near Beaver Dam Wash (Table 5) demonstrates the probability that it is similarly damaging to roundtail chubs. Other native fish species showed lower inci- dence of infection, but all cyprinids were in- fected at leveis that could severely damage the populations. The exotic red shiner was as susceptible to infection at the roundtail chub. Of the 17 chubs we examined from the river near St. George in July and August 1985, only 3 (18%) contained Asian fish tapeworms. We suggest that this is most likely a reflection of the fact that the tapeworm is just beginning to establish itself in the Utah segment of the river. The tapeworm probably arrived in the spring of 1984 along with the red shiner, Notropis lutrensis, which first appeared in April collections of fish from the Virgin River. Red shiners first became common in the St. George portion of the Virgin River during the summer of 1985. They were one of the most heavily infected species in the lower river in 1979 (Tables 5, 6). Twenty-four of 88 red shin- ers (27%) collected from the lower Virgin River near Mesquite, Nevada, on 15 August 1985 contained 37 adult Asian tapeworms. The red shiner, an excellent host for the Asian tapeworm, was recently established in the Utah portion of the Virgin River and probably brought the cestode into Utah. ACKNOWLEDGMENTS We thank Drs. Glenn Hoffman and Lauritz A. Jensen for the identification of the Asian fish tapeworm. This study was funded by the Utah Division of Wildlife Resources (Randy Radant). James Allen and the Electron Optics staff at Brigham Young University provided space and help for the scanning electron mi- croscopy part of this study. October 1986 LITERATURE CITED AVAULT, J. W., JR., AND R. ALLISON. 1965. Experimental biological control of a trematode parasite of bluegill. Expt. Parasitol. 17: 296-301. BAUER, O. N., S. EcusA, AND G. L. HOFFMAN. 1981. Para- sitic infections of economic importance in fishes. Pages 425-443 in W. Slusarski, ed., Review of advances in parasitology. Polish Academy of Sci- ence. BEDINGER, C. A., JR., AND T. G. MEADE. 1967. Biology ofa new cercaria for Posthodiplostomum minimum (Trematoda: Diplostomidae). J. Parasitol. 53: 985-988. Boyce, N. P., AND S. BEHRENS YAMADA. 1977. Effects of a parasite Eubothrium salvelini (Cedstoda: Pseudo- phyllidea) on the resistance of juvenile sockeye salmon (Oncorhynchis nerka) from Banine Lake, British Columbia. Canadian J. Fish. Aquat. Sci. 34: 706-709. Boyce, N. P., AND W. C. CLarKE. 1983. Eubothrium salvelini (Cestoda: Pseudophyllidea) impairs sea- ward adaptation of migrant sockeye salmon year- lings Oncorhynchus nerka from Banine Lake, British Columbia. Canadian J. Fish. Aquat. Sci. 40: 821-824. CHENG, T. C. 1973. General parasitology. Academic Press. 965 pp. CoLLey, F. C., AND A. C. OLSON. 1963. Posthodiplosto- mum minimum (Trematoda: Diplostomidae) in fishes of lower Otay Reservoir, San Diego County, California. J. Parasitol. 49: 149. Cor iss, J. O. 1953. The Chatton-Lwoff silver impregna- tion technique. Stain Tech. 28: 97-100. Davis, H. S. 1936. Pathological studies on the penetration of the cercaria of the strigeid trematode, Diplosto- mum flexicaudum. J. Parasitol. 22: 329-337. —___.. 1947. Studies of the protozoan parasites of fresh- water fishes. U.S. Dept. Interior Fishery Bull. #41: 1-29. DEACON, J. E., AND T. B. Harpy. 1984. Streamflow re- quirements of woundfin (Plagopterus argentis- simus ): Cyprinidae in the Virgin River, Utah, Ari- zona, Nevada. Festschrift for Walter W. Dalquest. Pages 45—56 in N. V. Horner, ed. Mid- western State University Press. DOoGIEL, V. A. 1958. Ecology of parasites of freshwater fishes. In Parasitology of fishes. Oliver and Boyd, Edinburgh. —___.. 1964. General parasitology. Oliver and Boyd, Ed- inburgh and London. DomsrOsKI, E. 1955. Cestode and nematode infection of sockeye smolts from Banine Lake, British Colum- bia. J. Fish. Res. Bd. Canada 12: 93-96: FERGUSON, M. S. 1938. Experimental studies on Posthodiplostomum minimum (MacCallum, 1921), a trematode from herons. J. Parasitol. 24 (Suppl.): 31. GREGER, P. D. 1983. Food partitioning among fishes of the Virgin River. Unpublished thesis, University of Nevada, Las Vegas. HECKMANN, R. A. 1983. Eye fluke (Diplostomum spathaceum) of fishes from the Upper Salmon River near Obsidian, Idaho. Great Basin Nat. 43: 675-683. HECKMANN ET AL.: WOUNDFIN MINNOW PARASITES 675 HOFFMAN, G. L. 1958. Experimental studies on the cercaria and metacercaria of a strigeid trematode, Postho- diplostomum minimum. Expt. Parasitol. 7: 23-50. . 1967. Parasites of North American freshwater fishes. University of California Press, Berkeley and Los Angeles. 486 pp. . 1976. The Asian tapeworm, Bothriocephalus gow- kongenesis, in the United States, and research needs in fish parasitology. Proceedings Fish Farming Con- ference and Annual Convention Catfish Farmers of Texas, Texas A & M University 1976: §4—90. . 1983. Asian fish tapeworm, Bothriocephalus op- sarichthydis, prevention and control. Fish Dis- ease Leaflet: USFWS: USDA: 1-4. HOFFMAN, G. L., AND G. SCHUBERT. 1984. Some parasites of exotic fishes. Pages 233-261 in W. R. Courtney and J. R. Staffer, eds., Distribution, biology, and management of exotic fishes. Johns Hopkins Uni- versity Press, Baltimore and London. Hucues, R. C. 1928. Studies on the trematode family Strigeidae (Holostomidae) No. IX. Neascus van cleavei (Agersborg). Trans. Amer. Micro. Soc. 47: 320-341. HUNTER, G. W., ITI. 1937. Parasitism of fishes in the lower Hudson area. Pages 264—273 in A biological sur- vey of the lower Hudson watershed. Biological Survey No. XI (1936), Suppl. to the Twenty-sixth Ann. Rept., New York State Conservation Dept. . 1940. Studies on the development of the metacer- caria and the nature of the cyst of Posthodoplosto- mum minimum (MacCallum, 1921) (Trematoda; Streigeidae). Trans. Amer. Micro. Soc. 59: 52-63. Lo, J. 1958. A contribution to the systematics and mor- phology of endoparasitic trichodinids from am- phibians, with a proposal of uniform specific char- acteristics. J. Protozool. 5: 251-263. . 1961. Protozoan parasites found in Czechoslo- vakian fishes. I. Myxosporidia, Suctoria. ool. Listy Fol. Zool. 10(24): 45-58. . 1963. The ciliates of the family Urceoloariidae inhibiting gills of fishes (the Trichodinella-group). Acta Soc. Zool. Bohemoslov. 27: 7-19. . 1970. Observations on trichodinid ciliates from freshwater fishes. Archiv f, Protistenkunde 112: 153-177. Lom, J.,ANDG. L. HOFFMAN. 1964. Geographical distribu- tion of some species of Trichodinids (Ciliata: Per- itricha) parasitic on fishes. J. Parasit. 50: 30-35. MILLER, J. H. 1954. Studies on the life history of Posthodiplostomum minimum (MacCallum, 1921). J. Parasitol. 40: 255-270. MUELLER, J. F. 1937. Some species of Trichodina (Ciliata) from freshwater fishes. Trans. Amer. Micr. Soc. 56: 117-184. ____. 1938. A new species of Trichodina (Ciliata) from the urinary tract of the muskalonge, with a reparti- tion of the genus. J. Parasit. 23: 251-258. NakajIMA, K., AND S. Ecusa. 1974a. Bothriocephalus op- sariichthydis Yamagutii (Cestoda: Pseudophyl- lidea) found in the gut of cultured carp, Cyprinus carpio (Linne)—I. Morphology and taxonomy. Fish Pathology 9(1): 31-39. . 1974b. Bothriocephalus opsariichthydis Yamagutii (Cestoda: Pseudophyllidea) found in the gut of cul- tured carp, Cyprinus carpio (Linne)—II. Incidence and histopathology. Fish Pathology 9(1): 40—45. 676 _____. 1974c. Bothriocephalus opsariichthydis Ya- magutii (Cestoda: Pseudophyllidea) found in the gut of cultured carp, Cyprinus carpio (Linne)— III. Anthelmintic effects of some chemicals. Fish Pathology 9(1): 46-49. .1976a. Bothriocephalus opsariichthydis Ya- magutii (Cestoda: Pseudophyllidea) found in the gut of cultured carp, Cyprinus carpio (Linne)— IV. Observations on the egg and coracidium. Fish Pathology 11(1): 17-22. .1976b. Bothriocephalus opsariichthydis Ya- magutii (Cestoda: Pseudophyllidea) found in the gut of cultured carp, Cyprinus carpio (Linne)—V. Ovicial effects of drying, freezing, ultraviolet rays, and some chemicals. Fish Pathology 11(1): 23-26. PapNnos, M., AND R. F. NIGRELLI. 1942. Trichodina spheroidesi and Trichodina halli spp. nov. para- sitic on the gills and skin of marine fishes, with special reference to the life-history of T. spheroidesi. Zoologica 27: 65-72. RaaBE, Z. 1950. Uwagi o Ureceolariidae (Ciliata-Per- itricha) skrzel ryb. Ann. Univ. M. Curie- Sklodowska, Lublin 5: 292-310. REES, G. 1967. Pathogenesis of adult cestodes. Helminth. Abstracts (36): 1-23. SANDBERG, G. W., AND L. G. SULTZ. 1985. Reconnaissance of the quality surface water in the Upper Virgin River Basin, Utah, Arizona, and Nevada. 1981-82. Utah Dept. of Natural Resources Tech. Bull. 83: 1-69. SMITH, H. D. 1973. Observations of the Cestode Euboth- rium salvelini in juvenile sockeye salmon (Oncorhynchus nerka Walbaum) at Banine Lake, British Columbia. J. Fish. Res. Bd. Canada 30: 947-964. GREAT BASIN NATURALIST Vol. 46, No. 4 SmiTH, H. D., AND L. MARGOLIS. 1970. Some effects of Eubothrium salvelini Schrank (1970) on sockeye salmon (Oncorhynchus nerka Walbaum) in Ba- nine Lake, British Columbia. J. Parasitol. 56(11): 321-322. SMITHERMAN, R. O., Jr. 1964. Effects of infections with the strigeid trematode, Posthodiplostomum minimum (MacCallum), upon the bluegill, Lepomis macro- chirus Rafinesque. Unpublished dissertation, Auburn University, Auburn, Alabama. 55 pp. SPALL, R. D., AND R. C. SUMMERFELT. 1969a. Host-para- site relations of certain endoparasitic helminths of the channel catfish and white crappie in an Okla- homa reservoir. Bull. Wild. Dis. Assoc. 5: 48-67. . 1969b. Life cycle of the white grub, Postho- diplostomum minimum (MacCallum 1921: Trema- toda, Diplostomatidae), and observations on host- parasite relationships of the metacercariae in fish. Pages 218-230 in S. F. Snieszko, ed., A sympo- sium of diseases of fishes and shellfishes. Special Publ. No. 5 AFS. TRIPATHI, Y. R. 1954. Studies on parasites of Indian fishes. III. Protozoa. 2. (Mastigophora and ciliophora). Rec. Indian Mus. 52: 221-230. WARDLE, R. A. 1933. The parasitic helminths of Canadian animals: the Cestodaria and Cestoda. Canadian J. Res. 8: 317-333. WELLBORN, T. L., JR. 1967. Tricodina (Ciliata: Urceolari- idae) of freshwater fishes of the southeastern United States. Journal of Protozoology 14(3): 399-412. 25 WituiAMS, H. H., AND O. HALvorsEN. 1971. The inci- dence and degree of infection of Gadus morhua L. 1758 with Aborthrium gadi Beneden 1871 (Ces- toda Pseudophyllidea). Norway J. of Zoology 19: 193-199. NEW SCLEROCACTUS (CACTACEAE) FROM NEVADA Ken Heil’ and Stanley L. Welsh” ABSTRACT.—Named as new is Sclerocactus schlesseri Heil & Welsh. The taxon is described and its relationships discussed. During the summer of 1981 a peculiarly adapted population of Sclerocactus was dis- covered by cactus enthusiast Dr. David Schlesser in the southeastern quadrant of Ne- vada, growing on a peculiar, Tertiary lacus- trian deposit. The substrate consists of sandy silts to silty clays and appears on the surface to be somewhat gypsiferous. Vegetation in the region consists of typical salt desert shrub community, with galleta (Hilaria jamesii) as the main perennial grass component. The long, ribbonlike, uppermost central spines simulate the leaves of the galleta, and the stems are difficult to discern among the grassy areas between the shrubs. The plants occur singly or in small clumps. General aspect is that of Sclerocactus whip- plei in a broad sense, but the stem diameter of the plants examined is not as great (4-8 cm, not 5-15 cm), flowers average smaller (8—4 cm, not 3.5—5 cm long), and the spines are densely pubescent, at least when young. The pubescent spines and characteristic flattened upper central spine approaches the condition found in S. pubispinus of the nearby Great Basin. The flowers average larger than in S. pubispinus (3—4 cm, not 2.5—3.5 cm) and the upper central spine is longer (8—5.5 cm, not 0.5-3.5 cm). The locality is intermediate be- tween that of S. pubispinus and S. whipplei var. roseus as interpreted by Welsh (1984). A specimen of the latter was taken along the Virgin River west of Bunkerville, Nevada, by _N. D. Atwood (7821b BRY) in May 1981. A locality for S. whipplei (as S. parviflorus var. intermedius) is mapped from Iron County in southwestern Utah by Benson (1982). Although some features of the plant discov- ered by Dr. Schlesser are intermediate be- | Department of Biology, San Juan College, Farmington, New Mexico, 87401. tween S. whipplei and S. pubispinus, there are some features that are evidently unique. The narrow stems suggest a parameter that is not shared by the two close geographical con- geners. Because of the discordant as well as intermediate features the plant is named as follows: Sclerocactus schlesseri Heil & Welsh sp. nov. Persimilis Sclerocacto whipplei (En- gelm.) Britt. & Rose sed in caulibus angus- tioribus (4—8 cm nec 5-15 cm), floribus mi- noribus (83-4 cm nec 3.5-5 cm), et spinis pubescentibus differt, et similis Sclerocacto pubispino (Engelm.) L. Benson in floribus majoribus (3—4 cm nec 2.5—3.5 cm) et spinis superioribus centralis differt. Stems dark green, typically solitary, ellip- soid to obovoid, 3—10 (14) cm tall, 4—6 (8) cm wide; ribs 13; tubercules 12-18 mm wide, 8-10 mm long; areoles woolly, finally glabrate; spines flexible, densely pubescent when young; upper central spine 1, ascend- ing, flat or trigonous, ribbonlike, curved, car- tillaginous to bony, pubescent to glabrous, 3-5.5 cm long, 1-2.5 mm wide, reddish brown to white; peripheral central spines 2, ascending, flat, ribbonlike, sometimes hooked, pubescent, 2-3 cm long, 0.5—1 mm wide, black to white; lower central spine 1, ascending, flexible, irregularly hooked, pubescent, 2.5—4.5 cm long, to 1 mm wide, black, gray, tan, or white; radial spines 6-9 (12), flattened, flexible, pubescent, 3-14 mm long, white; flowers apical on upper end of the tubercules near the top of the areoles and above the spines, funnelform, 3—4 cm long, 2-3 cm wide, violet pink; sepaloid perianth parts oblanceolate, finely irregularly toothed apically, mucronate, violet pink with brown- 2Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 677 678 ish midstripes, 1.5—2 cm long, 6-10 mm wide; petaloid perianth parts oblanceolate, entire or slightly undulate, minutely toothed apically, violet pink, 1.5—2 cm long, 4-5 mm wide, filaments red, anthers yellow; stigma lobes 7-9, light green; fruit dry, barrel shaped, naked or with 1 or 2 scales, 1-1.5 cm long, 9-13 mm wide, red to greenish red, dehiscing by a transverse break in the ovary wall, the perianth persistent; seeds 2 mm long (hilum to point opposite), 3 mm wide, ca 1 mm thick, pyriform with slightly flattened apex, GREAT BASIN NATURALIST Vol. 46, No. 4 shiny black, papillate, becoming less so near the hilum; hilum elliptic, 1 mm wide. TypE.—USA. Nevada. Lincoln County, Tertiary lacustrian deposit at ca 1,464 m in salt desert shrub-grass community, 16 June 1983, K. Heil s.n. (Holotype BRY; isotype NY). LITERATURE CITED BENSON, L. 1982. The cacti of the United States and Canada. Stanford University Press, California. 1044 pp. WELSH, S. L. 1984. Utah flora: Cactaceae. Great Basin Nat. 44:52-69. NEW SPECIES AND NEW RECORDS OF NORTH AMERICAN PITYOPHTHORUS (COLEOPTERA: SCOLYTIDAE) PART VII. Donald E. Bright! ABSTRACT. —Three species of Pityophthorus from Mexico are described: bravoi (Mexico), conscriptus (Morelos), and ostryacolens (Morelos). Additional host or locality data are given for: atomus, corruptus, deletus, diglyphyus, elimatus, exquisitus, furnissi, hylocuroides, molestus, montezumae, nebulosus, nocturnus, and vespertinus. Existing keys are modified to contain all newly described species. This is the final contribution to a series of papers describing the unnamed species and giving new data for the species of Pityophtho- rus collected by T. H. Atkinson and his col- leagues (Centro de Entomologia y Acarologia, Colegio de Postgraduados, Chapingo, Mex- ico). Unlike the previous papers of this series (Great Basin Nat. 45: 467—482) in which all the new species in a species group were described together and a new key written for the species of the group, each of the species described here is the only new one in a species group. Therefore, only the key couplets from my 1981 monograph (Ent. Soc. Canada Mem. 118) are modified to accommodate the new forms. Once again I wish to thank Dr. T. H. Atkin- son for sending the specimens to me and also thank him and his colleagues for their persi- tence in searching for Scolytidae in numerous different host plants. I also thank my col- leagues Dr. E. C. Becker and Dr. J. M. Campbell for reviewing the manuscript of this paper. Pityophthorus bravoi, n. sp. Length 1.9-2.2 mm, 2.7 times longer than wide. FEMALE.—Frons flattened on a semicircu- lar area extending laterally from eye to eye ‘and from epistoma to well above upper level of eyes; surface shining, with numerous fine punctures and with sparse, erect setae, these more abundant, longer, incurved on periph- ery. Antennal club broadly oval, 1.4 times longer than wide, widest through segments 2 and 3; suture | weakly arcuate, suture 2 more strongly arcuate; segment | slightly narrower than 2; first two segments occupy half of total club length. Pronotum less than 1.1 times longer than wide, widest behind summit; sides weakly arcuate; asperities on anterior slope small, scattered, isolated; summit strongly elevated; posterior area of disc dis- tinctly punctured, punctures small, deep, close, surface between punctures moderately shining, finely, minutely reticulate; median line not elevated, impunctate. Elytra 2.0 times longer than wide; apex broadly rounded; discal striae punctured in regular rows, punctures larger, deeper than those on posterior portion of pronotum; discal inter- striae only slightly wider than striae or as wide as striae, moderately shining, surface finely, minutely reticulate, interstriae 1, 3, 5, 7 each with a median row of sparse punctures and setae, punctures smaller than those in striae, setae longer than those in striae. Declivity convex, weakly bisulcate; interstriae 2 weakly impressed below 1 and 3, equal in width to discal width, surface as on disc; interstriae 1 very weakly impressed below level of 3, with median row of very small granules and setae; interstriae 3 bearing median row of granules, these slightly larger than those in interstriae 1 and with several long, stout setae (sometimes abraded); punctures in striae 1 and 2 much smaller than those on disc, barely visible. MALE.—Frons convex, median longitudi- nal carina weakly elevated from epistoma to upper level of eyes, surface on each side of carina shining, densely, finely punctured, se- tae absent except along epistomal margin. Pronotum and elytra essentially as described 'Biosystematics Research Institute, Agriculture Canada, Neatby Building, Ottawa, Ontario, Canada K1A 0C6. 679 680 for female except pronotal asperities some- what stronger and pronotal and elytral punc- tures larger, deeper. Declivity as in female. TYPE MATERIAL.—The holotype (@) is la- beled: “Carr. Mex. Popo, Km. 85, VIII-26- 1961, Col. H. Bravo M.” /2/ “HOLOTYPE Pityophthorus bravoi D. E. Bright, 1986, CNC No. 18719.” The allotype and 6 paratypes bear the same data. The holotype, allotype, and two paratypes are in the CNC; four paratypes have been returned to T. H. Atkinson. CoMMENTs.—Adults of this species closely resemble those of P. clivus Bright. They differ from those of P. clivus by the longer, more obscure carina on the male frons, by the more evenly pubescent female frons, by the slightly more deeply sulcate elytral declivity on which interstriae 2 is not wider than the discal width, and by the slightly larger size. The key to the species in the Blaudus group (pp. 238-241) in my 1981 monograph should be modified as follows: (Males) 5. Occurs in Mexico; carina on frons moderately or weakly elevated gequ wanes eins ceecPacie «seen 6 — Occurs in western United States and western Canada; carina on frons variable in height 6. Carina on frons distinctly, moderately elevated, short, extending to well below upper margin of eyes; 1.7-2.0 mm; Nuevo Leén...... clivus Bright — Carina on frons weakly elevated, extending to well above upper margin of eyes; 1.9-2.2 mm; NExiCOUL ERs er ee entrant bravoi Bright 6a. continue from couplet 6 in monograph. (Females) 27. Occurs in Mexico; setae on frons sparse; median pair of serrations on anterior pronotal margin longer than others; interstriae 2 equal to or only slightly wider on declivity than on disc ........ 28 — Occurs in western United States and western Canada; setae on frons abundant over entire sur- face; median pair of serrations on anterior prono- tal margin not especially longer than others; in- terstriae 2 distinctly wider on declivity than on Cab oloe tat Bt inrcre nia oie Oe CREE sierraensis Bright 28. Setae on frons absent or very sparse on small area in center; interstriae 2 slightly wider on declivity than on disc; 1.7—2.0 mm; Nuevo Leon — Setae on frons sparse, evenly distributed over surface; interstriae 2 equal in width on declivity and disc; 1.9—-2.2 mm; Mexico ...... bravoi Bright 29. continue as in couplet 28 in monograph. GREAT BASIN NATURALIST Vol. 46, No. 4 Pityophthorus conscriptus, n. sp. Length 1.7-1.9 mm, 2.8 times longer than wide; color black with light reddish legs and antennae. FEMALE.—Frons flattened on a large, semicircular area extending from epistoma to well above upper level of eyes, about half of flattened area above upper level of eyes, ris- ing slightly toward epistoma and with low, median granule on epistoma; surface shining, with fine, dense, scattered punctures and long, erect, yellowish setae, these slightly longer and incurved on periphery, denser along epistoma. Antennal club oval, 1.2—1.3 times longer than wide, widest through seg- ment 2; sutures 1 and 2 transverse, 1 more strongly chitinized; segments 1 and 2 occupy half of total club length. Pronotum 1.1 times longer than wide, widest on posterior half; asperities on anterior slope arranged in a se- ries of irregular concentric rows, the three or four rows behind anterior margin composed of individual, closely placed asperities generally arranged in broken rows, some rows not ex- tending completely from side to side, with occasional isolated asperities between rows, upper rows around summit in form of elon- gate, elevated costae, these sometimes weakly serrate but not divided to base to form individual asperities, each row extending up to half of distance across pronotum, some rows joining adjacent rows; summit not elevated; posterior area of disc distinctly punctured, punctures moderate in size and depth, sepa- rated by distance equal to or less than their diameters; surface between punctures smooth, moderately shining, with numerous impressed points; median line broad, im- punctate, not elevated. Elytra 1.8 times longer than wide; apex broadly rounded; dis- cal striae punctured in regular rows, punc- tures larger, deeper than those on posterior portion of pronotum; interstriae slightly nar- rower than striae, smooth, more brightly shining than posterior portion of pronotum with scattered fine points or lines. Declivity evenly convex, moderately steep; interstriae 1 moderately elevated, devoid of granules; interstriae 2 flat, slightly broader than discal width; interstriae 3 not elevated, unmodified; surface of all interstriae dull, densely micro- punctate; striae 1 narrowly impressed, slightly more strongly so on upper half. Inter- October 1986 striae 1 with a median row of erect setae ex- tending from base to apex; interstriae 3, 5, 7 with median row of similar setae extending from or posterior to midpoint of disc to apex. MALE.—Frons weakly flattened from epis- toma to above upper level of eyes, more dis- tinctly, transversely impressed just above epistoma, slightly convex on upper portions; surface densely, strongly punctured, dull on lower half, shining above, setae short, scat- tered, longer and more abundant in trans- verse impressions above epistoma. Pronotum and elytra essentially as on female except punctures on pronotum and in striae slightly larger. Declivity steep, deeply bisulcate; in- terstriae 1 strongly elevated, broad, with me- dian row of 4 or 5 large, acute granules; inter- striae 2 much broader than discal width, smooth, distinctly impressed; interstriae 3 strongly elevated, arcuate, with median row of 4 or 5 acute granules, these smaller than those on interstriae 1; surface and vestiture of interstriae as in female. TYPE MATERIAL.—The holotype (@) is la- beled: “LA HERRADURA, Mpio, Tepoztlan, Mor.|[elos], 10 Jie 1982, 1,750 m, (??)3-127, A. Burgos—E. Saucedo’/“HOLOTYPE Pity- ophthorus conscriptus D. E. Bright, 1986, CNC No. 18720.” The allotype and six paratypes bear the same labels. Some of the letters or numbers on the labels are illegible. The holotype, allotype, and two paratypes are in the CNC; four paratypes have been returned to T. H. Atkinson. COMMENTS.—Adults of this species are unique by the presence of broken concentric rows of asperities on the lower slope of the pronotum, by the short concentric, anasto- matic rows of elevated costae on the upper slope and around the summit of the prono- tum, by the deeply bisulcate elytral declivity that bears distinct granules on interstriae 1 and 3 of the male, and by the evenly sloping, not sulcate, not granulate elytral declivity of the female. This species does not readily fit into any of the species groups I used in my 1981 mono- graph. Its declivital characters indicate rela- tionship to P. guatemalensis, but its pronotal characters seem distinctly unrelated. Adults of this species differ from those of P. guatemalensis by the more deeply impressed male declivity, by the less deeply impresseda BRIGHT: AMERICAN PITYOPHTHORUS 681 female declivity that is devoid of granules on the first and third interstriae, by the charac- ters on the pronotum as described above, by the more extensively pubescent female frons, and by the less strongly convex male frons. The key to species group (p. 22) in my monograph should be modified as follows: 6. (as in monograph) — Segment 1 of antennal club nearly equal in width to others, club widest through segment 2....... Ta 7a. Asperities on lower slope of pronotum arranged in broken, concentric rows, those on upper slope and around summit developed into short, con- centric, anastomatic costae; male elytral declivity deeply bisulcate, with acute granules on inters- triae 1 and 3; female elytral declivity convex, not sulcate, devoid of granules on interstriae 1 and 3 RIPE ARTES aheAVENCe SIA MES. Hal meer Ana e Conscriptus group — Asperities on anterior slope and around summit arranged in even concentric rows, elevated costae not present; elytral declivity variable, not as above 7. Continue as in monograph. Pityophthorus ostryacolens, n. sp. Length 2.4-2.5 mm, 2.9 times longer than wide; color reddish brown, legs and antenna light reddish or yellowish brown. FEMALE.—Frons weakly convex, with a smooth, broad, weakly elevated, longitudi- nal, median elevation extending from epis- toma to level of upper margin of eyes; surface lateral to and above elevation densely punc- tured, punctures small, shallow, becoming larger, deeper toward vertex and laterally to- ward eyes, surface between punctures shin- ing, generally smooth but with a few, scat- tered minute granules; vestiture moderately abundant, scattered, consisting of downward pointing, moderately long, yellowish setae, these longer, denser along epistomal margin. Antennal club broadly oval, 1.2 times longer than wide, widest through segment 2; sutures 1 and 2 arcuate; segments 1 and 2 occupy about half of total club length. Pronotum 1.1 times longer than wide, widest at middle; sides very weakly arcuate on basal half, broadly rounded to the weakly serrate ante- rior margin; anterior slope with numerous, scattered, low asperities, these extending lat- erally to base and onto posterior discal por- tion; summit not elevated; posterior area of disc deeply punctured, punctures large, sepa- rated by a distance less than their diameters, lateral edges of punctures elevated, forming 682 low asperities except in very limited median area, surface between punctures smooth, shining, with minute impressed points; me- dian line not evident. Elytra 1.8 times longer than wide; apex broadly rounded; discal striae punctured in regular rows, punctures large, deeply impressed, each with a very short seta; interstriae about 1.5 times wider than striae, surface moderately shining, with numerous, scattered, impressed points. Declivity evenly convex, sloping: striae and interstriae essen- tially as on disc except interstriae 1, 3, 5, 7, etc., with a median row of very fine granules and a median row of erect setae and strial punctures slightly less distinct. MALE.—Identical to female except setae on frons less abundant and less distinct. TYPE MATERIAL.—The holotype (2) is la- beled: “Cuernavaca, Mor.|elos], 18.III.82, S- 396, 2,190 msnm, T. H. Atkinson’/“Ostrya virginiana (Ulmaceae)’/“HOLOTYPE Pity- ophthorus ostryacolens D. E. Bright 1986, CNC No. 18721.” The allotype bears the same locality and host label plus an allotype label. The holotype and allotype are in the CNC. COMMENTS.—This species belongs in the Alni group and will key out near P. alnicolens Wood. Adults differ by their larger size, by the more distinct longitudinal carina on the frons, by the more weakly serrate anterior pronotal margin, and by the different host and © distribution. The key to the species in the Alni group (pp. 92-93) in my 1981 monograph should be mod- ified as follows: 3. Declivital interstriae 2 bearing a median row of stout setae; elytral striae impressed on disc, inter- striae convex; lateral areas of pronotum asperate almost to base; pronotum and elytra shining; Weracnuzir accor ore oe ee oc alni Blackman — Declivital interstriae 2 not bearing a median row of setae; elytral striae not or only weakly im- pressed; lateral areas of pronotum punctate to subasperate; pronotum and elytra dull to moder- atelyshiningea) see cise ee eae Sess 3a 3a. Length 1.8—2.0 mm; longitudinal carina on frons indistinct, faintly elevated; elytral striae not im- pressed; in Alnus, Veracruz ..... alnicolens Wood — Length 2.4-2.5 mm; longitudinal carina on frons distinct, weakly elevated; elytral striae weakly impressed; in Ostrya, Morelos 4. Continue as in monograph. GREAT BASIN NATURALIST Vol. 46, No. 4 NEw Host or LOCALITY RECORDS Only locality records that significantly ex- tend the range or represent the first records since the species was described are listed be- low. All new host records seen are also listed below. Numerous new state records were seen in the material examined; these will be reported later. Pityophthorus atomus Wood Pityophthorus atomus Wood, 1964, Great Basin Nat. 24: 61; Bright, 1981, Ent. Soc. Canada Mem. 118: 44: Wood, 1982, Great Basin Nat. Mem. 6: 1137. This species was previously recorded only from Oaxaca and Veracruz from an unknown shrub. Two series totaling 17 specimens have been seen with the data: “Pachuca, Edo. de Hidalgo, S-463, 21.V.82, 2,400 m, Col. A. Equihua M.” (8) and “Jalapa, VERACRUZ, 25.11.84, FANM 143, Col. Felipe A. Noguera /“Hosp. Vernonia sp. (Compositae). ” Pityophthorus corruptus Wood Pityophthorus corruptus Wood, 1976, Great Basin Nat. 36: 363; Bright, 1981, Ent. Soc. Canada Mem. 118: 68; Wood, 1982, Great Basin Nat. Mem. 6: 1133. Known only from the type locality in Puebla from Toxicodendron (or Rhus sp.). Twenty- nine specimens have been seen with the data: “San Rafael, Mex.[ico], 4.1X.81, S-242, 2,400 m, Atkinson-Equihua’/“Hosp.: Rhus sp.” Pityophthorus deletus LeConte Pityophthorus deletus LeConte, 1879, Bull. U.S. Geol. Geog. Survey 5(3): 519; Bright, 1981, Ent. Soc. Canada Mem. 118: 110; Wood, 1982, Great Basin Nat. Mem. 6: 1040. This name includes a presently unresolved complex of one or more very closely related and variable species. Seven names (sensu Bright) or six names (sensu Wood) are in- cluded as synonyms under the above name. This species is known from California to South Dakota, south to Durango and Coahuila, Mexico. Five specimens which | have assigned to this species complex have been seen with the labels: (3) “Cerro Tlaloc, Tequesquinahuac, Edo. Mex.[ico], 22.1.80; 3,400 msnm, Col. T. H. Atkinson’/“Hosp.: Pinus hartwegii (Pinaceae) and (2) “Zoquia- pan, Edo. Mex.|ico], Altitud: 3,600 msnm, Fecha: 25-IJI-81, Col. A. Equihua M.’7/ “Hosp. Pinus hartwegii (Pinaceae). ” October 1986 Pityophthorus diglyphus Blandford Pityophthorus diglyphus Blandford 1904, Biol. Cent.- Amer. 4(6): 240; Bright, 1981, Ent. Soc. Canada Mem. 118: 121; Wood, 1982, Great Basin Nat. Mem. 6: 1058. Previously known only from Guatemala. Three specimens, all females, were seen with the labels: “Volcan Chichinautzin, Morelos, 30. Jie (?), 1982, 2,790 m, 73-133, J. Burgos”/ “(Pinaceae) Pinus sp.” These three specimens could possibly be considered examples of P. leiophyllae Black- man based on morphological similarity and locality. The presence of long setae on the third declivital interstriae place it in P. digly- phus as presently understood. The two names may be synonyms, but more specimens are needed before a definite conclusion can be made. Pityophthorus elimatus Bright Pityophthorus elimatus Bright, 1976, Great Basin Nat. 36: 432; Bright, 1981, Ent. Soc. Canada Mem. 118: 196; Wood, 1982, Great Basin Nat. Mem. 6: 1086. This species was previously known only from the type locality in Oaxaca. Eight speci- mens have been seen with the data: “Parque Nal. Zoquiapan, Edo. de Mexico, 3,200 msn, 24.V.80, S-099, Col. D. Cibrian To- var. Pityophthorus exquisitus (Blackman) Neodryocoetes exquisitus Blackman, 1942, Proc. U.S. Nat. Mus. 92(3147): 196. Pityophthorus exquisitus : Bright, 1981, Ent. Soc. Canada Mem. 118: 102; Wood, 1982, Great Basin Nat. Mem. 6: 1026. Pityophthorus inceptis Wood, 1975, Great Basin Nat. 35: 396; Bright, 1981, Ent. Soc. Canada Mem. 118: 102. This species was previously known from Jalisco and Michoacan. The host is listed as an unknown shrub or in wood crates or wood stems. Four specimens were examined that bore the following labels: “Rancho Tetela, Cuernavaca, Morelos], Compositae, 10-En- ero-1982, 1,350 m, Col. BUSA-SACE- MAFE.” Pityophthorus furnissi Bright Pityophthorus furnissi Bright, 1976, Great Basin Nat. 36: 433; Bright, 1981, Ent. Soc. Canada Mem. 118: 144; Wood. 1982, Great Basin Nat. 6: 1094. BRIGHT: AMERICAN PITYOPHTHORUS 683 Previously known only from the type local- ity of Amecameca, Mexico. Eight specimens were seen with the following data: “Cerro Tlaloc, Tequesquinahuac, Edo. Mex{ico], 22.1.80, 3,400 msnm, Col. T. H. Atkinson’/ “Hosp.: Pinus hartwegii (Pinaceae).” Pityophthorus hylocuroides Wood Pityophthorus hylocuroides. Wood, 1964, Great Basin Nat. 24: 69; Bright, 1981, Ent. Soc. Canada Mem. 118: 31; Wood, 1982, Great Basin Nat. Mem. 6: 1120. Previously known only from Hidalgo taken on Rhus sp. One specimen has been seen with the data: “Jalapa, VERACRUZ, 21.11.84, FANM 140, Col. Felipe A. Noguera ’/“Hosp.: Tithonia sp. (Compositae). ” Pityophthorus molestus Wood Pityophthorus molestus Wood, 1976, Great Basin Nat. 36: 362; Bright, 1981, Ent. Soc. Canada Mem. 118: 61; Wood, 1982, Great Basin Nat. Mem. 6: 1131. Previously known only from the type local- ity in San Luis Potosi. Three specimens were seen with the data: “Apulco Centre Zaca- poaxtla y Cuetzalan, Pue[bla], 4.V.81, 1,480 m, Col. T. H. Atkinson y A. Equihua, $2167/ “Hosp.: Liquidambar styraciflua.” Pityophthorus montezumae Bright Pityophthorus montezumae Bright, 1978, Great Basin Nat. 38: 81; Bright, 1981, Ent. Soc. Canada Mem. 118: 272: Wood, 1982, Great Basin Nat. Mem. 6: 1079. Known previously only from the type local- ity in Chiapas. Two specimens have been seen with the data: “Parque Nal. Zoquiapan, Edo. de Mexico, 3,200 msnm, 24.V.80, S-099, Col. D. Cibrian Tovar” (1) and “Parque Nal. Zo- quiapan, Edo. Mexico. Agosto 79, Hos. Pinus hartwegii. T. H. Atkinson’ (1). Pityophthorus nebulosus Wood Pityophthorus nebulosus Wood, 1976, Great Basin Nat. 36: 363; Bright, 1981, Ent. Soc. Canada Mem. 118: 100; Wood, 1982, Great Basin Nat. Mem. 6: 1126. This species was previously known only from the type series collected at Lake Catemaco, Veracruz from Bursera sp. A se- ries of 10 specimens was seen bearing the labels: “Campo Exptal. I.N.I.F., Escarcega, CAMP[ECHE], 14.1X.83, AEV. 51, Col. A. Estrada V.’/“Bursera semaruba (Burseraceae).” 684 Pityophthorus nocturnus Sched| Pityophthorus nocturnus Schedl, 1938, Archiv Naturgesch. 7: 185; Bright, 1981, Ent. Soc. Canada Mem. 118: 192; Wood, 1982, Great Basin Nat. Mem. 6: 1087. Pityophthorus hidalgoensis Blackman, 1942, Proc. U.S. Nat. Mus. 92: 215. This species was previously known from the states of Chiapas, Hidalgo, and Veracruz in Mexico and from Guatemala and Honduras. Three series have been seen with the lables: “Taxco, Guerrero, 22-11-82, S-325, 1,900 m, Col . Atkinson y Equihua’/“Hosp. Pinus sp.’; “La Herradura, Cuernavaca, Mor.|elos], 10 Diciembre 1982, 1,810 m, SM-104, E. GREAT BASIN NATURALIST Vol. 46, No. 4 Saucedo-A. Burgos’/“(Pinaceae, Pinus sp.” and “Acajete, Ver.[acruz]), 22-XI-83, FAMN 91, Col. Felipe A. Noguera’/“Hosp. Pinus patula (Pinaceae).” Pityophthorus vespertinus Bright Pityophthorus vespertinus Bright, 1978, Great Basin Nat. 38: 83; Bright, 1981, Ent. Soc. Canada Mem. 118: 119; Wood, 1982, Great Basin Nat. Mem. 6: 1058. This species was previously known from only four specimens collected from Pinus sp. in Durango. Six specimens that are referred to this species have been seen bearing the labels: “Acajete, Ver[acruz], 22-XI-83, FANM 91, Col. Felipe A. Noguera’/“Hosp. Pinus patula (Pinaceae). ” EFFECTS OF DWARF MISTLETOE ON SPRUCE IN THE WHITE MOUNTAINS, ARIZONA Robert L. Mathiasen', Frank G. Hawksworth?, and Carleton B. Edminster” ABSTRACT.—Mortality of spruce in mixed conifer stands moderately to heavily infested with western spruce dwarf mistletoe was two to five times greater than in healthy stands in the White Mountains, Arizona. Ten-year volume growth loss for heavily infected spruce trees ranged from 25% to 40%. Estimates of growth loss for spruce on a stand basis ranged from 10% to 20% in heavily infested stands. Because western spruce dwarf mistletoe is prevalent in the White Mountains and causes increased mortality and reduced growth, its control should be included in management of mixed conifer stands there. Western spruce dwarf mistletoe (Arceutho- bium microcarpum |Engelm.| Hawksw. & Wiens) is a damaging parasite of Engelmann spruce (Picea engelmannii Parry), blue spruce (P. pungens Engelm.), and bristlecone pine (Pinus aristata Engelm.) in the southwestern United States (Hawksworth and Wiens 1972, Mathiasen and Hawksworth 1980). The distri- bution of western spruce dwarf mistletoe is confined to Arizona (Pinaleno and White Mountains, San Francisco Peaks, Kendrick Peak, and the North Rim of Grand Canyon) and New Mexico (Mogollon and Sacramento Mountains) (Hawksworth and Wiens 1972, Mathiasen and Jones 1983). Western spruce dwarf mistletoe is most prevalent in the White Mountains, Arizona (Apache-Sitgreaves Na- tional Forest), where it has been reported to be in over 60% of the spruce type and is a primary factor associated with spruce mortal- ity (Hawksworth and Graham 1963). It is more common in the lower mixed conifer forests than in spruce-fir forests, possibly because its distribution is restricted to below approxi- mately 10,400 feet (Acciavatti and Weiss 1974, Mathiasen and Hawksworth 1980). Gottfried and Embry (1977) reported blue spruce was more heavily infected than Engel- mann spruce in a virgin mixed conifer stand in the White Mountains, Arizona, but overall infection of both species was relatively low (2% and 5% for Engelmann and blue spruce, respectively). However, Gottfried and Em- bry also reported that almost 20% of their sample points containing blue spruce had in- ISchool of F orestry, Northern Arizona University, Flagstaff, Arizona 86011. fected trees. Hawksworth and Graham (1963) reported 63%—70% of the spruce stands they surveyed in the White Mountains, Arizona, were infested with western spruce dwarf mistletoe. Our observations in the White Mountains also indicate blue spruce is heavily infected in many mixed conifer stands, partic- ularly along drainages. Jones (1974), Gottfried and Embry (1977), and Ronco et al. (1984) proposed general recommendations for silvi- cultural management of mixed conifer forests that consider the dwarf mistletoe problem. Although western spruce dwarf mistletoe represents the most damaging disease agent in southwestern mixed conifer forests domi- nated by Engelmann or blue spruce, little information is available regarding its effect on mortality and growth of its principal hosts. This study provides additional quantitative data on the mortality and growth loss caused by western spruce dwarf mistletoe in the White Mountains, Arizona. METHODS In 1981, 99 temporary rectangular plots ranging in size from 0.1 to 0.8 acre were placed in mixed conifer stands with various densities of Engelmann and/or blue spruce in the White Mountains, Arizona (Apache-Sit- greaves National Forest). Two-thirds of the plots were infested with various levels of west- ern spruce dwarf mistletoe. Plots were se- lected in an attempt to maintain a homoge- neous distribution of age classes, species 2USDA Forest Service, Rocky Mountain Forest and Range Experiment Station, Fort Collins, Colorado 80526. 685 686 composition, and dwarf mistletoe infection. Plots were only located in stands that had not been disturbed for at least 12 years prior to data collection. The following data were recorded for each tree greater than 4.5 feet in height in each plot: species, diameter breast height (DBH) to the nearest 0.1 inch, dwarf mistletoe rating (six-class system, Hawksworth 1977)’, crown class (dominant, co-dominant, intermediate, suppressed), tree condition (live or dead), and ten-year radial wood growth at DBH to the nearest 0.05 inch. Overstory DMR values were calculated using DMR for spruce trees only. Height and age data were measured as fol- lows: total height to the nearest 1.0 foot for two to three living or dead trees from each one-inch diameter class represented for each species in a plot; height to the base of the live crown to the nearest 1.0 foot for-live trees measured for total height; distance from the ground to the tenth whorl from the top of live trees measured for total height; age at DBH for two live trees from each two-inch diameter class represented for each species in a plot. Height versus diameter curves were devel- oped for each plot, and these curves were used to estimate present and past heights for trees with only measured diameters. Present volumes and past ten-year volumes were cal- culated from diameter and height data for En- gelmann and blue spruce using the volume equations for Engelmann spruce developed by Hann and Bare (1978). Ten-year periodic annual volume increment (cubic feet/year) was calculated using present and past volumes for spruces greater than 6.0 inches at DBH. RESULTS THE SAMPLE. —Of the 99 plots sampled, 34 were predominantly Engelmann spruce, 42 were predominantly blue spruce, and 23 had approximately equal representation of Engel- mann and blue spruce based on stems/acre. Basal areas (square feet/acre) ranged from 10 “The six-class dwarf mistletoe rating system divides the live crown ofa tree into thirds, and each third is rated separately as: 0, no infected live branches; 1, less than 50% of the live branches infected; 2, more than 50% of the live branches infected. The ratings for each third are totaled to obtain a dwarf mistletoe rating (DMR) for the tree. Adding the DMRs for all live trees in a stand and dividing the total by the number of trees is the average stand dwarf mistletoe rating (DMR). The six-class system is a standard method for quantifying the intensity of mistletoe infection for individual trees or stands. GREAT BASIN NATURALIST Vol. 46, No. 4 to 134 for blue spruce and from 6 to 156 for Engelmann spruce. However, approximately one-third of the plots sampled had Engel- mann or blue spruce basal areas of less than 50 square feet/acre: Number of plots Basal area Engelmann Blue (square feet/acre) spruce spruce <100 ) Wy The majority of the plots sampled had spruce densities (stems/acre) of less than 200: Number of plots Engelmann Blue Stems/acre spruce spruce <100 Ia 19 101—200 13 21 201-300 i) ) >300 q 2 One-third of the plots had no western spruce dwarf mistletoe infection and the re- mainder ranged from light to heavy: DMR Number of plots 0 33 0.1-1.0 i 1.1-2.0 20 2.1-3.0 10 >3.0 9 MorrTALiTy.—Percent mortality was deter- mined for the following size classes: small saplings (DBH 0.1-—1.0 inches), large saplings (DBH 1.1-5.0 inches), poles (DBH 5.1—10.0 inches), small sawtimber (DBH 10.1—16.0), and large sawtimber (DBH greater than 16.0 inches) by 1.0 DM R classes (Table 1). Mor- tality of small and large saplings was approxi- mately the same for healthy plots (DM R 0) and for plots with a DMR less than 2.0. However, percent mortality for these size classes was approximately two and four times greater than in healthy plots for DM R classes 2.1-3.0 and greater than 3.0, respectively. Mortality of pole size spruce increased rapidly as DMR increased. Mortality of pole-sized Engelmann spruce was higher than for blue spruce in the higher DMR classes (DM R greater than 2.0). This trend was reversed in the small and large sawtimber size classes, October 1986 MATHIASEN ET AL.: ARIZONA DWARF MISTLETOE 687 TABLE 1. Percent mortality of Engelmann and blue spruce by 1.0 DMR classes and size classes. Size class (Inches) Small Large Small Large sapling sapling Poles sawtimber sawtimber OME (0.1-1.0) (1.1-5.0) (5.1-10.0) (10. 116.0) (>16.0) All trees Class E? Be E B E B E B E B E B 0 BY 4 u 9 7 4 9 BY 6 4 6 6 0.1-1.0 0 1 5 6 17 10 11 3 % 2 0 8 4 1.1-2.0 2 3 U 6 18 18 12 23 10 10 10 14 2.1-3.0 10 9 12 16 20 15 17 OT si 20 16 17 >3.0 20 17 32 25 34 24 20 33 19 33 23 28 lEngelmann spruce Blue spruce TABLE 2. Percentage of dead trees with DMR 2-6 by size class.' Dwarf Mistletoe Rating’ Size class Total (Inches) trees 2 3 4 5 6 (Percent) Small sapling 20 5 10 20 30 35 (0.1-1.0) Large sapling 63 8 11 14 Th 40 (1.1-5.0) Poles 29 0 0 14 28 58 (5.1-10.0) Small sawtimber 33 0 0 6 36 58 (10. 116.0) Large sawtimber 14 0 0 14 28 58 (>16.0) Total 159 4 6 13 30 47 Includes both Engelmann and blue spruce ?Includes only dead trees that could be assigned an accurate DMR TABLE 3. Mean ten-year periodic annual volume increment for Engelmann and blue spruce greater than 6.0 inches DBH by DMR class. DMR Mean ten-year periodic annual Percent difference class N volume increment (cubic feet/year) from DMRO 0 620 0.32 A’ — 1 48 0.34 A +6 2 94 0.31 A —3 3 127 0.30A 6 4 100 0.28 B —12 5 128 0.24 C —25 6 107 0.20 D —38 Numbers followed by different letters are significantly different. Oneway AOV, a = 0.05, Student-Newman-Kuels. where blue spruce had a more rapid increase in mortality as DM R increased. Mortality of small sawtimber was from two to five times greater in the most heavily infested plots than in healthy plots. Mortality of large sawtimber was from three to eight times greater in the most heavily infested plots (Table 1). Nearly half of the dead spruce that could be accurately assigned a DMR were rated as class 6 trees (Table 2). This was true for all size classes except the small sapling class where approximately one-third of the trees were rated as class 5 or 6. The percentage of dead trees rated as class 4 ranged from 6% for the small sawtimber size class to 20% for the small saplings. Few dead spruce were rated as class 2 or 3 in the sapling size classes, and none were rated 2 or 3 in the pole and sawtimber size classes. EFFECT ON VOLUME GROWTH.—Mean ten- 688 GREAT BASIN NATURALIST Vol. 46, No. 4 TABLE 4. Percent infection of live trees by DMR and 0.5 DMR classes’. Dwarf mistletoe rating DMR 0 1 2 class 0.1-0.5 82 13 4 0.6-1.0 50 32 6 Ts 40 25 14 1.6—2.0 23 32 17 OMEOES 16 26 11 2.6-3.0 8 19 19 Sul= 35 6 19 12 3.6—4.0 2 a 12 4.0+ 1 6 10 ‘ll spruce combined year periodic annual volume increment (cubic feet/year) was determined for all spruce greater than 6.0 inches (DBH) for DMR classes 0-6 (Table 3). The results do not in- clude growth loss due to mortality of individ- ual trees. INFECTION AND GROWTH Loss ON A STAND Basis.—The percentage of live trees infected for both Engelmann and blue spruce by indi- vidual tree DMR and by 0.5 DMR classes are presented in Table 4. The percentage of trees in the heaviest infection class (DMR 6) ranged from 0% in DMR class 0.1-0.5 to 23% in DM R class 4.0+. A summary of the percentage of trees in DMR classes 4—6 (those in which significant growth loss occurs) is as follows: DMR Percentage of trees in class DMR class 4-6 0.1-0.5 0 0.6-1.0 4 1.1-1.5 12 1.6—2.0 cs} 2.1-2.5 28 2.6-3.0 35 3.1-3.5 4l 3.6—4.0 57 4.0 + @ Stand growth loss was estimated by the dis- tribution of infected spruce by DM R classes (Table 4), and based on the following esti- mates of growth loss for individual trees by DMR class (Page 6): DMR 1—0%, DMR 2— 0%, DMR 3—5%, DMR 4— 10%, DMR 5— 25%, DMR 6—40%. Stand growth loss by 1.0 DM RB classes based on the above estimates was: 3 4 5 6 (Percent) 1 0 0 0 8 2 ] 1 9 5 4 3 15 6 4 3 19 10 ll 7 19 9 12 14 22 19 14 8 22, 21 20 16 ll 20 29 23 DMR Estimated percent loss class on a stand basis 0.1-1.0 i We ee (0) 3 230) 9 3.1-—4.0 1; ADE 20 DISCUSSION Mortality of Engelmann and blue spruce in mixed conifer stands moderately to heavily infested with western spruce dwarf mistletoe is from two to five times greater than for healthy stands in the White Mountains, Ari- zona. Approximately 20% to 35% of the spruce sampled in heavily infested stands were dead, indicating western spruce dwarf mistletoe is a primary factor associated with spruce mortality. Hawksworth and Graham (1963) also reported high mortality rates for spruce in western spruce dwarf mistle- toe—infested mixed conifer stands in the White Mountains. Nearly half of the dead trees that could be accurately assigned dwarf mistletoe ratings were class 6 trees. Approximately one-tenth and one-third of these dead spruce were in DMR class 4 and 5, respectively. This was true for all size classes of spruce except the small sapling size class, where more dead saplings were rated as class 4. The high mor- tality rate in class 5 trees for spruce contrasts to mortality patterns in dwarf mistletoe-in- fected pines, where mortality is predomi- nantly in class 6 trees (Hawksworth and Lusher 1956). Heavy dwarf mistletoe infection (DMR 5-6) severely reduces volume increment of spruce October 1986 in the White Mountains. Lightly infected spruce (DMR 1-2) do not suffer any detectable growth loss and moderately infected spruce (DMR 3-4) only suffer losses ranging from approximately 5% to 10%. These results are similar to those reported for southwestern dwarf mistletoe (Arceuthobium vaginatum subsp. cryptopodum [Engelm.] Hawksw. & Wiens) parasitizing ponderosa pine (Pinus ponderosa Laws.) in the Southwest (Hawks- worth 1961). Estimates of growth losses on a stand basis ranged from approximately 10% to 20% for heavily infested stands (DMR greater than 2.0). Lightly to moderately infested stands are estimated to have losses less than 3%. Our estimates of the effects of western spruce dwarf mistletoe on the growth of spruce are the first reported for this parasite-host combi- nation. Western spruce dwarf mistletoe is a com- mon parasite of spruce in mixed conifer stands in the White Mountains, Arizona. Because heavy infection by western spruce dwarf mistletoe severely reduces the growth of trees and stands and is associated with increased spruce mortality, silvicultural control of the parasite should be a primary concern of re- source managers. Heavily infected spruce should be removed from infested stands whenever possible to reduce the impact of this parasite on forest productivity. ACKNOWLEDGMENTS The authors acknowledge the assistance of Mr. David Conklin with the field data collec- tion. This study was conducted as part of a cooperative research project between the USDA Forest Service, Rocky Mountain Forest and Range Experiment Station, Fort MATHIASEN ET AL.: ARIZONA DWARF MISTLETOE 689 Collins, Colorado, and the Department of Plant Pathology, University of Arizona, Tuc- son, Arizona. LITERATURE CITED AccIAvaTTI, R. E., AND M. J. WEIss. 1974. Evaluation of dwarf mistletoe on Engelmann spruce, Fort Apache Indian Reservation, Arizona. Plant Dis. Rep. 58: 418-419. — GOTTFRIED, G. J., AND R. S. Emsry. 1977. Distribution of Douglas-fir and ponderosa pine dwarf mistletoes in a virgin Arizona mixed conifer stand. USDA Forest Service, Res. Pap. RM-192. 16 pp. HANN, D. W., AND B. B. BARE. 1978. Comprehensive tree volume equations for major species of New Mexico and Arizona: I. Results and methodology. USDA Forest Service, Res. Pap. INT-209. 43 pp. Hawkswokrty, F. G. 1961. Dwarfmistletoe of ponderosa pine in the Southwest. USDA Forest Service, Tech. Bull. 1246. 112 pp. ____. 1977. The 6-class dwarf mistletoe rating system. USDA Forest Service, Gen. Tech. Rep. RM-48. 7 pp. Hawkswokrty, F. G., AND D. P. GraHaM. 1963. Dwarf- mistletoes on spruce in the Western United States. Northwest Sci. 37: 31-38. HAwkswokrtH, F. G., AND A. A. LUSHER. 1956. Dwarf mistletoe survey and control on the Mescalero Apache Reservation, New Mexico. J. For. 54: 384-390. HaAwkswokrty, F. G., AND D. WIENS. 1972. Biology and classification of dwarf mistletoes (Arceuthobium). USDA Forest Service, Agriculture Handbook 401. 234 pp- Jones, J. R. 1974. Silviculture of southwestern mixed conifer and aspen: the status of our knowledge. USDA Forest Service, Res. Pap. RM-122. 44 pp. MATHIASEN, R. L., AND K. H. Jones. 1983. Range exten- sions for two dwarf mistletoes (Arceuthobium spp.) in the Southwest. Great Basin Nat. 43: 741- 746. MarTHIASEN, R. L., AND F. G. HAwKsworrTu. 1980. Taxon- omy and effects of dwarf mistletoe on bristlecone pine on the San Francisco Peaks, Arizona. USDA Forest Service, Res. Pap. RM-224. 10 pp. RONCO, F., G. J. GOTTFRIED, AND R. R. ALEXANDER. 1984. Silviculture of mixed conifer forests in the South- west. USDA Forest Service, RM-TT-6. 72 pp. HYDROLOGY OF BEAR LAKE BASIN AND ITS IMPACT ON THE TROPHIC STATE OF BEAR LAKE, UTAH-IDAHO Vincent Lamarra’, Chuck Liff’, and John Carter’ ABSTRACT.—Bear Lake is a large, relatively pristine lake located in a graben valley. The lacustrine environment is more than 35,000 years old. Over that period of time, the Bear River intermittently flowed into Bear Lake. Approximately 10,000 to 8,000 BP, the Bear River ceased flowing directly into the lake. Between 1912 and 1924, channels were dug that diverted Bear River flows into the lake. An analysis was conducted to determine the impacts of Bear River flows upon the hydrologic and nutrient budgets of the Bear Lake ecosystem. In addition, the resulting limnological conditions were evaluated. Based upon eight years of historical data (1976 to 1984), regression relation- ships were developed that allowed an estimation of the historical conditions in Bear Lake (1923 to present) with and without the influence of the Bear River. Bear Lake, located on the northern border of Utah, is a 282 km’ natural body of water. The lake presently occupies most of the south- ern half of the Bear Lake Valley. The geology of the area was first mapped by Richardson (1913) and Mansfield (1927) and more recently by Armstrong and Cressman (1963), McClurg (1970), and Kaliser (1972). The Bear Lake basin is a graben valley bordered on the east and west by normal faults, with Mesozoic and Cenozoic rocks on the east and Paleozoic, Mesozoic, and Cenozoic rocks on the west. The exact age of the lake is presently un- known; however, stratigraphic studies by Robertson (1978) have verified previous mor- phologic interpretations by Mansfield (1927), which suggest a glacial age for the origin of Bear Lake. Robertson (1978) suggested the lake has had a continuous lacustrine history of at least 35,000 years (BP). However, unlike pluvial lakes Thatcher and Bonneville, which formed in closed basins and were therefore regulated by climatic fluctuations, the early conditions within the Bear Lake Valley re- mained opened with a northward drainage along the Bear River. Over the last 28,000 years, the major water level fluctuations in the Bear Lake Valley have been the result of downcuttings of the north- ern valley outlet and two periods of faulting within the southern Bear Lake Valley. Early conditions within the lake indicated a wide- spread bay and marsh ecosystem. Tectonic activity lowered the valley differently, result- ing in marshes and shallow bays occupying the northern Bear Lake Valley and a deep lake to the south (Robertson 1978). Current condi- tions have continued over the last 8,000 years with the present outlet of the Bear River northward along the east side of the valley. During most of this time, the lake has been isolated from the major drainage networks (primarily the Bear River). This has led to the occurrence of four endemic fish species that still inhabit the lake in large numbers, and a unique macrochemistry with magnesium as the predominant cation (Kemmerer et al. 1923). At the present time, Bear Lake is no longer considered a closed basin. Its isolations ended in 1912 with the development of Stewart Dam, Lifton Station, and the diversion of the Bear River into Dingle Marsh. Although his- torically the marsh (and therefore the Bear River) was separated from the lake by a natu- rally occurring sandbar, it now serves as a water storage and transfer facility. Water is diverted from the Bear River into the marsh during spring runoff (March—July) and al- lowed to flow into Bear Lake. When irrigation demands increase during the summer, water flows into the marsh from Bear Lake and then into the Bear River downstream from the di- version dam. Previous studies (Nunan 1972) have indi- cated that the storage of Bear River water has altered the macrochemistry of Bear Lake, re- ducing the TDS (total dissolved solids) levels ‘Ecosystem Research Institute, 975 South State Highway, Logan, Utah 84321. 690 October 1986 LAMARRA ET AL.: BEAR LAKE BASIN HYDROLOGY OUTLET RAINBOW oe ae ANAL CANAL ~ ——~ PARIS \ he ; } “= s fe Gu eS \MERKLEY Ven Pepe esis MTNS. \ = = \ \ BIOOMINGTON ; ‘oO } Nan aS Dosa oc _ LIFTON \ ty / HOT MeN ee Oe \ STATION | 4 SPRINGS ¢ ST.CHARLES ( jae N / ye! INDIAN ae ( CREEK ane ! Sa wl TNR rR i TNT “N) FISH HAVEN / \ COOLEY ) a0 CANYON ( j a | es = ; if TKS La. SWAN ( NORTH i inant Ce _? EDEN 2. = UTAH ‘ Fe = Neo \ \ ? * Me i a ARDEN Be SOUT iy pS Y EDEN oa ( (a4 So Aa ‘_--- - (Ge PICKLEVILLE Uf x ee - De : in . CANYON (ROUND \\ VALLEY \ SY = eae \ \ a ‘ & \ A ‘ \ X ; / ( 7) N MR & ( \ ({LAKETOWN ) 4 — ( A aed = — rtf yf o | oe (iy Fig. 1. Watershed and bathemetric map of Bear Lake, Utah-Idaho. Depth contours are in meters. 691 692 from over 1000 mg/l to 500 mg/l during a 50-year period. Recently Lamarra et al. (1984) has noted that the Bear River has also altered the trophic state of Bear Lake. Because of the dominant role of the Bear River in the histori- cal as well as the current hydrological and limnological conditions of Bear Lake, a more detailed description of the impacts upon the lake is needed. It is therefore the purpose of this study to quantitatively describe the cur- rent impact of the Bear River upon the hydrol- ogy and trophic state of Bear Lake over the last decade and to empirically describe the historical conditions that have existed within the lake during the last 60 years. METHODS To distinguish the water quality impacts of the Bear River from the historical Bear Lake watershed, mass nutrient loadings were de- termined for each major tributary basin and the Bear River (Figs. 1, 2). Water quality analyses (total phosphorus, total nitrogen, to- tal organic carbon, ortho-phosphate, ammo- nia, nitrate, and nitrite) were performed ac- cording to standard methods (APHA 1980). Flow measurements were determined on site or obtained from Utah Power and Light Com- pany. Water samples were also collected at eight depths at a limnetic site corresponding to the deepest area in Bear Lake (63 m). Analyses included temperature, oxygen, pH, and con- ductivity in addition to the nutrients previ- ously mentioned. Algal biomass (chlorophyll a) was determined by the Fluorometric pro- cedure using a Turner Model III Fluorome- ter. Meteorological data were obtained from the NOAA Station at Lifton. Water quality of rain events was previously determined (Her- ron et al. 1984) and used in this study. RESULTS The Bear Lake ecosystem and its associated watersheds cover approximately 8,250 km’, with 7,000 km’ in the upper Bear River basin and the remaining 1,250 km’ within the natu- ral Bear Lake drainage. These major water- sheds are within the states of Idaho, Utah, and Wyoming (Fig. 2) and lie within the Great Basin. The results of this study will be pre- GREAT BASIN NATURALIST Vol. 46, No. 4 ST. CHARLESO™ WYOMING ee Fig. 2. A location map for Bear Lake and its watershed. sented separately by drainage basin and Bear Lake limnology. BEAR RIVER DRAINAGE.—A special portion of the Bear Lake watershed is located above the Dingle Marsh system. This watershed has only been impacting Bear Lake since 1912, when the Stewart Dam and the associated canal system was constructed. The water from the upper Bear River basin is diverted into Bear Lake during spring runoff (March— June). The annual flows at Stewart Dam for 1975 to 1984 can be seen in Figure 3. The dominant portion of the flows occurs between October 1986 LAMARRA ETAL.: BEAR LAKE BASIN HYDROLOGY 693 Bear River Flow at Stewart Dam 225 200 175 150 125 100 Flow (1000 ac—ft) 75 50 25 1975 1976 1977 1978 1979 1980 1981 1982 19835 Time (years) Fig. 3. The flow in the Bear River at Stewart Dam between 1975 and 1984. March and June. In the last 10 years, the lowest volume entering the marsh was during 1977 (11,400 ac-ft) and 1981 (37,240 ac-ft). The highest volumes occurred in 1983 (411,330 ac-ft) and 1984 (492,400 ac-ft). Because of the water control structures present around the marsh, not all the water entering the system exits into Bear Lake during the spring period. For example, although 492,400 ac-ft entered Stewart Dam in 1984, 240,900 ac-ft was di- verted through the outlet canal back into the Bear River, whereas the remaining 244, 100 ac-ft was released through Lifton Station into Bear Lake (Table 1). The comparison of Lifton inflows to all other water sources into Bear Lake can be seen in Table 2 for the period 1975 to 1984. Between 1975 and 1984 more than 66 sets of water quality data have been collected at the three key water control structures in Dingle Marsh (March to September). These data are also summarized in Table 1. The most com- plete data set are represented by total phos- phorus and total inorganic nitrogen. The total phosphorus data at Lifton represents the mass loading by the Bear River into Bear Lake for 1975 and 1978 to 1984. The difference in the range of loading is significant, with 1981 being only 1,300 kg TP but 1983 being 25,838 kg TP. Ortho-phosphate was from 6.8% to 19.8% of the total phosphorus loading. In all cases, the marsh tended to remove phosphorus from the Bear River prior to its entrance into the lake. This was markedly different when compared to nitrogen and organic carbon (i.e., 1981, 1982, and 1984), when the marsh was in bal- ance or actually increased the mass of these materials to the Bear River as it moved through the system. BEAR LAKE WATERSHEDS AND PRECIPITA- TION.—Land in the Bear Lake watersheds has 694 GREAT BASIN NATURALIST TABLE 1. The mass movements of water and nutrients through key control structures around the Dingle Marsh system. The material moving through Lifton Station is the mass actually entering Bear Lake. (TP = total phospho- rous; TOC = total organic carbon; TN = total nitrogen; TSS = total suspended solids; PO,—P = orthophosphate; TIN = total inorganic nitrogen). Flow (AC-ftx1000) (March—June) Year STD LFT Outlet 1975 119.61 179.34 20.27 1976 253.96 221.08 52.45 1977 11.43 977 111.11 1978 185.26 183.61 1.65 1979 90.36 86.44 22.28 1980 329.73 279.58 o2el3 1981 37.24 34.03 28.79 1982 267.68 264.84 2.83 1983 411.33 251.50 198.97 1984 492.40 244.10 240.90 Kg TP (March—June) Year STD LAP Outlet 1975 11,367 9,982 10 1978 17,366 13,343 0 1979 20,669 8,551 IL 1980 56,310 27,679 10 1981 6,068 1,300 3,480 1982 59,151 18,460 254 1983 75,951 25,838 26,403 1984 115,450 23,680 51,990 Kg TOCx1000 (March—June) Year STD LFT Outlet 1975 — — — 1978 — — = 1979 — — — 1980 —— = on 1981 328 290 432 1982 3,310 7,808 32 1983 — — — 1984 — == zee Kg TN (March—June) Year STD LFT Outlet 1975 — 289, 042 — 1978 — — = 1979 — — = 1980 — = a 1981 20,693 16,579 29,284 1982 2S NO 153,438 2,219 1983 578,087 137,337 138,669 1984 834,370 183,645 234,241 Kg TSSx1000 (March—June) Year STD LFT Outlet 1975 — = z= 1978 == os 1979 — — — 1980 — eae ae 1981 2,289 456 1,396 Vol. 46, No. 4 Table 1 continued. Kg TSSx1000 (March—June) Year SD Era Outlet 1982 63,403 13,388 193 1983 38,535 18,850 12,586 1984 73,991 22.655 41,645 Kg PO,—P (March—June) Year STD EAL Outlet 1975 — 1,602 — 1978 — — — 1979 2,703 582 .08 1980 13,442 5,494 0 1981 249 195 376 1982 4,263 1,244 20 1983 16,188 Set 2,883 1984 5,435 1,155 Sul Kg TIN (March—June) Year STD LFT Outlet 1975 — 7 TALL —_— 1978 54,951 33,877 0.40 1979 15,323 13,965 1.00 1980 83,485 30,347 0 1981 5,193 3,173 4,042 1982 61,847 30,257 471 1983 24,791 12,607 6,968 1984 68,475 29,286 36,536 traditionally been used almost exclusively for rural-agricultural purposes. The high moun- tain lands are used primarily for grazing, wa- tershed protection, and some recreation, whereas the land uses in the foothills sur- rounding the lake are grazing, dry farming, and recreational home sites. The valley floor adjacent to the lake is used for irrigated crop- lands, pasture for native grasses, and the ma- jor sites for summer homes and subdivisions, which are being developed at a rapid rate. The tributary discharges from the watershed for the years 1975 to 1984 can be seen in Table 2 and Figure 4. As with the Bear River, 1977 and 1981 were dry years and 1983 and 1984 were wetter than average. The associated to- tal phosphorus budgets for these watersheds can be seen in Table 3. As can be seen from these data, 13% of the total phosphorus input into Bear Lake is by wet fall from atmospheric precipitation, whereas 20% is from the en- demic watershed. The vast majority (67%) is from the Bear River at Lifton. The areal phos- phorus loading (g P/m*? Bear Lake surface/ year) ranges from 0.045 g P/m?/year to 0.136 g _ P/m’/year for the five years studied. October 1986 LAMARRA ETAL.: BEAR LAKE BASIN HYDROLOGY 695 TABLE 2. The hydrologic inputs to Bear Lake between 1975 and 1984. Flows (ac-ft x 1000) Year Bear River (Lifton) % Bear Lake watersheds (%) Precipitation (%) Total 1975 179.34 (48) 140.6 (37) 55.6 (15) 375.5 1976 221.1 (53) , 119.3 (29) 74.6 (18) 415.0 1977 9.8 ( 8) 49.2 (41) 61.3 (51) 120.3 1978 183.6 (47) 127.6 (33) 765 (20) 387.7 1979 86.4 (39) 98.8 (44) SIS (17) 2228S 1980 279.6 (51) 174.1 (32) 96.2 (17) 549.9 1981 34.0 (23) 60.5 (41) 52.8 (36) 147.3 1982 264.8 (48) 169.0 (30) 120.3 (22) 554.1 1983 251.5 (45) 203.1 (36) 106.3 (19) 560.9 1984 244.1 (42) 264.1 (45) 73.9 (13) 582.1 Bear Lake Iributary Inflow 60 50 40 30 Flow (1000 ac—ft) 20 1975 1976 1977 1978 1979 1980 1981 1982 19835 Time (years) Fig. 4. The flows from the endemic Bear Lake watershed between 1975 and 1984. BEAR LAKE LIMNOLOGY.—The first re- ported limnological investigation of Bear Lake was conducted in 1912 by Kemmerer et al. (1923). Since then numerous studies have been made of Bear Lake. An attempt will be made here to summarize the results of limno- logical investigations on the key physical, chemical, and biological characteristics of the Bear Lake ecosystem. Bear Lake is oval shaped, about 34 km long and 14 km wide. It has an 81 km shoreline and a surface area of 284 km° (Fig. 1). Bear Lake 696 GREAT BASIN NATURALIST Vol. 46, No. 4 TABLE 3. The total phosphorus budgets for the major tributaries to Bear Lake for the years 1975 and 1981 to 1984. Years (Kg P/Year) Streams 1975! 1981° 1982° 1983° 1984° Average percent of total Lifton (Bear River) 9,982 8,340 18,910 28,362 24,410 67 Swan Creek 495 710 1,860 1,976 2,913 6 Big Creek 1,405 840 1,250 1,716 1,481 6 Fish Haven 125 30 570 = — Little Creek 190 310 600 — = North Eden 920 100 120 = == Falula Spring 780 190 190 — — 8 South Eden — 29 40 = = Irrigation return and other streams 640 88 81 2,653 2,320 Precipitation 1,901 1,860 2,680 3,798 2,551 13 Septic tank leakage 95 260 260 260 260 << Total 16,533 12,757 26,561 38,765 33,935 100 Loading (g P/m’/yr) .058 045 094 136 nei "EPA 1975 “Lamarra et al. 1984 his study TABLE 4. The physical and chemical characteristics of Bear Lake, Utah. Chemical characteristics are from Lamarra et al. (1984). Physical-morphometric characteristics Surface area 282 km? Mean depth 27m Maximum depth 63.4 m Volume 7.89 x 10° m? Mean hydraulic retention time 92 years Chemical characteristics Alkalinity 265 mg as CaCO, /] Cae 25 mg Ca+ +/] MG++ 75 mg Mg+ +/] K+ 3.1 mg/l Na+ 39.1 mg/] Cl- 54.2 mg/] SO, 19.7 mg/l Suspended solids 5.0 mg/] Total dissolved solids 457 mg/] Volatile suspended solids 1.5 mg/l Total solids 475 mg/l] has been described as dimictic with a distinct thermocline at 15-17 m. Summer surface temperatures range bewteen 20C and 22C, and hypolimnetic temperatures are usually below 7C. The maximum temperature fluctu- ations of hypolimnetic water below 50 m have been found to be 2C to 7C. Part of the north and northwest shores of the lake are covered with emergent plants. The remaining shore- line is composed of sand and rock. However, the rocky zone is not extensive, extending only 4 m into the lake. Hypolimnetic sedi- ments are made of marl. TABLE 5. The mean summer areal oxygen deficits (mg O,/cm?/day) Chl a (ug/l) concentrations in Bear Lake from 1975 to 1984. Oxygen deficits Chlorophyll a(+ S. E.) Year (mg O;/cm*/day) (g/l) 1975 .026 - —_ 1976 .049 0.41 + .05 1977 .012 0.30 + .05 1978 .049 0.66 + .13 1979 .037 0.62 + .06 1980 .057 0.78 + .11 1981 .031 0.39 + .03 1982 .043 0.59 + .05 1983 — 0.90 + .09 1984 .054 0.71 + .10 Oligotrophic .055 >7.4 Hutchinson 1957 Vollenweider and Kerekes 1980 The macrochemical constituents found in Bear Lake are rather unique in their relative abundance (Table 4). Each investigation on Bear Lake has shown that Mg’ *>CA‘*> Na'>K° and HCO;>Cla>SOp_COw Conductivities range between 720 and 680 umhos/cm at 25C and pH between 8.3 and 9.0. The surface oxygen concentrations dur- ing the summer in Bear Lake are near satura- tion, based on temperature and pressure. However, hypolimnetic concentrations were found to be less than 50% of saturation. The mean summer areal oxygen deficits (rate of oxygen loss in the hypolimnion) between 1975 and 1984 can be seen in Table 5. October 1986 LAMARRA ET AL.: BEAR LAKE BASIN HYDROLOGY 697 Bear Lake Limnetic Station: TN/TP a TN/TP Ratio 10 1982 1983 1984 Time (years) Fig. 5. The total nitrogen to total phosphorus ratio for the surface and 10 m station in Bear Lake between 1981 and 1984. Phosphorus was found to be limiting TN:TP ratio >17, 88% of the time. An initial analysis of the limnetic nutrient data has indicated that phosphorus appears to be the dominant limiting element in the Bear Lake system (Fig. 5) and therefore the nutri- ent of most concern. Furthermore, the mean summer total phosphorus concentrations in the epilimnion of Bear Lake have been steadily increasing over the last nine years (Fig. 6). Because of the uniform shoreline in Bear Lake, rooted plants in the littoral zone of the lake are scarce, therefore relegating the domi- nant primary production in the lake to the limnetic phytoplankton. The seasonal distri- _ bution of the surface (epilimnetic) and subsur- face (metalimnetic) phytoplankton biomass | ] | | | | } } 1 can be seen in Figure 7. These data indicate that during the summer months the highest density of phytoplankton occurs between 20 and 30 m below the surface (metalimnetic). In addition, the average summer surface chloro- phyll a concentrations for 1976 through 1984 are provided in Table 5. Although these data are not representative of the highest algal densities, they do provide a historical per- spective of water quality changes within the surface waters of Bear Lake. DISCUSSION The limnological conditions present in Bear Lake over the last decade provides some in- teresting insights into the temporal dynamics of this lake ecosystem. The algal biomass in the lake (expressed as a spring-summer aver- age for chlorophyll a) has increased in concen- tration since 1976, reaching a maximum in 1983 and declining slightly the following year 698 16 14 12 10 TOTAL PHOSPHORUS(ug P/1) (ee) 1976 1977 1978 GREAT BASIN NATURALIST 1979 Vol. 46, No. 4 Y:.085X + 6.42 N:28 12.70 1980 1981 1982 1983 YEARS Fig. 6. The mean summer months (April-September) total phosphorus concentrations from the epilimnion of Bear Lake. Sources of the data are from Lamarra et al. 1984. (Table 5). In a similar manner, areal oxygen deficits (Table 4) and phosphorus loading (Table 2) have demonstrated a year-to-year fluctuation. The concept of a trophic classification for lake ecosystems has long been recognized. Early studies investigated the quality of plankton and have been summarized by Raw- son (1956). More recently, trophic state has been defined by nutrient loading (Vollen- weider 1976), complex ecosystem models (Si- mon and Lam 1980, Ditoro and Matystik 1980), and the interrelationships ofa variety of parameters (Porcella 1980). A coarse resolu- tion technique used by Carlson (1977) re- sulted in using single but interrelated parameters. Total phosphorus, chlorophyll a, and Secchi disk transparency have been shown to provide an excellent basis for a trophic state index (TSI). However, because of the presence of CaCO, precipitates in Bear Lake and its effect upon phosphorus availabil- ity, Chl a was determined to be the most representative parameter for a TSI calcula- tion. A comparison has therefore been made in Table 6 between the Chl a TSI value, areal phosphorus loadings (g P/m?/year), and areal hypolimnetic oxygen deficits (mg O,/em’*/ day). In each case the static (TSI), dynamic (areal oxygen deficits), and predictive (areal phosphorus loadings) trophic state classifica- tions indicate that Bear Lake is upper olig- October 1986 | LAMARRA ET AL.: BEAR LAKE BASIN HYDROLOGY Chl a Concentration at BL—M Site 699 3.0 LEGEND © Epilimnetic ® Metalimnetic 2.5 2.0 Chl a (ug/)) z 0.5 0.0 JJASONDJIFMAM 1981 JJASONDJFMAM 1982 / e 1 ! 1 1 J 1 1 ! ! 1 1 ! 1 I 1 1 1 i 1 1 4 ! 1 t ! ! ! 1 \ JJASONDJSFMA 1983 1984 Fig. 7. The temporal distribution of chlorophyll a in the epilimnion (surface and 10 m) and metalimnion (20 and 30 m) in Bear Lake, Utah-Idaho. TABLE 6. The estimated TSI values (Carlson 1977) for chlorophyll a, the areal oxygen deficits (mg O./cm?/day), and the total phosphorus areal loadings (g P/m’/yr) for Bear Lake between 1975 and 1984. I. TSI parameter (chlorophyll a wg/l) N = 140 TSI values Average (1975-1984) 0.60 29 Range .19-3.0 14-45 Oligotrophic”” <.80 <30 Mesotrophic .80-7.4 30-60 Eutrophic >7.4 >60 II. Areal oxygen deficits (mgO,/cem?7/day) N=9 Average (1975-1984) .040 _ Range .012—.057 | Oligotrophic ” <.025 _ Mesotrophic .025-.055 Eutrophic >.055 Ill. Areal phosphorus loading | (g P/m*/yr) IN| = & | Average 1975 (1981-1984) 090 | Range .058—. 136 Oligotrophic® <07, _ Mesotrophic .07-.15 | Eutrophic >.15 Carlson 1977 _- @ Hutchinson 1957 _ Vollenweider 1976 otrophic to strongly mesotrophic. Because Bear Lake has been previously classified as oligotrophic (Kemmerer et al. 1923), the driv- ing factors for the observed trophic changes need to be elucidated. The limnological trends presently observed in the lake may be the result of increased human activity within the basin, increases in the hydrologic inputs, or a combination of these factors. Compre- hensive sets of water quality data for the Bear River, Bear Lake watersheds, and Bear Lake do not exist prior to 1975. As an alternative to a historical data base, inferences to previous water quality conditions in Bear Lake can be made from the extensive hydrological data available. Based upon the data presented here, a series of regression equations were produced that indicated the watershed load- ings of phosphorus and inlake water quality parameters (summer chl a and oxygen deficits) were significantly related to mass flows from the watersheds (Table 7). Based upon these statistical relationships and the historical flow data, hydrologic and nutrient budgets were developed for the recent history Vol. 46, No. 4 GREAT BASIN NATURALIST Bear Lake Surface Elevation 700 Legend (ZA With Bear River 9, Keg reetererereecee KX) Without Bear River DOP Do PoP PPP Poo PoP PoP PoP Poo PPPS PPP PPPS 195255255255 OC OO LO 255050505099 C COCO OOOO OOO IO OS A IW AAAAAANSAAANAANAAASAANASSAAAANSSAN] DRLAEARARARABRARBAARBARAEABRAARARACAARESA IX.AANANANANANSSAANAAN Time (years) KAAANANANANANASASAN KANAAANAANAANANAN KANANANNAANS aN _ b@ re O O Q “29 oO x O) ‘ee 5) O O se O oO oe Oo O ee O) oe O © O O o O O oe O O Q) > o 0 O O O <4 ‘ee o we O ee o. LEGEND Recorded Elevation qeloee AAA nggecncescceeceencet” __.Simulated W/O Bear River _ 1922 1926 1930 1934 1938 19411944 1948 19511954 195819611964 1968 197119741977 1981 5910 5900 ° N fez) wo ¢ S bo) re) ra) 25 Ww (e) ve) Oo }Y4) UORDAA!A a = a Fig. 8. The simulated elevations in Bear Lake without the Bear River and the natural elevations with the operation of the Bear River storage system between 1924 and 1984. (quacued) Aouenbe.4 3929" 3950" sogs5 5920 5915 Elevation (feet) Fig. 9. A frequency distribution of monthly Bear Lake elevations (1924 to 1984), with a simulated watershed 5910 excluding the Bear River and actual elevations with the Bear River. SEO) SENOS October 1986 75000 60000 45000 30000 Phosphorus Loading (Kg P/year) 15000 LAMARRA ET AL.: BEAR LAKE BASIN HYDROLOGY 701 Phosphorus Loading to Bear Lake LEGEND Bear River Bear Lake Tributaries sree: 19241927 1931 1935 19391942 1945 194919521955 1959 1963 1967 1970 1973 1976 1979 1983 Water Year Fig. 10. The estimated mass (kg P/year) of phosphorus entering the Bear Lake system between 1924 and 1984. Frequency (percent) Frequency Histogram of P Loading 100 Legend ZZ] Without Bear River RX With Bear River 80 60 | 40 Oligotrophic Mesotrophic Eutrophic 20 RRR as "@ =e GO S62 400 “iSO. 8200 , 280 30.0 - Phosphorus Loading (g P/m* /year) *10 Fig. 11. The frequency distribution of areal total phosphorus loading (g P/m*/yr) for Bear Lake with and without the Bear River for the period 1924-1984. I S bo GREAT BASIN NATURALIST Vol. 46, No. 4 Bear Lake Chlorophyll a 0.75 0.70 0.65 0.60 — S D =e ea 0'55 ol Ss a fe) 0.50 2 1S O 0.45 0.40 0.35 0.30 1925 1928 19311934 1938 1941 1944 19471950 19541957 19611964 1967 1970 1973 1976 1979 1982 Water Year Fig. 12. The historical distribution of mean summer chlorophyll a concentrations as simulated from the empirical relationships developed from the years 1975 through 1984. (1923—-present) in Bear Lake with and without Bear River inflows. The major assumption made in this analysis was that the historical water quality in the Bear River had not signifi- cantly changed and is similar to the period 1975-1984. The hydrologic budget for Bear Lake was developed using all sources and losses on a mass balance basis. Input data for equation (1) was obtained from Utah Power and Light Company. KE=T=O\a where: AAS =Annual change in Bear Lake storage (ac-ft/year) I = Rainbow canal inflow (ac-ft/ year) O = Outlet canal flow (ac-ft/year) S = Actual Bear Lake storage from elevation capacity curves (ac- ft/year) The results of this analysis, with and with- out the Bear River inflows can be seen in Figure 8. The data indicate that the simulated elevations in Bear Lake without the river were higher (except for the years 1944-1950) than the lake elevations with the river inflow- outflow manipulations. The estimated eleva- tions of the lake indicated that the threshold of 5,927.0 ft between Bear Lake and the marsh complex would have been exceeded about 24% of the time during the last 60 years (Fig. 9), providing a direct connection between the shallow marsh in the northern valley and the lake to the south. In addition, the simulation indicates that during the 1970s the lake would have had a steady increase in elevation above 5,924 ft to a high elevation of 5,935 ft in 1984. This increase in lake elevation would inundate the confluence of the Bear River and the Bear Lake valley, thus naturally adding a 7,000 km” watershed to the Bear Lake drainage. During the same time period simulated in the hydrologic budgets, the annual phospho- rus loading (kg/year) by source was estimated (Fig. 10). It appears that about 60% of the historical loading to the lake can be attributed October 1986 LAMARRA ET AL.: BEAR LAKE BASIN HYDROLOGY 703 Bear Lake Oxygen Deficit Oxygen Deficit (mg 0,,/em* day) 0.00 19241927 1931 1935 193919421945 194919521955 1959 1963 1967 19701973 1976 1979 1983 Water Year Fig. 13. The historical distribution of mean summer areal oxygen deficits (mg O,/cm*/day) as simulated from the empirical relationships developed from the years 1975 through 1984. to the Bear River, and in only 10 years of the last 60 has the endemic watershed produced more phosphorus loading than the Bear River. A frequency analysis of areal phospho- rus loadings indicates that with the presence of the Bear River, 58% of the last 60 years the loading could be considered oligotrophic, while 42% could be considered mesotrophic or eutrophic. In contrast, without the Bear River inflows 100% of the annual loadings would be oligotrophic (Fig. 11). In a similar manner, the simulation of the historical chlorophyll a and areal oxygen deficits (Fig. 12, 13) with the presence of the Bear River demonstrates the importance of this water source in modifying the Bear Lake _ environment. Frequency histograms for both _ parameters (Fig. 14, 15) demonstrate patterns _ similar to those expressed by areal phospho- rus loadings, indications that the Bear River may have shifted the trophic state of Bear Lake from oligotrophic to mesotrophic. The Bear Lake ecosystem is a unique envi- ronment. Because of its isolation for more than 8,000 years, the biological community has evolved into a simple, coexisting trophic structure, with four endemic species of fish. The uniqueness of the Bear Lake community lies in the adaptations of the organisms to one another and the importance of the endemic fish to the overall trophic structure. For exam- ple, the cisco is a dominant food item of the large predators and is, itself a planktivore, feeding exclusively on zooplankton within the metalimnion during summer stratification. In turn, the zooplankton community has few large cladocerans, with its structure domi- nated by a large Epischura sp. This organism has adapted a swift predatory escape mecha- nism. Because the effect of water quality changes upon these species is unknown, defining the driving factors and their degree of impact upon changes in water quality may provide management alternatives for this ecosystem. The results of this preliminary investigation have inferred the historical impacts of the Bear River inflows upon the limnological con- Vol. 46, No. 4 GREAT BASIN NATURALIST 704 Bear Lake Chlorophyll a SG GCC[O Quedied) Aouenbe.4 0.6 Gaorestarl a (ug /\) Fig. 14. A frequency histogram of the simulated mean sum NJ : v (e) ugh 1984. (wg/l) for the years 1924 thro Bear Lake Oxygen Deficits (Qjuacsied) Aouenbe4 Nie) Nps aS 5 aa OF O_O IS ~ @ on) oO L3 = cee st OC Gq ) N S ro) on) SS x (mg O,/em?/d ay) for the years 1924 eal deficits cy histogram of the simulated mean summer ar Fig. 15. A frequen ugh 1984. thro October 1986 ditions in Bear Lake, based upon 9 years of empirical limnological data and 60 years of historical flows. This impact has been exten- sive. In addition, the future human develop- ments of the Bear Lake basin will only in- crease the nutrient export from the watershed to the lake environment. Mitigation measures that directly address sources of nitrogen and phosphorus within the watersheds must be developed. Increased eutrophication may re- sult in the loss of several if not all of the endemic species. In a similar manner, the investigation of alternative hydrologic storage of the Bear River as it relates to Bear Lake seems advisable. The development of 100,000 ac-ft of storage above Bear Lake may reduce previously described oxygen deficits and cut the phosphorus loading by 7,000 kg/year. LITERATURE CITED AMERICAN PUBLIC HEALTH ASSOCIATION. 1980. Standard methods for the examination of water and waste- water. Edition 15. APHA. Washington, D. C. 1134 pp. ARMSTRONG, F. G., AND E. R. CRESSMAN. 1963. The Ban- nock thrust zone in southeastern Idaho: U.S. Geol. Survey Prof. Paper 374-J. CarRSLON, R. E. 1977. A trophic state index for lakes. Limnol. Oceanogr. 22: 361-9. Diroro, D. M., AND W. F. MatysTIK. 1980. Mathematical model of water quality in large lakes, part 1: Lake Huron and Saginaw Bay. EPA-6000/3-80-056. U.S. Environ. Prot. Agency, Duluth, Minnesota. HERRON, R., V. A. LAMARRA, AND V. D. ADAMS. 1984. The nitrogen, phosphorus, and carbon budgets of a large riverine marsh and their impacts on the Bear Lake ecosystem. In Lake and reservoir manage- ment. EPA 440/5/84-001. HUTCHINSON, G. E. 1957. A treatise on Limnology. Vol. I. Geography, physics, and chemistry. John Wiley and Sons Inc. New York. 1015 pp. LAMARRA ET AL.: BEAR LAKE BASIN HYDROLOGY 705 KALISER, B. N. 1972. Environmental geology of Bear Lake, Rich County, Utah. Utah Geological and Mineralogical Survey, Bulletin 96. Salt Lake City, Utah. 32 pp. KEMMERER, G., J. R. BOVARD, AND W. R. BoorMaN. 1923. Northwestern lakes of the United States; biologi- cal and chemical studies with reference to possibil- ities to production of fish. U.S. Bur. Fish., Bull. 39: 51-140. LamarRaA, V. A., V. D. ADams, C. THomas, R. HERRON, P. BIRDSEY, V. KOLLOCK, AND M. PILLs. 1984. A his- torical perspective and present water quality con- ditions in Bear Lake, Utah-Idaho. Pages 213-218 in Lake and reservoir management. EPA 440/5/ 84-001. MANSFIELD, G. R. 1927. Geography, geology, and min- eral resources of part of south-western Idaho: U.S. Geol. Survey, Prof. Paper 152. McC ure, L. W. 1970. Source rocks and sediments in drainage area of North Eden Creek, Bear Lake Plateau, Utah-Idaho. Unpublished thesis, Utah State University, Logan. 84 pp. NuNAN, J. 1972. Effects of Bear River storage on water quality of Bear Lake. Unpublished thesis, Utah State University, Logan. PoRCELLA, B. 1980. Index to evaluate lake restoration. J. Environ. Eng. Div. Proc. Amer. Soc. Civil Eng. 106: 1151. Rawson, D. W. 1956. Algal indicators of trophic state types. Limnol. Oceanogr. 1: 18-25. RICHARDSON, G. B. 1913. The Paleozoic section in north- ern Utah: Amer. Jour. Sci., Ser. 4, 36: 406-416. ROBERTSON, G. C. 1978. Surficial deposits and geological history, northern Bear Lake Valley, Idaho, Un- published thesis, Utah State University, Logan. SIMON, T. J., AND D. C. Lam. 1980. Some limitations on water quality models for large lakes: a case study of Lake Ontario. Water Resour. Bull. 16: 105. VOLLENWEIDER, R. A. 1976. Advances in defining critical loading levels for phosphorus in lake eutrophica- tion. Mem. Ist Ital. Idrobiol. 33: 53-83. VOLLENWEIDER, R. A., AND J. S. KEREKES. 1980. Back- ground and summary results of the OECD cooper- ative-program on eutrophication. Pages 25-36 in International symposium on inland waters and lake restoration. September 8—12, 1980. Portland, Maine. USEPA EPA 440/5/81-010. UNDERSTORY SEED RAIN IN HARVESTED PINYON-JUNIPER WOODLANDS Richard L. Everett! ABSTRACT.—Seed rain was collected on six paired tree harvest and undisturbed plots in singleleaf pinyon (Pinus monophylla)—Utah juniper (Juniperus osteosperma) stands. Approximately 14,600 seeds were collected during four years. Seed rain in undisturbed plots was similar to levels in mixed forest communities. Seed rain on harvest plots was similar to disturbed sites and grasslands. Seed rain levels reflect the current successional stage rather than the climax community type for the site. Seed rain increased in numbers and seed production per unit of plant cover following tree removal and especially on transition soil microsites. Only three to four of the plant species present on a site contributed greater than 10% of the total seed rain. Seed rain composition was similar on harvest and undisturbed plots (Jaccard Similarity Index Values = 47% to 67%) and explains in part the rapid reestablishment of predisturbance understory communities. Understory response to tree harvesting is dependent upon remnant plants (Dyrness 1973, Lyon and Stickney 1976), soil seed re- serves (Oosting and Humphreys 1940), and seed rain (Rice et al. 1960). Seed rain, the dissemination of seed and its dispersal, varies among species and plant communities (Harper 1977). Plant strategies in part control the amount of resources a plant commits to seed production and the impact the species has on community seed rain (Grime 1977, Everett and Sharrow 1983). Seed rain declines exponentially with dis- tance from the parent plant (Werner 1975, Harper 1977). But seeds are often transported across the soil surface until they lodge against protuberances or depressions (Knipe and Springfield 1972). Nelson and Chew (1977) found greatest concentrations of soil seed re- serves in the Mojave Desert, adjacent to shrub canopies, and hypothesized that shrub litter areas acted as seed rain catchments. In pinyon-juniper woodlands soil seed reserves were greatest at the edge of the tree crown (Koniak and Everett 1982). Seed rain composition changes abruptly from one year to the next in forests. This in part is due to the abrupt appearance and dis- appearance of species during forest succession (Oosting and Humphreys 1940) and variable annual precipitation (Duba and Norton 1976). This paper reports on a four-year study of seed rain following tree harvesting in single- leaf pinyon (Pinus monophylla Torr. & ‘Intermountain Research Station, USDA Forest Service, Ogden, Utah 84401. Frem.)—Utah juniper (Juniperus osteosperma [Torr.] Little) woodlands. Seed rain was ob- served on three grass and annual forb under- story sites for four years and three shrub and annual forb understory sites for two years. The null hypotheses to be tested were: (1) there are no differences in seed rain numbers or composition between harvest and undis- turbed plots or soil microsites, (2) seed rain is evenly distributed among plant forms and re- flects species composition, and (3) seed rain production per unit of cover remains constant following tree harvest. Increased seed rain following tree harvest would promote rapid establishment of understory and if related to floristic composition would increase pre- dictability of plant response. FIELD METHODS AND DATA ANALYSIS Previous study sites in stands of singleleaf pinyon—Utah juniper on the Sweetwater and Shoshone mountain ranges were selected for study. The Sweetwater site lies above alluvial fans that are dominated by mountain big sage- brush (Artemisia tridentata Nutt. ssp. vaseyana) and below a ponderosa pine (Pinus ponderosa Doug}. ex Laws.) community. The site has an average annual precipitation of 266 mm. Fully stocked stands have a remnant mountain big sagebrush—annual forb under- story. The three plots used for the study had a similar east exposure (N 70° E) but varied in elevation from 2,040 to 2,280 m. 706 October 1986 The Shoshone site lies between communi- ties of higher elevation mountain big sage- brush and lower elevation Wyoming big sage- brush (Artemisia tridentata Nutt. ssp. wyomingensis ). The site has an annual precip- itation of 330 mm. The three study plots are on north- (N 20° E), west- (S 84° W), and south-facing (S 16° E) aspects at approxi- mately the same elevation (2,300 m). Under- story is sparse but includes perennial grasses: Idaho fescue (Festuca idahoensis Elmer), Sandberg bluegrass (Poa sandbergii Vasey), bottlebrush squirreltail (Sitanion hystrix Nutt.); perennial forbs: Hood’s phlox (Phlox hoodii Rich.), hollyleaf clover (Trifolium gym- nocarpum Nutt.); and an array of annual forbs. The soil surface in both woodland study areas is a mosaic of soil microsites: a dense needle duff under the tree, scattered needle cover in a transition zone at the crown edge, and bare ground in the interspace between trees. These soil microsites vary in species composition and productivity (Everett 1984). Three 30 by 30 m square plots previously clear-cut of all trees greater than | m in height were used at Shoshone and Sweetwater sites. Sweetwater plots were cut in 1977 and Shoshone plots in 1979. Trees and slash were removed from the sites by hand with minimal disturbance to the plots. Areas of similar floristic and topographic appearance were se- lected adjacent to cut areas to serve as undis- turbed controls. Shoshone plots were sampled for seed rain in the first and second years following harvest (1980 and 1981) using 12 seed traps randomly placed on four duff, four transition, and four interspace soil microsites at each of the three harvest and control plots. The study was ex- panded in 1982 and 1983 to include three paired harvest and control plots at the Sweet- water site (five and six years following har- vest). Sampling intensity was increased to 24 seed traps in each harvest and undisturbed plot at Sweetwater and Shoshone sites in 1982. Seed traps were emptied of seed every 2 to 3 weeks from May to October. Traps were left in the field during the winter months and inspected the following spring. A seed voucher was made for all plant species en- countered on the sites. Seeds collected in the traps were compared with voucher specimens for correct identification. EVERETT: PINYON-JUNIPER WOODLANDS 707 Seed traps consisted of 90 mm dia. petri dishes coated inside with an adhesive (Tangle- foot’), as suggested by Werner (1975). Each dish had a central hole (3 mm dia.). We placed a screen (5 mm mesh) over the traps to pre- vent predation by rodents and inserted a nail through the screen and dish to hold the trap on the soil surface. Differences in seed rain among harvest and undisturbed plots and among soil microsites were compared in analysis of variance tests using Hartley's Sequential method of testing to identify significant differences between means (Snedecor and Cochran 1978). To measure plot understory cover, five per- manent transects (20 m) were established in each plot. Shrub cover was estimated by line intercept (Canfield 1941). Herbaceous spe- cies cover was estimated by the Daubenmire (1959) canopy coverage method using sam- pling frames (50 x 50 cm) laid down at each meter mark. In addition, we centered circular sampling frames (50 cm dia.) over each seed trap and estimated plant cover of seed produc- ing species. Relationships between seed rain and plant cover of seed-producing species were evalu- ated by linear regression. Similarity between plot seed rain and floristic composition was evaluated using Jaccard Similarity Index Value (SIV) (Mueller-Dombois and Ellenberg 1974). CS x 100 (CS + SSR + SFC) Where: CS = species common to seed rain and floristic composition SSR = species in seed rain only SFC = species in floristic composi- tion only SIV = Similarities in seed rain and floristic composi- tion between harvest and undisturbed plots were also compared. RESULTS AND DISCUSSION We collected 14,676 seeds in seed traps from 1980 to 1983. Understory seed rain was >The use of trade, firm, or corporation names in this paper is for the information and convenience of the reader. Such use does not constitute an official endorsement or approval by the U.S. Department of Agriculture of any product or service to the exclusion of others that may be suitable. 708 TABLE 1. Mean seed rain (seeds/dm’) on tree harvest and undisturbed plots. Year Shoshone Sweetwater Harvest Undisturbed Harvest Undisturbed (seeds/dm?) 1980 20.3°° 2.9 a = ISL SEO 1.6 = _ 1982 23.9° 5.0 10.4° 1.6 1983. 98.3°° 3.9 61.9 19.9 ‘Significant difference (p = 0.05) between harvest and undisturbed plots. TABLE 2. Seed rain (seed/dm’) by community types Seed rain Plant community Citation 1,000 Meadow Mortimer 1974 115-500 Alien annuals Duba and Norton 1976 20-98 Disturbed woodland (this study) 24-42 Mine waste-prairie Archibold 1980a 22 Grassland 2-20 Woodland 3-9 Mixed forest Knipe and Springfield 1972 (this study) Archibold 1980b TABLE 3. Mean proportion of seed rain by plant form for Shoshone and Sweetwater sites for the study period. Shoshone Sweetwater Harvest Undisturbed Harvest Undisturbed (percent) Annual 0.0 0.5° 0.0 0.0 grass Perennial 36.8? 59.3" 9° 3.0° grass Annual 63.2? 39.5° 99.1? 97.0° forb Values in the same column with different superscripts are significantly different (p = 0.05). estimated at 1.6 to 20 seed/dm? in undis- turbed stands and from 10 to 98 seeds/dm? in harvested plots (Table 1). Seed rain was al- ways greater on harvested than on undis- turbed stands. This in part explains previous findings that soil seed reserves decline rapidly from early to late successional stages (Koniak and Everett 1983). Seeds/dm’? equates to mil- lions of seeds per hectare. Undisturbed pinyon-juniper woodlands had seed rain totals similar to mixed forest communities in Canada (Archibold 1980a). These levels are lower than in other community types (Table 2). Seed rain on tree harvest sites approxi- mates seed rain levels reported for disturbed sites and grasslands. On the Shoshone site, plant-form composi- tion of seed rain on harvested sites was grass GREAT BASIN NATURALIST Vol. 46, No. 4 37% and forbs 63%, and on the undisturbed sites grass 59% and forbs 39% (Table 3). Seed rain on Sweetwater sites was composed al- most entirely of annual forb seeds (97% to 99%) on harvested and undisturbed sites. An- nual forbs produce copious amounts of seed (Biswell and Graham 1956, Archibold 1980a, b), but populations of annuals change dramati- cally from one year to the next (Treshow and Allan 1979). Although shrub and perennial forb species were present on both sites, their seed made up less than 1% of the seed rain. Lodging of mountain sagebrush seed stalks by snow was observed and seed dissemination may not have occurred. At both Shoshone and Sweetwater sites the proportion of annuals in seed rain declined in undisturbed stands. Only three to five species made greater than 10% contribution to total seed rain on any site. On Shoshone plots squirreltail bot- tlebrush and Sandberg bluegrass each con- tributed 10% to 50% of the total seed rain. The annual forb Cryptantha watsonii (A. Gray) Greene made up 16% to 25% of total seed rain on harvest plots but only 1% on undisturbed sites. Microsteris gracilis (Hook.) Greene contributed to the seed rain (11% to 21%) on both harvested and undisturbed plots. Seed rain on the Sweetwater site was domi- nated by an array of annual species including Collinsia parviflora Dougl. (7% to 54%), Cryptantha watsonii (0% to 22%), Gayophy- tum ramosissimum Nutt. (5% to 32%), Gilia gilioides (Benth.) Greene (1% to 17%), Mi- crosteris gracilis (1% to 23%), and Phacelia humulis Torr. and Gray (2% to 20%). Archi- bold (1980a,b) previously reported the domi- nance of seed rain by annual forb species on disturbed sites. Seed rain composition does not mirror floristic composition. Shoshone and Sweetwa- ter sites had Jaccard Similarity Index Values (SIV) of 23% to 78% between floristic compo- sition and seed rain (Table 4). The proportion of perennial species that contributed to seed rain (58%) was significantly less (p=0.05) than the proportion of annual species (80%). Annu- als must set seed crops frequently, whereas perennials can forego seed production and still maintain a position in the plant commu- nity. SIV values were somewhat lower on the Shoshone site because of a greater proportion of perennial species (55% to 59%) than oc- curred on Sweetwater sites (26% to 48%). i . | | | : i October 1986 EVERETT: PINYON-J UNIPER WOODLANDS 709 TABLE 4. Similarity index values between seed rain and plant composition on harvest and undisturbed plots. 1982 1983 Harvest Undisturbed Harvest Undisturbed (percent) Sweetwater 78 71 23 65 Shoshone 29 31 39 33 Jaccard’s Similarity Index Values (Mueller-Dombois and Ellenberg 1974). Maximum SIV = 100. TABLES. Seed rain per dm’ cover of seed-producing species by soil microsite. Duff Transition Interspace Harvest Undisturbed Harvest Undisturbed Harvest Undisturbed Shoshone Bae SY 4.18% 157 2.05" 97 Sweetwater Bola 1.43 9.14 .65 1.56 OY 1Slope (b) from regression of species seed rain on species cover significantly different (p = 0.05) than 0 based on t-values. TABLE 6. Proportion of seed rain by soil microsites on harvest and undisturbed plots. Harvested Undisturbed Duff Transition Interspace Duff Transition Interspace Shoshone 28.0° 45.0° 26.8? 17.0 40.5 42.8 Sweetwater O5e5u BilW 5s DSP 22.0 34.0 44.0 Site values in the same row with different superscripts are significantly different (p = 0.05). Cover of seed-producing species adjacent to the seed traps was significantly (regression slope greater than 0) related to seed rain on harvest plot soil microsites (Table 5). Seed rain per unit area of plant cover was greater on harvested than on undisturbed plots for all soil microsites. Cover on harvested plots pro- duced 1.56 to 9.14 seeds per dm’ of cover, whereas undisturbed plots produced 0.27 to 1.47 seeds per dm’? of cover. In 1983 under- story cover on harvest plots (mean = 24.4%) was significantly greater (p=0.05) than that on undisturbed plots (mean=7.7%) (five of six plots). Both the quantity of understory and its ability to produce seed increased following tree removal. Seed rain was greater on the transition soil microsite than duff or interspace in the Sho- shone and Sweetwater harvest plots (Table 6). Differences in seed production among mi- crosites were less apparent in undisturbed stands. Tree competition in undisturbed stands may reduce differences in potential un- derstory seed production among microsites. Differences in seed rain among microsites may reflect and maintain previously de- scribed differences in understory species dis- tribution (Everett 1984). Similarity values for seed rain between har- vested and undisturbed plots ranged from 47% to 67% for both sites in 1982 and 1983. Similarity values for floristic composition ranged from 43% to 68% during this time. Tree harvest alters understory composition, but the similarity in seed rain is likely to be a contributing factor to the rapid reestablish- ment of predisturbance woodland communi- ties following tree harvest and burning (Ev- erett 1984, Everett and Ward 1984). CONCLUSIONS Seed rain increased following tree harvest- ing. Seed rain of 10 million to 98 million seeds per hectare fell to the soil surface in harvested pinyon-juniper woodlands. On undisturbed plots seed rain was less than 20 million seeds per hectare. Increased seed rain was a result of increased plant cover and increased seed production per unit of cover. Seed rain on 710 pinyon-juniper harvested plots had a large annual forb component as reported for other disturbed areas. Seed rain reflects the current successional stage and vigor of plants more than the climax community type. Prediction of seed rain composition from general species floristics would be difficult. Seed rain and floristic composition were not in close agreement (SIV = 20% to 70%). The character of seed rain was determined by the relative cover of a few species producing a majority of the seed. A significant linear rela- tionship existed for seed-producing species cover and its seed rain. But the proportion of perennial species contributing to seed rain was less than the proportion of annual species. The variability in annual populations reduces predictability of seed rain numbers and com- position. Similar seed rain composition on harvest and undisturbed plots (SIV = 47% to 70%) may in part explain the rapid reestablishment of predisturbance understory in the wood- land. Reduced seed rain in undisturbed stands in part explains the previously re- ported decline in soil seed reserves as succes- sion proceeds. Increased seed rain in the tran- sition microsite reflects and perhaps main- tains differences in understory distribution among soil microsites. LITERATURE CITED ARCHIBOLD, O. W. 1980a. Seed input into postfire forest site in northern Saskatchewan. Canadian J. For. Res. 10: 129-134. _____. 1980b. Seed input as a factor in the regeneration of strip-mine wastes in Saskatchewan. Canadian J. Bot. 58: 1490-1495. BISWELL, H. H., AND C. A. GRAHAM. 1956. Plant counts and seed production on California annual-type ranges. J. Range Manage. 9: 116-118. CANFIELD, R. H. 1941. Application of the line interception method in sampling range vegetation. J. For. 39: 388-394. DAUBENMIRE, R. 1959. Canopy coverage method of vege- tation analysis. Northwest Sci. 33: 43-64. Dua, D. R., AND B. E. Norton. 1976. Plant demographic studies of desert annual communities in northern Utah dominated by nonnative weedy species. Pages 68-100 in US/iBP Desert Biome Res. Memo. 76-6. GREAT BASIN NATURALIST Vol. 46, No. 4 Dyrness, G. T. 1973. Early stages of plant succession following logging and burning in the western Cas- cades of Oregon. Ecology 54: 57-69, EVERETT, R. L. 1984. Understory response to tree har- vesting in pinyon-juniper woodlands. Unpub- lished dissertation, Oregon State University, Cor- vallis, Oregon. 174 pp. EVERETT, R. L., AND S. H. SHARROW. 1985. Understory response to tree harvesting of singleleaf pinyon and Utah juniper. Great Basin Nat. 45: 105-112. EVERETT, R. L., AND K. Warp. 1984. Early plant succes- sion on pinyon-juniper controlled burns. North- west Sci. 58: 57-68. GriME, J. P. 1977. Plant strategies and vegetation pro- cesses. J. Wiley and Sons, New York. 222 pp. HARPER, J. L. 1977. Population biology of plants. Aca- demic Press, New York. 892 pp. KNIPE, O. D., AND H. W. SPRINGFIELD. 1972. Germinable alkali sacaton seed content of soils in the Rio Puerco Basin, west central New Mexico. Ecology 53: 965-968. Koniak, S. D., AND R. L. EVERETT. 1983. Soil seed reserves in successional stages of pinyon woodland. Amer. Midl. Nat. 108: 295-303. Lyon, L. J., AND P. F. STICKNEY. 1976. Early vegetal suc- cession following large northern Rocky Mountain wildfires. Pages 355-375 in Proceedings of the Montana Tall Timbers Fire Ecology Conference and Land Management Symposium, November 14, 1974. Tall Timbers Research Station, Tallahas- see, Florida. MORTIMER, A. M. 1974. Studies of germination and estab- lishment of selected species with special refer- ences to the fates of seed. Unpublished disserta- tion, University of Wales. MUELLER-Domsols, D., AND H. E. ELLENBERG. 1974. Aims and methods of vegetation ecology. J. Wiley and Sons, New York. 547 pp. NELSON, J. F., AND R. M. CHEw. 1977. Factors affecting seed reserves in the soil of a Mojave Desert ecosystem, Rocky Valley, Nye County, Nevada. Amer. Midl. Nat. 97: 300-320. OosTING, H. J., AND M. E. Humpureys. 1940. Buried vi- able seeds in a successional series of old field and forest soils. Torrey Bot. Club Bull. 67(4): 253-273. Rice, E. L., W. T. PENFOUND, AND L. M. ROHRBAUGH. 1960. Seed dispersal and mineral nutrition in suc- cession in abandoned fields in central Oklahoma. Ecology 41: 224-228. SNEDECOR, G. W., AND W. G. Cocuran. 1978. Statistical methods. Iowa State University, Ames, Iowa. TRESHOW, M., AND J. ALLAN, JR. 1979. Annual variation in the dynamics of a woodland plant community. Environ. Conserv. 6: 231-236. WERNER, P. A. 1975. A seed trap for determining patterns of seed deposition in terrestrial plants. Can. J. Bot. 53: 810-813. | | SPECIES DIVERSITY AND HABITAT COMPLEXITY: DOES VEGETATION ORGANIZE VERTEBRATE COMMUNITIES IN THE GREAT BASIN? David J. Germano! and David N. Lawhead” ABSTRACT. —In this study, we have examined the effect of vegetation structure on the three major vertebrate taxa in Great Basin habitats of southwestern Utah. The effect of increasing vegetation heterogeneity, both horizontally and vertically, on the diversities of lizards, rodents, and postbreeding birds was investigated. We found no statistically significant relationship between diversity of all animal taxa and horizontal vegetation heterogeneity, although lizard diversity tended to decrease with increasing heterogeneity and rodent diversity tended to increase. Bird species diversity was positively correlated with vertical habitat heterogeneity. Abundances were highest for rodents in pinyon/juniper habitat and highest for lizards and birds in areas with the highest grass cover. Species richness was highest in sagebrush habitat for rodents but highest for lizards and birds in pinyon/juniper. Evenness values were relatively similar and high for birds and rodents and were relatively high for lizards in all habitats except for pinyon/juniper, which had an evenness value of 0.38. For rodents and lizards, abundance was significantly correlated with the index for horizontal habitat heterogeneity. After logarithmic transformation, abundance of lizards was positively correlated with increasing vegetation complexity. Combined abundance of lizards and rodents was also positively correlated with vegetation complexity. Rodent and lizard abundances, however, were affected by different aspects of the habitat. After logarithmic transformation, lizard abundances increased significantly with increasing grass cover, whereas rodent abundances increased significantly with increasing shrub cover. Spatial heterogeneity, or simply the com- plexity of vegetation structure both horizon- tally and vertically, appears to predict species diversity in some instances, and many authors have felt that this is the primary factor causing differences in species diversity in communi- ties (Pianka 1967, Rosenzweig and Winakur 1969, Karr 1971). MacArthur was the first to indicate that species diversity could be corre- lated with habitat diversity (MacArthur and MacArthur 1961, MacArthur et al. 1962). Others have found a similar trend of increas- ing animal diversity with increasing habitat complexity. This trend has been seen for birds (Karr 1971, Karr and Roth 1971, Tomoff 1974, Willson 1974, Lancaster and Rees 1979, Beedy 1981), lizards (Pianka 1966), rodents (Rosenzweig and Winakur 1969, Feldhamer 1979, Pizzimenti and De Salle 1981), and spi- ders (Hatley and MacMahon 1980). By far the greatest amount of literature on this topic deals with the relationship between breeding bird communities and habitat com- plexity. This is the first study to consider (1) the relationship between vegetation complex- ity and postbreeding bird assemblages and (2) to consider more than one vertebrate class in an area. This allows us to ask several questions about species diversity and habitat complex- ity. Do postbreeding assemblages of birds conform to the pattern of increasing diversity with increasing habitat complexity seen for many breeding bird assemblages? Do diversi- ties of several major taxa in the same habitats respond in the same way to vegetation struc- ture? If measures of species diversity do not correlate with vegetation structure, are other measures of the relationship between a taxon and habitat more meaningful and predictive? METHODS Study Area The study area is in the Escalante Desert of Utah, in the southeastern portion of the Great Basin (Fig. 1). We set up four 1,000 m tran- sects in each of the five habitats that are the dominant vegetation types in this area. These habitats were uniform areas of pinyon/ juniper, sagebrush, greasewood/shadscale, grassland, and an area we termed mixed shrub because it was a heterogeneous mix of small shrubs and grasses different from the other four habitats. These habitats generally fol- lowed an elevational gradient from approxi- mately 1,550 to 1,785 m, with greasewood/ Department of Biology, Museum of Southwestern Biology, University of New Mexico, Albuquerque, New Mexico 87131. School of Dentistry, University of California, Los Angeles, California 90024. 711 iS 712 GREAT BASIN NATURALIST Vol. 46, No. MILLARD CO. BEAVER CO. MILFORD STUDY AREA MN 7, IRON CO. RIZONA Fig. 1. The study site in southwestern Utah in the Escalante Desert. Four 1,000 m transects were established in each of five Great Basin habitats within the shaded area. October 1986 shadscale in the valley bottom and pinyon/ju- niper woodland on the foothills of the moun- tain range bordering this valley. The other habitats were at intermediate elevations. This area is characterized by hot summer tempera- tures and cold winters. Annual rainfall aver- ages approximately 200 mm, with precipita- tion falling in all months. The highest amount of precipitation falls in March and April, ap- proximately 50 mm, 23% of the total. Each of the other months averages about 15 mm of precipitation, approximately 7% of total. Field Methods Five experienced investigators carried out censuses of rodents, lizards, and birds in July and August 1981. We used visual walking cen- suses to determine the densities of lizards and birds. We censused birds between 0530 and 0800 and lizards between 0900 and 1200. We recorded the species and number of animals sighted, distance of each observation from the transect line, and the compass direction of each observation. This information was en- tered into a computer program (Burnham et al. 1980) that gave a density estimate for each species. This method of line-transect sam- pling takes into account differential visibility of individual animals in different habitats. Where cover is more dense, the effective dis- tance of sighting a bird or lizard is reduced. This, in turn, reduces the width of the area censused on either side of the transect line. A smaller belt of area on either side of the tran- sect line gives a smaller area sampled for the number of observations and therefore cor- rects for decreased visibility. This computer program generates a different size of area cen- sused for each species and for each habitat. Density estimates were, therefore, deter- mined with visibility being an integral part of that estimate. Rodents were live-trapped at night using the assessment line technique (O Farrell et al. 1977) to determine rodent density. This tech- nique also includes the movement behavior of the animal at the time of censusing in making density estimates. The assessment lines are trapping stations located perpendicular to the two main parallel lines of trapping stations. The assessment lines give a maximum boundary around the main census lines for each species by recording the farthest dis- tance individuals of a species are caught from GERMANO, LAWHEAD: GREAT BASIN ECOLOGY 713 the main census lines. The length of the main census lines multiplied by the width, as deter- mined by trapping on the assessment lines, gives an estimate of the area censused for each species. Vegetation sampling was done using the Daubenmire Nested Quadrat method (Mueller-Dombois and Ellenberg 1974) on the same transects used to census animals. Sampling yielded plant species abundance, percent density, and percent cover. Data Analysis Species diversity and evenness values were calculated from the richness and abundance data using indices from Hill (1973). These in- dices define diversity as N, = 1/2(p;), where p, is the relative abundance of the i™ species, and evenness as N,/N,, with N, = exp(— =p, In p;). The diversity index (N,) expresses diver- sity with “species” as the basic unit and still includes an evenness component of species abundance pattern as well as richness. The diversity value calculated by this index is in- fluenced more by the number and abundance of common species than rare species, al- though both are included in determining a diversity value. Hill (1973) points out that N, allows a more straightforward comparison among communities with different diversities and sample sizes (see Rotenberry 1978 for a summary of the advantages of using N, as a diversity index). For vegetation we calculated indices for both horizontal and vertical heterogeneity. We determined horizontal heterogeneity us- ing a habitat physiognomic complexity index (PCI) for each habitat type similar to that of Tomoff (1974). We determined this diversity index for each habitat again using the index N,, where p, equals the proportional cover value of each physiognomic component in the habitat (i.e., grass, cacti, forbs, shrubs, and trees). A habitat with only one or two of these components composing the majority of cover, or being the only components in the habitat, is not likely to have as much horizontal hetero- geneity as a habitat that contains a somewhat equal mixture of components. We deter- mined vertical heterogeneity using the Shan- non-Weaver Information index, HX = —Xp, In p; to give foliage height diversity (FHD) where p, is the proportion of the total cover of the foliage that lies in the i” vertical layer 714 GREAT BASIN NATURALIST Vol. 46, No. 4 TABLE 1. Cover values (percent total cover) of physiognomic components and diversity indices for each habitat type. Habitats are listed in order of elevation from lowest to highest. Physiognomic component Habitat Forb Grass Cactus Greasewood/ 0.01 0.04 0 Shadscale Mixed shrub 0.28 14.62 0.08 Grassland 0.48 14.06 0 Sagebrush 0.03 5.34 0.21 Pinyon/Juniper 0.31 LL, 0.02 Diversity index Shrub Tree PCI FHD 28.18 0 1.00 0.06 17.08 0 2.03 0 12.97 0 2.07 0 22.49 0 1.47 0.50 3.95 22.02 1.50 0.61 TABLE 2. Species richness, abundance (number per hectare), evenness, and diversity of the three taxa for a gradient of habitats. The habitat types are listed in the order of their PCI value from highest to lowest. PCI values for each habitat are listed in parentheses below the habitat type. Mixed Grassland shrub Taxon (2.07) (2.03) LIZARDS Species richness 5 2 Abundance 27.87 22.12 Evenness .69 .94 Diversity 2.02 1.03 RODENTS Species richness 6 5 Abundance 2.66 6.22 Evenness 84 .93 Diversity 3.24 2.50 BIRDS Species richness 3 6 Abundance 8.96 1.95 Evenness .94 85 Diversity 1.92 3.39 (MacArthur and MacArthur 1961). Vegetation was divided into layers of 0 to 0.5 m, 0.5 to 1.0 m, and > 1.0 m. We used correlation and regression statistics to find relationships be- tween PCI, FHD, and animal diversity. RESULTS Values of PCI for each habitat type were highest in the two habitats with abundant grass cover and lowest in the greasewood/ shadscale, where virtually all the cover was composed of similar-height shrubs (Table 1). The grassland and mixed shrub habitats had highest PCI values by virtue of having an even mix of two physiognomic components, whereas the other three habitats were domi- nated by only one physiognomic component. The pinyon/juniper habitat contained all five vegetation components, but only the tree cat- Habitat Type Pinyon/ Greasewood/ Juniper Sagebrush Shadscale (1.50) (1.47) (1.00) 0 5 3 4.25 8.69 4.43 38 wo .86 Meal 3.05 BOT bs) 9 5 18.32 8.01 11.36 .96 74 mo 2.80 2.62 2.04 25 6 7 3.19 0.22 1.21 .70 1.00 83 8.67 4.79 Seal egory gave a significant cover value. Cover densities of forbs and cacti were low in all five habitats. Trees were present only in the pinyon/juniper habitat. However, values of FHD gave a different trend. The grassland and mixed shrub habitats, which gave the highest values for PCI, had values of 0 for FHD. Not too surprisingly, the highest value for FHD was for pinyon/juniper habitat. For each animal taxon, the highest diversity indices occurred in different habitat types (Table 2). Birds showed the widest range in diversity values, with a high of 8.67 in pinyon/ juniper habitat and a low of 1.92 in grassland habitat. Diversities of rodents were the most similar, with a range of 2.04 to 3.24. Species richness was highest in the pinyon/ juniper for lizards and birds, but highest in the sagebrush for rodents. Birds also showed the widest range in species richness, with val- October 1986 9.0 @ 7.0 5.0 @ @ 3641 Ww a) ry Dy < > 1.0 DIVERSITY o ro) \ 2.0 PG | Fig. 2. Diversity values for birds (circles), rodents (tri- angles), and lizards (squares) versus the physiognomic complexity index (PCI) for five Great Basin habitats. Al- though the regression lines are not significant at the 0.05 level, trends are evident for lizards and rodents. ues from 3 to 25. Abundances were highest in the pinyon/juniper for rodents, but for birds and lizards they were highest in areas with abundant grass cover (Table 2, Appendices A, B, C). Although there is not a clear pattern, GERMANO, LAWHEAD: GREAT BASIN ECOLOGY 715 9.0 7.0 5.0 A Y=2.59+7.75X r=.88 tJ SS) P<.05 a] tO > > te te ae WW A > a 10 3.0 - B a 1.0 0.1 0.3 0.5 0.7 F H D Fig. 3. Diversity values for birds (circles), rodents (tri- angles), and lizards (squares) versus foliage height diver- sity (FHD) for five Great Basin habitats. Lizards and rodents are not correlated to FHD, but postbreeding birds are significantly correlated. the areas with the lowest species richness pro- duced the highest, or nearly the highest, abundance for each taxon. Evenness values were relatively high for birds and rodents in all habitats, ranging from 0.70 to 1.00 for birds 716 1.50 q uJ oO a qx © 1.00 za = @a 10.0 wo A. a § Yi=0.37+13.89X r=.93 Fe 1.0 1.5 2.0 Cr Fig. 4. The logarithmic transformation of lizard abundance (number per hectare) versus the physiognomic complex- ity index (PCI) for five Great Basin habitats on the left and the combined abundance (number per hectare) of rodents and lizards versus PCI for the same habitats on the right. Both regression lines are significant. and 0.72 to 0.96 for rodents. For lizards, an unusually low evenness value of 0.38 was ob- tained for the pinyon/juniper habitat. No statistically significant relationships came from plotting diversity indices for each taxon against PCI values for the five habitat types (Fig. 2), although rodent diversity was highly correlated to PCI and tended to in- crease with increasing PCI, and lizard diver- sity tended to decrease with increasing PCI. Bird diversity was positively correlated to FHD); lizard and rodent diversities were un- correlated (Fig. 3). Although the relationships of diversity to horizontal habitat heterogene- ity for rodents and lizards are suggestive, nei- ther is statistically significant and therefore not wholly satisfying. We did find, however, that a component of diversity, abundance, was related to PCI in some instances. For lizards the logarithmic regression of abun- dance to PCI was significant and highly corre- lated, as was the regression of combined abundance for rodents and reptiles plotted against PCI (Fig. 4). We looked at the above relationship more closely and found that lizard abundance in- creased with increasing grass cover. There is a significant (P < .05) negative relationship be- tween percent grass cover and percent shrub and tree cover (r = —.96). As grass cover increased, there was a linear decline in over- story cover. Reptile abundance plotted against the ratio of percent grass cover over the percent shrub and tree cover gave a signif- icant (P < .01) logarithmic relationship (r = .98, Fig. 5). As grass cover increased and overstory cover dropped, reptile abundance increased. We also found a significant (P < .05) inverse relationship between the loga- rithmic transformation of rodent abundance and the grass/overstory ratio (r = —.91 Fig. 5.) Rodent abundance decreased with increasing grass cover and increased with increasing shrub and tree cover. No pattern existed for bird abundances when plotted against the ra- tio of percent grass cover to percent shrub and tree cover. DISCUSSION In this part of the Great Basin, both lizard and rodent assemblages seem to be struc- tured, at least in part, by the horizontal het- erogeneity of the vegetation. Postbreeding bird assemblages are correlated with vertical heterogeneity. For lizards there was a trend of decreasing diversity with increasing vegetation complex- ity. This trend is in contrast to the positive relationship Pianka (1966) found between the number of lizard species and plant volume diversity. Comparisons with this study are weak, however, because Pianka used species richness as his measure of animal diversity, October 1986 1.50 uJ a (S) za ) fua} eocoooroqoorrocwrcocRnNOCCS I a I I I ON fo, og al Foe Ita I mae femal Fo OONFORNAOOCHOF SO y= LW ODS ~——=~—_—“/__— a Data from bummed site are in parentheses; otherwise, data are from unburned site. Many of the moths taken at the burn site were Euxoa spp. The larvae, called cutworms, have a wide host range yet are rarely encountered be- cause they inhabit soil. Agromyzids were common at both sites (Table 1). They are an important component of range- land because their larvae mine leaves and stems of grasses, forbs, and shrubs. Yet, agromyzids were at the burn site even though vegetation was poorly developed when the flies were collected. Parasitic hymenopterans were regularly col- lected at both sites. Chalcids were never abun- dant, yet they represented ca 30 species in eight families. Mutillids, external parasites of larvae and pupae of various wasps and bees, were col- lected more often at the burned site. Later in the season, braconids and ichneumonids also were more common at the burned site; they parasitize caterpillars, beetle and sawfly larvae, maggots, various bugs, aphids, spiders, and other wasps. Other hymenopterans more abundant in the burned site were pompilids, which are spider- hunting wasps, and sphecids, which are preda- tors of aphids, bugs, grasshoppers, planthop- pers, leafhoppers, flies, caterpillars, beetles, bees, and spiders. Adults of all these ento- mophagous wasps are attracted to flowers. Flow- ers, prey, and potential hosts presumably were scarce at the burned site, yet wasps were com- mon there. Pipunculids were found only at the unburned site (Table 2). Their larvae are solitary internal parasites of nymphs and adults of Homoptera, particularly leafhoppers. Nevertheless, the burned site Malaise trap collections contained many potential hosts. Pipunculid biology, how- ever, is poorly known and factors other than food supply may have influenced the flies to avoid the burned area. These flies may be good indicators of undisturbed areas because they were consis- tently absent from the burned site. Cursorial insects were collected with pitfall traps. Although pitfall traps are limited in effec- tiveness (Greenslade 1964, Luff 1975), they have been successfully used to collect and compare surface arthropods from different sites (Fitcher 1941, Morrill 1975). The traps indicate that ants and ground-dwelling beetles survived the fire, probably escaping the heat by being below the ground surface. The fire did not destroy all or- ganic material, such as brome seeds, so that food resources were available for ants to maintain their colonies and for the polyphagous beetles. Other studies have verified that ground insect populations are unharmed by fire or changes in vegetative architecture. Rice (1932) collected more ants on burned prairie in Illinois than on nearby control sites. Removal of shrubs from a shrub-steppe site in Wyoming did not adversely affect the abundance of tenebrionids and cara- ] October 1986 bids (Parmenter and MacMahon 1984). The collection data suggested that A. immacu- lata is attracted to stressed environments be- cause specimens were only found in pitfall traps on the burned site. Many buprestid species are sensitive to smoke and heat, and the beetles may have been attracted by volatiles from burned winterfat, the host plant for the larvae. Further- more, larvae may survive by feeding on the roots of damaged plants. Pitfall traps commonly collected two other types of insects. Silphids may have entered pit- fall traps to feed on the dead bodies of other insects. No apparent reason explained why the omnivorous Jerusalem crickets, S. fuscus, were not collected at the unburned site. The present study raises several questions about the relationship between fire and the in- sect community. For example, why were so many predaceous and parasitic hymenopterans in the burned area (Table 3), especially when so few flowers and, presumably, potential hosts were present? Why did parasitic flies avoid the burned site that contained abundant potential hosts? Some groups, particularly leafhoppers and moths, apparently are attracted to burned areas, but unfortunately their means of orienta- tion are not well known. Although only adults were examined in this study, this is the main life stage at which many insects disperse into various habitats. Although fire is a common management tool for rangeland, this study raises important con- siderations of its use. Herbivorous insects seem very attracted to burned sites, yet their natural enemies, particularly parasitic flies, avoid those locations. Consequently, vegetative regrowth is highly susceptible to plant feeders and may be so severely stressed as to inhibit stand reestablish- ment. The abundance of insect herbivores also presents a danger to reseeding programs. Young vegetation is often highly susceptible to insect herbivory. Obviously, more research is needed to determine the long-term effects of range fires on insect and plant communities. ACKNOWLEDGMENTS Appreciation is extended to: D. C. Nielson, C. L. Nowak, and S. F. Parker for their assis- tance; W. J. Hanson, G. E. Bohart, and F. D. Parker for identifying specimens; and USDI, Bureau of Land Management for use of study sites. This research was the result of cooperative HANSEN: INSECT ECOLOGY 127 investigations of the USDA-ARS and the Utah Agricultural Experiment Station, Logan, Utah 84322. Approved as Journal Paper No. 3027. LITERATURE CITED ALLAN, J. D., H. J. ALEXANDER, AND R. GREENBERG. 1975. Fo- liage arthropod communities of crop and fallow fields. Decologia (Berl.) 22: 49-56. CaNCELADO, R., AND T. R. YONKE. 1970. Effect of prairie burn- ing on insect populations. J. Kansas Entomol. Soc. 43: 274-281. EVANS, E. W., R. A. ROGERS, AND D. J. OPFERMANN. 1983. Sam- pling grasshoppers (Orthoptera: Acrididae) on burned and unburned tallgrass prairie: night trapping vs. sweeping. Environ. Entomol. 12: 1449-1454. Evans, F. C., AND W. W. Murpocu. 1968. Taxonomic compo- sition, trophic structure, and seasonal occurrence in a grassland insect community. J. Anim. Ecol. 37: 259-273. Evans, F. C., AND D. F. OWEN. 1965. Measuring insect flight activity with a Malaise trap. Pap. Michigan Acad. Sci., Arts, and Letters 50: 89-94. GREENSLADE, P. J. M. 1964. Pitfall trapping as a method for studying populations of Carabidae (Coleop-tera). J. Anim. Ecol. 33: 301-310. HAWKINS, B. A., ANDE. A. Cross. 1982. Patterns of refaunation of reclaimed strip mine spoils by nonterricolous arthropods, Environ. Entomol. 11: 762-775. Hewitt, G. B., AND W. H. BURLESON. 1976. An inventory of arthropods from three rangeland sites in central Mon- tana. J. Range Manage. 29: 232-237. JANZEN, D. H., AND T. W. SCHOENER. 1968. Differences in insect abundance and diversity between wetter and drier sites during a tropical dry season. Ecology 49: 96-110. Knutson, H., AND J. B. CAMPBELL. 1976. Relationships of grasshoppers (Acrididae) to burning, grazing, and range sites of native tallgrass prairie in Kansas. Proc. Tall Timbers Conf. Ecol. Anim. Control Habitat Man- age. 6: 107-120. Lurr, M. L. 1975. Some features influencing the efficiency of pitfall traps. Oecologia (Berl.) 19: 345-357. MATTHEWS, R. W., AND J. R. MaTrHEws. 1971. The malaise trap: its utility and potential for sampling insect popu- lations. Michigan Entomol. 4: 117-122. Morri, W. L. 1975. Plastic pitfall trap. Environ. Entomol. 4: 596. Murpocy, W. W., F. C. EVANS, AND C. H. PETERSON. 1972. Diversity and pattern in plants and insects. Ecology 53: 819-828. NaGEL, H. G. 1973. Effect of spring prairie burning on herbiv- orous and non-herbivorous arthropod populations. J. Kansas Entomol. Soc. 46: 485-496. PARMENTER, R. R., AND J. A. MACManon. 1984. Factors influ- encing the distribution and abundance of ground- dwelling beetles (Coleoptera) in a shrub-steppe ecosystem: the role of shrub architecture. Pedobiolo- gia 26: 21-34. Rice, L. A. 1932. Effect of fire on the prairie animal communi- ties. Ecology 13: 392-401. SEASTEDT, T. R. 1984. Microarthropods of burned and un- burned tallgrass prairie. J. Kansas Entomol. Soc. 57: 468-476. Tuomas, D. B., AND F. G. WERNER. 1981. Grass feeding insects of the western range: an annotated checklist. Univ. Arizona Tech. Bull. 243: 1-50. SIZE, STRUCTURE, AND HABITAT CHARACTERISTICS OF POPULATIONS OF BRAYA HUMILIS VAR. HUMILIS (BRASSICACEAE): AN ALPINE DISJUNCT FROM COLORADO Elizabeth E. Neely’ and Alan T. Carpenter” ABSTRACT.—Size, structure, and habitat characteristics were studied in three populations of Braya humilis var. humilis (C. A. Meyer) Robins. in Gray & Wats. (Brassicaceae), a small, herbaceous perennial of the alpine tundra in central Colorado. There was a significant association between numbers of reproductive, juvenile, and seedling individuals and population location. Plant size within reproductive, juvenile, and seedling size classes varied signifi- cantly among three populations. Plots containing Braya had significantly lower total plant cover, a different set of dominant plant species, more rock, bare ground, and less litter than plots without Braya. Braya appears to be restricted to calcareous substrates that experience a moderate level of disturbance, such as solifluction lobes and abandoned roads. Populations are small despite the existence of much potential habitat. Population studies are necessary for active conservation management of Braya. Braya humilis var. humilis (C. A. Meyer) Robins. in Gray & Wats. (Brassicaceae) is a small, herbaceous perennial that occurs in the alpine tundra of the Rocky Mountains in cen- tral Colorado. It is a rare taxon, disjunct from its nearest relatives in Canada by approxi- mately 1,600 km. The isolated populations in Colorado were previously treated by Rollins (1953) as B. humilis ssp. ventosa. However, recent monographic work by Harris (1985), based on greenhouse and common garden studies, indicates that the Colorado plants should be treated with B. humilis var. hu- milis. In North America Braya humilis var. humilis occurs from Alaska, south through the northern Rocky Mountains to Alberta and British Columbia, north through the western Canadian Arctic Archipelago, east to Green- land, Newfoundland, Anticosti Island, Ver- mont, and the north shore of Lake Superior (Harris 1985). Colorado populations of B. hu- milis may represent isolated relicts left behind on small areas of calcareous alpine habitat as glaciers retreated about 12,000 years before present (Harris 1985). Hereinafter, Braya will refer to Braya humilis var. humilis. In Colorado, Braya is restricted to cal- careous soils derived from Paleozoic rock for- mations such as the Mississipian Leadville Limestone and Ordovician Manitou Dolomite (Tweto 1974). The plant commonly grows in 1303 E. Plum, Fort Collins, Colorado 80524. association with Dryas octopetala, Carex ru- pestris, and Kobresia myosuroides on exposed slopes without late-lying snowbanks. It is of- ten found growing in solifluction lobes, on low-angle scree slopes, and on gravel with minor amounts of soil movement, but it also grows on man-made disturbances such as old mining roads and prospecis. In Colorado, Braya is known to exist only in 19 small isolated populations at 12 general locations in the Mosquito, Ten Mile, Elk, and Collegiate ranges. At nearly all of its known occurrences in Colorado, Braya populations are small despite the existence of much appar- ent potential habitat. Approximately 3,900 in- dividuals exist in Colorado, based on esti- mates for each known population. It is a taxon of special concern in Colorado (O'Kane 1986) and is currently a candidate for listing (Cate- gory 2) by the U.S. Fish and Wildlife Service under the Endangered Species Act (Fay 1985). Counts of one population west of Hoosier Pass, Colorado, have varied greatly, suggest- ing considerable year-to-year fluctuation in numbers (Harmon 1980, Johnston 1984, Neely 1985). Unfortunately, the accuracy of these counts is questionable because the plants are small and easily overlooked. Accu- rate counts are important for determining the size and dynamics of populations and are fun- Department of Range Science, Colorado State University, Fort Collins, Colorado 80523. 728 October 1986 BUENA VISTA NEELY, CARPENTER: COLORADO BRAYA 729 COLORADO SPRINGS Fig. 1. Locations of the three study sites in central Colorado. 1 = Mt. Bross, 2 = West Hoosier, and 3 = Spout Lake. damental to the conservation management of the taxon (Bradshaw and Doody 1978). Relatively few population studies have been undertaken on long-lived perennials, particularly those that are rare. There is in- creasing recognition by plant conservationists of the need to monitor rare plant populations to determine population structure, flux, and modes of population regulation (Bradshaw and Doody 1978, Whitson and Massey 1981, Kruckeberg and Rabinowitz, unpublished manuscript). We initiated in 1985 a long-term study of Braya population dynamics, selecting three populations in locations that span a gradient from minimal to substantial human distur- bance. First-year objectives were to deter- mine if significant variation among and within populations existed for (a) numbers of plants in three size classes and (b) size characteristics of plants. Additional objectives were to deter- mine if vegetation and substrate characteris- tics at one of the locations differed between (c) microsites containing Braya and those that did not, and (d) microsites containing Braya on and off an old vehicle track. METHODS Three populations were chosen for study (Fig. 1, Table 1). Both the Mt. Bross and West Hoosier sites have experienced great histori- cal mining activity and are dissected by old roads and prospects. Mt. Bross is particularly important because it has the largest known population, with an estimated 1, 160 individu- als (Johnston 1984). The West Hoosier site is significant because it has the second largest population (approximately 430 individuals). Part of the West Hoosier site is owned and managed as a preserve by The Nature Conser- vancy. The Spout Lake population, the only 730 GREAT BASIN NATURALIST Vol. 46, No. 4 TaBLE 1. General location descriptions for three Braya humilis var. humilis populations in central Colorado. Descriptor Mt. Bross County Park Latitude/longitude 39°19’N, 106°06’E Mountain range Mosquito Elevation (m) 3758 Aspect East-southeast Slope (degrees) 0-30 Human disturbance High Dominant vascular plant species Poa rupicola, Draba aurea, Arenaria obtusiloba record from the Collegiate Range, consists of approximately 210 individuals and has the largest known population on a site with no evident human disturbance. Monitoring Thirty-two permanent, l-m’ plots were es- tablished at the three locations in August 1985 (22 plots at Hoosier, 4 plots at Mt. Bross, and 6 plots at Spout Lake). Alternate corners of the plots were marked with iron spikes 25 cm long. An iron nail was driven into the ground adjacent to each Braya plant; an aluminum tag was attached to each nail. Plant coordinates within the plots were recorded on data forms to facilitate relocation of individual plants in future years. Measurements of rosette diame- ter and height and counts of numbers of stems, leaves, flowers and fruits were recorded for all Braya individuals within the plots. Sampling methodology was adapted from that used by D. W. Inouye and M. B. Cruzan (personal communication) in central Colorado and was conducted during 10 to 15 August 1985. Plants were separated into three size classes based on their development; ages could not be determined. Seedlings were de- fined as nonreproductive plants with five or fewer leaves. Juveniles were plants with more than five leaves but without current year’s flowers or fruits. Reproductive individuals had current year’s flowers or fruits present. G-tests were used to test for significant as- sociation between numbers of Braya plants in the three size classes and location. Plant size data were tested for normality using an al- gorithm in Minitab (Ryan et al. 1982) and were normalized using logarithmic transfor- Location West Hoosier Spout Lake Park/Summit Chaffee 39°22'N, 106°05'E 38°47'N, 106°25’E Ten Mile Collegiate 3695 3750 East-southeast North 20-30 30-40 High-low Low Dryas octopetala, Dryas octopetala, Carex rupestris, Carex rupestris, Kobresia myosuroides Kobresia myosuroides mation where appropriate. Data were sub- jected to analysis of variance (ANOVA) to de- termine if plant size characteristics differed significantly among locations. T-tests were used to assess statistical significance of differ- ences in plant size characteristics on and off the old vehicle tract (termed the cutoff road) within the West Hoosier population. Vegetation and Substrate Sampling Total cover of vascular plant species, cover by species, and cover of substrate components (rock, bare ground, and litter) were estimated to the nearest percent for 22 permanent 1-m? plots containing Braya and 22 randomly placed 1-m’ plots that did not contain Braya at the West Hoosier location. Similar data were collected for plots containing Braya on and off the cutoff road. Data were tested for normal- ity as outlined above, transformed as needed, and analyzed using ANOVA. T-tests were used to assess significance of differences in total vegetal cover, cover of dominant vascu- lar plant species, and substrate cover between plots (a) containing or not containing Braya and (b) on or off the cutoff road. RESULTS Size Structure of Populations Size structure of Braya populations differed among the three locations with a highly signif- icant association between the number of plants in each size class and location (Table 2). Mt. Bross had the largest proportion of seedlings, whereas West Hoosier had the low- est. West Hoosier had the highest proportion of reproductive individuals. Spout Lake had the same proportion of juveniles as did West October 1986 NEELY, CARPENTER: COLORADO BRAYA 731 TaBLE 2. Number (and percentage) of Braya plants in three size classes at three locations in central Colorado. Reproductive class includes individuals with current year’s flowers or fruits. Juvenile and seedling classes include individuals possessing no current years reproductive material and that have >5 leaves or 1—5 leaves per plant, respectively. Plant size class Mt. Bross Reproductive 95 (29%) Juvenile 72 (22%) Seedling 164 (49%) Total 331 Location West Hoosier Spout Lake 139 (47%) 24 (18%) 139 (47%) 63 (47%) 19 (6%) 47 (35%) 297 134 There is a highly significant association (P<0.005) between plant class and location (G-test, Sokal and Rohlf 1981). TABLE 3. Measures of Braya plant size and reproductive output in three size classes at three locations in central Colorado. Means +1 SEM. For definitions of plant classes, see Table 2. Different letters in parentheses within each row denote significantly different means (P<.05, least significant difference test, Zar 1974). Plant size class Mt. Bross REPRODUCTIVE Height (mm) 34.8 + 1.5(a # of stems/plant 4.4 + 0.5(a # of fruits/plant # of leaves/plant Rosette diameter (mm) 27.4 + 1.0(a JUVENILE # of leaves/plant 9.9 + 0.7(a) Rosette diameter (mm) 12.8 + 0.7(a) SEEDLING # of leaves/plant 4.1 + 0.1(a) Rosette diameter (mm) 5.5 + 0.2 Hoosier, but it had a much higher proportion of seedlings. The height, number of stems, fruits, and leaves per plant and rosette diameter of repro- ductive individuals were significantly differ- ent among the three locations (Table 3). Re- productive plants at Mt. Bross were consist- ently the largest and produced the most re- productive material. Spout Lake consistently had the smallest reproductive individuals; they also produced the smallest amount of reproductive material. The number of leaves per plant and rosette diameter of juvenile individuals were signifi- cantly different among the three locations (Table 3). The Spout Lake juvenile plants were the largest. There were significant dif- ferences in the number of leaves per seedling among sites, but not in rosette diameter. The proportions of plants in the three size classes were significantly different on and off the cutoff road at West Hoosier (Table 4). The plants on the road were mostly reproductive, whereas the plants off the road were mostly Location West Hoosier Spout Lake P-value 30.2 + 1.2(b) 20.7 + 1.6(c) <0.001 2.7 + 0.2(b) ee) <0.001 12.1 + 1.4(b) 6.7 + 1.2(c) <0.001 23.6 + 1.0(b) 16.8 2 1.5) <0.001 21.4 + 0.8(b) 20.4 + 1.1(b) <0.001 12.7 + 0.6(b) 13.1 + 0.8(b) <0.001 10.8 + 0.4(b) 13.4 + 0.6(a) <0.006 4.9 + 0.1(b) 35 + 0.1(c) <0.001 5.9 + 0.6 6.2 + 0.4 0.178 juveniles. Although the proportion of seed- lings on the road was twice that off the road, seedlings were scarce in both areas. The num- ber of leaves per plant and the diameter of reproductive individuals, as well as the rosette diameter of juveniles, were signifi- cantly greater on the road (Table 5). Vegetal and Substrate Cover Total vascular plant cover and cover of Dryas octopetala, Carex rupestris, and Ko- bresia myosuroides were significantly lower on the cutoff road at West Hoosier (Table 6). Plots on the road had significantly greater cover of rock but significantly less litter cover than plots off the road. There was little evi- dence of disturbance off the road. Plots containing Braya had significantly less total vascular plant cover and cover of Dryas octopetala, Kobresia myosuroides, and Polygonum viviparum than plots lacking Braya (Table 7). The rank order of the three plant species with the greatest cover was dif- ferent in Braya plots compared to plots lack- 732 TABLE 4. Number (and percentage) of Braya plants in three size classes on and off the cutoff road at the West Hoosier location in central Colorado. For definitions of plant classes, see Table 2. Location Plant size class On cutoff road Off cutoff road Reproductive 61 (59%) 78 (40%) Juvenile 39 (37%) 100 (52%) Seedling 4 ( 4%) 15 ( 8%) Total 104 193 There is a highly significant association (P<0.005) between plant class and location (G-test, Sokal and Rohlf 1981). ing the plant. The substrate of Braya plots was characterized by significantly greater rock and bare ground but lower litter cover. DISCUSSION Size Structure and Populations The different proportions of reproductive, juvenile, and seedling individuals that were observed at the three locations could arise from differences in recruitment, survival, or growth of Braya plants. Recruitment of seedlings in harsh environments such as the alpine is probably episodic for many plant species (Billings 1974, Jolls 1982). Factors af- fecting recruitment could vary substantially over distances on the order of several kilome- ters. Thus, Braya seedling recruitment at the three sites could vary widely among and within years. The small size and large num- bers of seedlings at Mt. Bross may indicate more recent establishment at this location, with the few seedlings at West Hoosier result- ing from lack of recent recruitment. Survival of plants within all size classes could also vary greatly among locations and years. The combined effects of differential re- cruitment and survival could perhaps account for the large observed differences in propor- tions of size classes at the three locations. The variation in plant size structure among populations could arise from differential plant growth rates, caused by site quality differ- ences such as soil fertility, soil moisture, length of the growing season, or competition from other species. Difference in rosette di- ameter may be a phenotypic response to these varying site conditions. Very little is known about recruitment, sur- vival, and growth of Braya individuals. Thus, it is unknown how rapidly the present class GREAT BASIN NATURALIST Vol. 46, No. 4 structure could change, implications of which are important for the conservation of the taxon. For example, the West Hoosier popu- lation consists of reproductive and juvenile individuals with very few seedlings. This pop- ulation might be senescent with poor prospects for long-term survival. Alterna- tively, the present dearth of seedlings could simply reflect several recent years of poor seedling establishment, with former seed- lings moving into the juvenile and reproduc- tive classes. We presently lack the informa- tion necessary to make rational management prescriptions for Braya. Subpopulations only afew meters apart also had different size structures. The lower pro- portions of juvenile and seedling individuals on the cutoff road may have resulted from increased mortality in the smaller size classes caused by soil erosion on the road. Reproduc- tive individuals, which were nearly always larger than juveniles or seedlings, may have survived better because of their more exten- sive root systems. Harris (1985) has reported that Braya is octoploid (2n=56). Polyploidy may be an im- portant factor in Braya’s success on old roads. Polyploids are more likely to become adapted to environments disturbed by human activity than are their related diploids (Clegg and Brown 1983). Polyploidy helps to buffer against inbreeding depression and may en- hance wide environmental tolerance. Auto- gamy and polyploidy may help maintain the extremely isolated populations in Colorado (Harris 1985). Substrate and Plant Associates Braya microsites at West Hoosier are rocky with much bare ground, minimal litter, and sparse vegetal cover. Typical substrate con- sists of about 50% rock fragments 1-3 cm in length and about 50% bare ground. Our ob- servations of substrates at other Braya loca- tions were consistent with these findings. The low total vegetal cover on the cutoff road is consistent with the findings of Greller (1974), who found total plant cover on 40- to 50-year- old roadcut slopes in the alpine tundra of Rocky Mountain National Park to be less than half of the surrounding natural sites. Past and Present Disturbance Braya appears to be a pioneer species of | | { October 1986 NEELY, CARPENTER: COLORADO BRAYA 733 TABLE 5. Measures of Braya plant size and reproductive output in plots on and off the cutoff road at West Hoosier. Means + 1SEM. For definitions of plant classes, see Table 2. Location Plant size class On cutoff road Off cutoff road P-value REPRODUCTIVE Height (mm) 30.3 + 1.9 30.2 + 1.5 0.96 Number of stems/plant 3.0 + 0.4 2.4 0:3 0.17 Number of fruits/plant 13.9 + 2.3 1O7/ 22 16 0.24 Number of leaves/plant 26.9 + 1.7 *Al, I 32 ILO. 0.003 Rosette diameter (mm) SY G) se 11.8} 18.8 + 0.7 0.000 JUVENILE Number of leaves/plant 13.6 + 1.3 13,8) a= 0,7 0.35 Rosette diameter (mm) 13.2 + 0.9 10.0 + 0.5 0.001 SEEDLING Number of leaves/plant 4.8 + 0.2 5.@ s2 11,7 0.49 Rosette diameter (mm) Uo = (VY By ae (0,7 0.23 TABLE 6. Cover of dominant (>1% cover) vascular plant species and substrate components in Braya plots on and off of the cutoff road and plots off the cutoff road not containing Braya at West Hoosier. Means (%) + 1SEM. Location Braya plots Braya plots Plots with- on road off road out Braya P-value PLANT SPECIES Dryas octopetala 0.1 + 0.0 193,83 s= SO) S06) a2 HY 0.001 Carex rupestris 0.8 + 0.3 Grol ==253 Pe) 25 IO) 0.060 Kobresia myosuriodes 0.9 + 0.4 5.8 + 2.3 10.8 + 2.5 0.030 Erigeron pinnatisectus 2.0 + 0.6 1.8 + 0.6 2.8 + 0.8 0.056 Polygonum viviparum 13+ 0.3 DAD) = (0). {5) 45+ 0.8 0.556 Hymenoxys acaulis absent 2.2 + 0.6 1.7 + 0.6 — Calamagrostis purpurascens 1.0 + 0.6 1.8+0.5 2.4 + 0.8 0.483 Silene acaulis absent 143 23 1165 1.3 + 0.9 — Total vascular plants 9.8 + 1.2 39.8 + 5.3 Osa 33 Ext 0.000 SUBSTRATE COMPONENTS Rock 58.1 + 5.5 25.1 + 5.3 16.4 + 3.2 0.000 Bare ground 35.0 + 4.9 46.5 + 4.9 22.5 + 4.2 0.002 Litter 2.0+ 0.5 To®) 28 Weo4$ ae 492 0.000 TABLE 7. Cover of dominant (>1% cover) vascular plant species and substrate components in plots with and without Braya at West Hoosier. Means (%) + 1SEM. Location Plots with Braya Plots without Braya P-value PLANT SPECIES Dryas octopetala U8) 22 BO S08 23 SL 0.001 Carex rupestris 42+ 1.4 Po) as ILO 0.45 Kobresia myosuroides S98 2 ILS 10.8 + 2.5 0.02 Erigeron pinnatisectus 1.9 + 0.4 2.8 + 0.8 0.29 Polygonum viviparum 1.8 + 0.4 45+ 0.8 0.005 Hymenoxys acaulis 13+ 0.4 Ly ss OG 0.58 Calamagrostis purpurascens 14+ 0.5 2.4 + 0.8 0.29 Silene acaulis 10+0.9 8} 2s 0,9 0.86 Total vascular plants 27.5 + 4.5 70.7 + 5.7 0.000 SUBSTRATE COMPONENTS Rock 37.6 + 5.2 16.4 + 3.2 0.0015 Bare ground AON 22 BW DDS se ZN) 0.0011 Litter 5.4 + 1.0 W4 + 2:2 0.0000 734 disturbed areas. In some populations, only a few individuals have been found off these dis- turbances (E. E. Neely, personal observa- tion). Congeners grow on unstable substrates, such as scree slopes, gravel bars, shorelines, and solifluction lobes (Harris 1985). Many rare taxa in the western flora of North America and their common relatives colonize dis- turbed habitats (Stebbins 1980). Braya may inhabit unstable or disturbed areas because of an inability to compete with other species, as suggested by Griggs (1940) for other species of rare plants. Of the three populations, Mt. Bross plants appear to be the most vigorous, perhaps be- cause past disturbance has reduced the den- sity or size of other plants, leaving more re- sources available to Braya. The largest plants and those with the greatest amout of repro- ductive output at Mt. Bross occur mostly on the margins of a rough vehicle path and on spoil banks adjacent to a ditch. The path is level, and the surface is apparently stable. At West Hoosier, the cutoff road is considerably more disturbed than the adjacent areas. Possi- bly the degree of disturbance on the road is greater than optimum for Braya, given the virtual absence of seedlings and small propor- tion of juveniles. Observations of Braya in the Spout Lake population reinforce the importance of soil disturbance. Here it typically grows in small gravels, scree slopes, and solifluction lobes that have been demonstrated in Rocky Moun- tain National Park, Colorado, to move down- hill at a rate of 3-4 cm year ‘ (Benedict 1970). Braya appears to be preadapted to unstable substrates, making it most successful where there has been some moderate level of natural or man-made disturbance. The sizes of Braya populations before hu- man intervention began is unknown, but if populations at relatively undisturbed sites such as Spout Lake are any indication, popula- tions must have been small. In some cases human disturbance may simulate natural pro- cesses that create suitable habitat; however, drastic disturbances such as mine-related ac- tivities could greatly reduce or eliminate pop- ulations. Because Braya is found on cal- careous soils derived from rocks such as limestone, which are often highly mineral- ized, it may be threatened by potential min- ing activities. GREAT BASIN NATURALIST Vol. 46, No. 4 ACKNOWLEDGMENTS This research was conducted while the se- nior author was supported by the Colorado Field Office and the Rocky Mountain Her- itage Task Force of the The Nature Conser- vancy, the U.S. Fish and Wildlife Service, and the Colorado Native Plant Society. The authors thank J. G. Harris, M. K. Owens, J. S. Peterson, and L. M. Shultz for critically read- ing the manuscript and TNC Stewardship Committee, Colorado Chapter, volunteers for assistance with data collection. C. M. Warner drafted the figure. LITERATURE CITED BENEDICT, J. B. 1970. Downslope soil movement in the Colorado alpine region: rates, processes and cli- matic significance. Arctic Alp. Res. 2: 165-226. BILLINGs, W. D. 1974. Arctic and alpine vegetation: plant adaptations to cold summer climates. Pages 404—443 in J. D. Ives and R. G. Barry, eds., Arctic and alpine environments. Metheun, London. BRADSHAW, M. E., AND J. P. Doopy. 1978. Population studies and their relevance to nature conserva- tion. Biol. Conserv. 14: 223-242. CLEGG, M. T., AND A. H. D. Brown. 1983. The founding of plant populations. Pages 216-228 in C. M. Schonewald-Cox, S$. M. Chambers, and B. Mac- Bryde, eds., Genetics and conservation. Ben- jamin-Cummings Publ. Co., Inc., London. Fay, J. J. 1985. Endangered and threatened wildlife and plants; review of plant taxa for listing as endan- gered or threatened species; notice of review. Federal Register 50(188): 39526-39584. GRELLER, A. M. 1974. Vegetation of roadcut slopes in the tundra of Rocky Mountain National Park, Colo- rado. Biol. Conserv. 6: 84-93. Griccs, R. F. 1940. The ecology of rare plants. Bull. Torrey Bot. Club 67: 575-594. Harmon, W. 1980. Field data summary for B. humilis ssp. ventosa. Report prepared for the Colorado Natu- ral Areas Program, Department of Natural Re- sources, Denver, Colorado. Harris, J. G. 1985. A revision of the genus Braya (Cruci- fereae) in North Ameria. Unpublished disserta- tion, University of Alberta, Edmonton. JoHNSTON, B. 1984. Revised status report of Braya hu- milis ssp. ventosa. U.S. Forest Service, Region II, Denver, Colorado. Jouts, C. L. 1982. Plant population biology above timber- line: biotic selective pressures and plant reproduc- tive success. Pages 83-95 in J. C. Halfpenny, ed., Ecological studies in the Colorado alpine: a festschrift for John W. Marr. Occasional Paper 37. Institute of Arctic and Alpine Research, Boulder, Colorado. KRUCKEBERG, A. R., AND D. RABINOWITZ. Biological as- pects of endemism in higher plants. Unpublished manuscript. October 1986 NEELY, E. E. 1985. West Hoosier Braya preserve, Colo- rado, stewardship plan. The Nature Conservancy, Denver, Colorado. O’KANE, S. L. 1986. Plant species of special concern in Colorado. Report prepared for Colorado Natural Areas Program, Department of Natural Re- sources, Denver, Colorado. ROLLINS, R. C. 1953. Braya in Colorado. Rhodora 55: 109-116. Ryan, T. A., B. L. JOINER, AND B. F. Ryan. 1982. Minitab reference manual. Pennsylvania State University, University Park. SOKAL, R. R., AND J. F. ROHLF, 1981. Biometry, 2d. ed., W. H. Freeman Co., San Francisco. NEELY, CARPENTER: COLORADO BRAYA 735 STEBBINS, G. L. 1980. Rarity of plant species: a synthetic viewpoint. Rhodora 82: 77—86. TweETo, O. 1974. Geologic map of the Mt. Lincoln 15- minute quadrangle, Eagle, Lake, Park and Sum- mit counties, Colorado. USGS Misc. Field Stud., Map MF-556. Whitson, P. D., AND J. R. MAssEy. 1981. Information sys- tems for use in studying the population status of threatened and endangered plants. Pages 217- 236 in L. E. Morse and M. S. Henefin, eds., Rare plant conservation: geographical data organization. New York Botanical Garden, Bronx, New York. ZAR, J. H. 1974. Biostatistical analysis. Prentice Hall, En- glewood Cliffs, New Jersey. 620 pp. BIOGEOGRAPHIC ASPECTS OF LEECHES, MOLLUSKS, AND AMPHIBIANS IN THE INTERMOUNTAIN REGION Peter Hovingh’ ABSTRACT.—Some biogeographical and paleobiological aspects of leeches, mollusks, and amphibians in the Inter- mountain Region are reviewed. Areas of eastern Nevada and western Bonneville Basin as well as the tristate region of Nevada, Utah, and Idaho are poorly inventoried with respect to many aquatic-dependent animals. Observations of Batracobdella picta in the Wasatch Mountains and Erpobdella punctata in Tule Valley in the Bonneville Basin extends the western ranges of these leeches in the Great Basin. Life history and size of leeches varies among the study sites in the northern hemisphere. Aquatic mollusk species have diminished greatly in both prehistoric and historic times, as demonstrated by Utah Lake where some 30 species once lived. Eight genera survived into historic times, and perhaps only one species presently lives there. Extinction of numerous mollusks in the Bonneville Basin is still unknown with respect to cause and time. The finding of the Western Spotted Frog (Rana pretiosa) in Tule Valley reveals both a different habitat for this species when compared to other study sites and that this species must have occupied the region during Lake Bonneville times. With the exception of the Leopard Frog (Rana pipiens), most other amphibians probably migrated into the Bonneville Basin after the desiccation of Lake Bonneville. For eight years I have examined many ponds and springs to determine the distribu- tion of amphibians and their breeding habitat requirements in such rather diverse arid re- gions as the Bonneville Basin and the Colo- rado Plateau as well as the regions of high precipitation, such as the Wasatch and Uinta mountains. It soon became apparent that, with the exception of the threatened and en- dangered species, very little systematic work had been done on the distribution of native aquatic species in the Intermountain Region. Most of the work was done before 1940. To- day, with more and better roads and a very extensive inventory of the water resources, it seemed that new attempts should be made, especially in view of the recent efforts in un- derstanding the hydrological basins and their paleo-history. This paper reviews certain aspects of leeches, mollusks, and amphibians with the idea that with more information one might better understand their present distribution as well as their past distribution during the era of glaciers and the pluvial lakes. This paper is divided into three separate sections: (1) bio- geographical distribution and life history vari- ations of leeches, (2) review of mollusks in the Bonneville Basin, and (3) notes on the distri- bution of amphibians in Utah and Nevada. LEECHES With the exception of Herrmann’s work (1970) in Colorado, neither Nevada nor Utah have been methodically investigated for leeches. Twenty-one species were identified from Colorado (Klemm 1982, Herrmann 1970). Ten of these occur in western Colorado in the Middle Rocky Mountain Province and the Colorado Plateau Province (Herrmann 1970). Eight of these western Colorado spe- cies were found in Utah (Beck 1954, Barnes and Toole 1981) and four of them were found in Nevada (Klemm 1982). Table 1 lists the distribution. Note the lack of Erpobdellidae in Nevada. Observations of Placobdella ornata (Verrill 1872) in western Colorado, P. multilineata (Moore 1953) in the Uinta Basin of Utah, and P. parasitica (Say 1824) in Nevada need fur- ther clarification. Only in Nevada (Truckee River drainage) does the turtle host exist within this region. Theromyzon rude (Baird 1869) was found in both Nevada and Colorado and should be found in Utah. The above leeches will not be discussed further. Batracobdella picta (Verrill 1872) was re- ported from Current Creek, Wasatch County, Utah at 1,980 m elevation in a bog (Beck 1954). I found this leech very numerous on larval salamanders of Ambystoma tigrinum in ‘Department of Biochemistry, University of Utah, Salt Lake City, Utah. Correspondence should be sent to 721 Second Avenue, Salt Lake City, Utah 84103. 736 October 1986 HOVINGH: INTERMOUNTAIN BIOGEOGRAPHY 737 TABLE 1. Distribution of leeches in the Intermountain Region. Species Glossiphoniidae Batracobdella picta (Verrill, 1872) Glossiphonia complanata (Linnaeus, 1758) Helobdella stagnalis (Linnaeus, 1758) Placobdella ornata (Verrill, 1872) Placobdella multilineata (Moore, 1953) Placobdella parasitica (Say, 1824) Theromyzon rude (Baird, 1859) Hirudinidae Haemopis marmorata (Say, 1824) Erpobdellidae Dina dubia (Moore and Meyer, 1951) Dina parva (Moore, 1912) Erpobdella punctata (Leidy, 1870) Nephelopsis obscura (Verrill, 1872) Dog Lake (2,660 m elevation, Salt Lake County) and on breeding adults and larval salamanders in Red Pine Lake (2,680 m eleva- tion, Summit County, Weber River drainage) in the Wasatch Mountains. These two obser- vations extend the range of B. picta into the Great Basin. This leech was not found in 10 other salamander-inhabited ponds in the Wasatch Mountains (Provo, Weber, and Jor- dan River drainages) and two ponds of the Uinta Mountains (upper Duchesne River drainage). It was found between 2,062 and 3,224 m elevation in Colorado in 6 lentic wa- ter sources (Herrmann, 1970). The scattered distribution of B. picta could reflect both the distribution of the amphibian host as well as past mountain glacier distribution. Glossiphonia complanata (Linnaeus 1758) was found in the bench region of Utah County and at Deer Creek Dam in Wasatch County (Beck 1954). Beck (1954) found G. complanata and Helobdella stagnalis (Linnaeus 1758) in the “same general distribution of quiet pools of water or slowly moving shallow streams’. Glossiphonia complanata was found up to 3,610 m elevation in Colorado (Herrmann 1970). Helobdella stagnalis (Linnaeus 1758) was found in a stream near Laketown (Bear Lake), Utah, the bench region of Utah County, and in Utah Lake (Beck 1954, Tillman and Barnes 1973). In Colorado H. stagnalis was found between 1,000 and 3,200 m elevation (Herr- mann 1970). I found it feeding on Nephelopsis obscura (Verrill 1872) in the mountain ponds Western Colorado Utah Nevada 4. + + 7 + + = + + ? + + + + - + +: + - . + + + + of the Uinta Mountains (upper Duchesne River drainage, elevation 3,060 m). Tillman and Barnes (1973) showed that in- dividual Helobdella stagnalis (Linnaeus 1758) produced two broods of young during May and June in Utah Lake. This variation in life history is different from those studied in Canada (Davies and Reynoldson 1976) and the British Isles (Murphy and Learner 1982) (Fig. 1). At these latter locations, the adults died after producing young; in some locations two generations per year occurred and in other locations only one generation per year oc- curred. Water temperature may be a determi- nant of the two life history patterns in Canada (Davies and Reynoldson 1976). It is unknown what the determinants of H. stagnalis life his- tory in Utah Lake are, or if this life history variation is limited to Utah Lake. Nephelopsis obscura (Verrill 1872) is per- haps the most common leech in Uinta Moun- tain ponds and may be the top predator in the ponds that do not contain salamanders. In the Uinta Mountain ponds, I observed N. obscura at night with densities of four to six leeches per m’ of surface water in a pond that was at the most 1 m deep. Daytime observations were common in June and July, with densities less than one leech per m’ and rare in Septem- ber. Cocoons were deposited from early June to autumn, with the prevalent deposition oc- curring during late July. Growth patterns show, in late June, sizes between 0.01 and 0.85 g, with a group between 0.2 and 0.3 g (Fig. 2). During July, the large individuals 738 LIFE HISTORY VARIATIONS IN HELOBDELLA STAGNALIS British Columbia Alberta Utah Lake British Isles January Adult Adult Adult February j \ N March \ \ \ April Young \ N Ma \ \ \ \ Z \ Ngee \ Young J Adult acs ov SEI Wweiae woe ‘ Goa July \ \ 1 . XY August qe \ N \ September \ \ \ \ October \ \ \ \ November \ \ \ \ December Adult Adult Adult Adult LIFE HISTORY VARIATIONS IN NEPHELOPSIS OBSCURA Minnesota Alberta Uinta Mountains Spring VV Hatch Summer \ Autumn \ \ ve \ Winter \ \ \ \ Spring Bre 4y \ \ Hatch \ Hatch Summer \ \ \ \ Breed \ Autumn Breed BreedN \ \ Winter \ \ Spring meth Erced Summer \ Breed Autumn Winter Fig. 1. Life history patterns of Helobdella stagnalis in British Columbia and Alberta (Davies and Reynoldson 1976), British Isles (Murphy and Learner 1982), and Utah Lake, Utah (Tillman and Barnes 1973) (upper figure). Life history patterns of Nephelopsis obscura in Alberta (Davies and Everett 1977), Minnesota (Peterson 1983), and the Uinta Mountains, Utah (based on size alone) (lower figure). (larger than 0.5 g) die and can be seen in the bottom of the ponds. Growth continues to September. These data indicate that there is only one generation of young each year. One generation of young each year is similar to populations in Minnesota (Peterson 1983) and contrasts with the two-generation strategy in Alberta (Davies and Everett 1977) (Fig. 1). Sizes of leeches may not be an appropriate indicator of generation (Collins and Hohm- strand 1984a, b). Nephelopsis obscura reaches sizes of up to 1.2 g in the Uinta Mountains compared with populations in Alberta (0.41 g) and Minnesota (over 4.0 g) (Davies and Ev- erett 1977, Collins and Hohmstrand 1984). The scarceness of leeches less than 0.1 g in the Uinta Mountain ponds contrasts sharply with the abundance of leeches in this class size in Alberta. It was found in the bench regions of Utah and Salt Lake counties in freely running GREAT BASIN NATURALIST Vol. 46, No. 4 20 JUNE 22,1980 n=92 JulyeZoulsse@ n=6 20 PERCENTAGE OF SAMPLE O SEPTEMBER 6, IS80 20 n=27 500 WEIGHT (mg) 1000 Fig. 2. Weights of Nephelopsis obscura in a Uinta Mountain pond during the summer. The leeches were weighed to the nearest 10 mg. For comparison, consult with Davies and Everett (1977) and Peterson (1983). streams or in shallow ponds (Beck 1954). It has not been recorded in Utah Lake. It was found only in lentic habitats in Colorado between 1,650 and 3,200 m elevation (Herrmann 1970) and only in lotic habitats in Michigan (Klemm 1972). Reynoldson and Davies (1980) noted that N. obscura was rather sensitive and intol- erant to the changes in osmolarity of the aquatic medium. Erpobdella punctata (Leidy 1870) was October 1986 found in the bench region of Utah County and at 1,740 m on Cove Mountain, Sevier County (Beck 1954). It has been found in Utah Lake (Barnes and Toole 1981). I found it in Red Pine Lake (2,680 m elevation, Weber drain- age, Summit County) and in Solitude Lake (2,740 m elevation, Big Cottonwood Creek drainage, Salt Lake County) in the Wasatch Mountains and in five springs-wetlands in Tule (White) Valley (elevation 1,350 m, Mil- lard County). The aquatic systems in Tule Valley have been isolated from other Bon- neville Basin aquatic systems for some 13,000 years. In Tule Valley the specific conductance of the inhabited waters varied between 1,200 and 2,500 umhos/cm at 25 C, depending on the location in each wetland as well as the time of the year. Reynoldson and Davies (1980) noted that E. punctata was much more toler- ant to variations in osmolarity than Nephelop- sis obscura (Verrill 1872). Such tolerance might explain the presence of E. punctata in the interior of the Bonneville Basin, where it may exist as a relict species. Herrmann (1970) found E. punctata in both lentic and lotic habitats between 1,044 and 3,232 m elevation in Colorado. The leech was found in waters with pH variations of 6.3 to 10.3 (Herrmann 1970) and as low as pH 5.0 in Michigan (Klemm 1972). Davies et al. (1977) noted vari- ations in life history patterns with E. punctata populations in Alberta and that the differ- ences might be explained by interspecific competition with N. obscura. Two other Erpobdellidae, Dina dubia (Moore and Meyer 1951) and D. parva (Moore 1912), have been reported from Utah. Dina dubia was found in association with Nepholopsis obscura (Verrill 1872) in lotic habitats in the bench region of Utah County (Beck 1954). Dina parva was found in Utah Lake (Barnes and Toole 1982). Haemopis marmorata (Say 1824) was found in the bench region of Utah County, eastern slope of the Aquarius Plateau in Garfield County, and at Deer Creek Dam in Wasatch County (Beck 1954). It was found to tolerate the greatest total dissolved solids (2,807 mg/l) of all the other leeches and was found between 1,044 and 2,975 m elevation in Colorado (Herrmann 1970). In the Colorado mountains, Pennak (1968) described the semidrainage lakes. In these lakes certain fauna and flora form a character- HOVINGH: INTERMOUNTAIN BIOGEOGRAPHY 739 istic assemblage. The tiger salamander (Am- bystoma tigrinum) and the yellow pond lily (Nuphar polysepalum) are the characteristic animal and plant. Glossiphonia complanata, Nephelopsis obscura, and Helobdella stag- nalis occur together in these lakes (Pennak 1968). Other leeches found in semidrainage lakes include Batracobdella picta, Batracob- della phalera, Dina dubia, Erpobdella punc- tata, Haemopis marmorata, Haemopis kingi, and Theromyson rude (Herrmann 1970). The semidrainage ponds in the Uinta Mountains occur with the tiger salamander, yellow pond lily, N. obscura, and H. stagnalis. It would be interesting to study these ponds over the en- tire Uinta Mountains to determine how these ponds relate to the Colorado semidrainage ponds. During the glaciation some 25,000 years ago, all of these ponds and lakes were under ice fields and, thus, the ecology of the Uinta Mountain ponds and lakes may have evolved separately from the Colorado Rocky Mountain ponds and lakes. MOLLUSKS Of some 100 species of mollusks in the Great Basin, some 50 species are found along the Highway 89-91 axis from Idaho to Arizona. With only Fish Springs National Wildlife Refuge being extensively inventoried in the western Bonneville Basin, much of the Bon- neville Basin and eastern Nevada are still very fertile areas for studying the assemblage of molluscan species. The desiccation of the pluvial lakes left many “semifossils,” a name coined by early collectors who gathered empty shells from arid regions in the Bonneville Basin (Call 1884). Other collectors gathered shells from aquatic regions such as Utah Lake and Bear Lake and noted that they were fresh. In re- viewing the literature it is difficult to know what specimens were found living. Many mol- luscan species require the examination of the soft parts for identification, and soft parts were not collected. A major problem today is to determine what mollusks were present in the region be- fore Lake Bonneville, the distribution of mol- lusks with the rising and desiccating waters of Lake Bonneville, and the present distribution of mollusks in the numerous basins of the Bonneville Basin and adjacent basins of Ne- 740 vada. This information could tell us very much about the evolution of the present-day aquatic flora and fauna of the water sources in the numerous basins. Both Utah Lake and Bear Lake have an abundant record of mollusks. Some 30 species of mollusks have been identified from Utah Lake environs. The taxa and their references are: Unionidae: Anodonta oregonensis Lea 1838 (Chamberlin and Jones 1929, Jones 1940a), Anodonta nuttalliana Lea 1838 (Call 1884, Chamberlin and Jones 1929, Henderson 1931, Jones 1940a), Anodonta wahlametensis Lea 1838 (Chamberlin and Jones 1929, Jones 1940a). Sphaeriidae: Sphaerium pilsbryanum Sterki 1909 (Baily and Baily 1951-1952, Call 1884, Chamberlin and Jones 1929, Jones 1940a), Pisidium compressum Prime 1851 (Baily and Baily 1951-1952, Call 1884, Cham- berlin and Jones 1929, Jones 1940a), Pisidium casertanum Poli 1791 (Baily and Baily 1951-1952, Jones 1940a). Pisidium variabile Prime 1851 (Jones 1940a). Valvatidae: Valvata humeralis Say 1829 (Baily and Baily 1951-1952, Chamberlin and Jones 1929), Valvata utahensis Call 1884 (Baily and Baily 1951-1952, Bickel 1977, Call 1884, Chamberlin and Jones 1929, Taylor 1966, Jones 1940a). Hydrobiidae: Fluminicola fusca Haldeman 1847 (Baily and Baily 1951-1952, Call 1884, Chamberlin and Jones 1929, Jones 1940a), Fluminicola seminalis Hinds 1842 (Jones 1940a), Amnicola limosa Say 1817 (Baily and Baily 1951-1952, Chamberlin and Jones 1929, Henderson 1931, Jones 1940a), Fonteli- cella (Paludestrina) longinqua Gould 1855 (Jones 1940a), and Tyonia (Paludestrina) protea Gould 1855 (Jones 1940a). Lymnaeidae: Lymnaea stagnalis Say 1821 (Call 1884, Chamberlin and Jones 1929), Lymneus elodes Say 1821 (Baily and Baily 1951-1952, Chamberlin and Jones 1929), Fos- saria modicella Say 1825 (Chamberlin and Jones 1929, Jones 1940a), Fossaria obrussa Say 1825 (Chamberlin and Jones 1929, Jones 1940a), Fossaria parva Lea 1841 (Baily and Baily 1951-1952), Stagnicola utahensis Call 1884 (Baily and Baily 1951-1952, Bickel 1977, Chamberlin 1933, Chamberlin and Jones 1929, Jones 1940a), Stagnicola caperata Say 1929 (Baily and Baily 1951-1952), Stagnicola GREAT BASIN NATURALIST Vol. 46, No. 4 proxima Lea 1856 (Baily and Baily 1951- 1952), and Stagnicola hemphilla Baker 1934 (Baily and Baily 1951-1952). Physidae: Physella propinqua triticea Lea 1856 (Baily and Baily 1951-1952), Physella utahensis Clench 1925 (Baily and Baily 1951-1952, Barnes and Toole 1981, Bickel 1977, Chamberlin and Jones 1929, Jones 1940a). Planorbidae: Gyraulus similaris Baker 1917 (Baily and Baily 1951-1952), Gyraulus ver- micularis Gould 1847 (Chamberlin and Jones 1929, Jones 1940a), Helisoma (Carinifex) newberryi Lea 1858 (Baily and Baily 1951— 1952, Call 1884, Chamberlin and Jones 1929, Taylor 1966, Jones 1940a), Promenetus (Men- etus) exacuous Say 1821 (Baily and Baily 1951-1952, Chamberlin and Jones 1929), and Planorbella (Helisoma) trivolvis Say 1817 (Chamberlin and Jones 1929, Jones 1940a). Ancylidae: Ferrissia fragilis Tryon 1863 (Baily and Baily 1951-1952), and Ferrissia rivularis Say 1817 (Chamberlin and Jones 1929). Of these mollusks in Utah Lake only An- odonta (Henderson 1931), Pisidium compres- sum (Call 1884), Physella propinqua triticea (Baily and Baily 1951-1952), Physella utahen- sis (Barnes and Toole 1981, Bickel 1977), He- lisoma newberryi (Center for Health and En- vironmental Studies 1975, Call 1884), Fluminicola fusca (Call 1884), Valvata uta- hensis (Call 1884), Lymnaea stagnalis (Call 1884), and Stagnicola utahensis (Chamberlin 1933) have been found living in Utah Lake. Presently Physella utahensis may be the only living species in Utah Lake (Barnes and Toole 1981). The extinction of mollusks at Utah Lake and Bear Lake raises some questions. Was there a — general extinction of mollusks in the Great — Basin lakes during some specific period, or | did each species become extinct with species specific causes and with lake specific causes? | Did the rising waters of the prehistoric pluvial _ lakes cause any extinction of mollusks in the | lakes and isolated springs, or did the rising | waters distribute the isolated mollusks throughout the basin? Has there been any postpluvial evolution of mollusks? The Bear Lake molluscan assemblage | showed that Helisoma newberryi (the most common gastropod) was radiodated at 8270 | B.P. (from 1.2 m deep at the Willis Ranch October 1986 terrace, elevation 1,814 m), 7700 B.P. (0.3 m deep at the Lifton bar shoreline, elevation, 1,808 m), and 7880 B.P. (less than 1 m above Bear Lake shoreline, 1,806 m_ elevation) (Williams et al. 1962). These dates indicate that H. newberryi became extinct at a rather specific time and perhaps its extinction was related to the lowering of Bear Lake (Williams et al. 1962). A second report listed the age of H. newberryi and Sphaerium sp. at the Bear Lake shoreline at 12,000 B.P. (Smart 1963). This would suggest that the extinction of mol- lusks at Bear Lake included an assemblage of species. Unfortunately, the dates are in con- flict (fictitious results can result under several circumstances, see Keith and Anderson 1963, Riggs 1984, Rubin and Taylor 1963). It would be important to reexamine the ages of mol- lusks of Bear Lake and to examine the ages of mollusks at Utah Lake and other lakes in the Great Basin. Whereas many molluscan species have lim- ited present-day distribution, some of these species may have a widespread fossil distribu- tion. Stagnicola pilsbryi Hemphill, 1890, is an exception, with its distribution being limited to Fish Springs National Wildlife Refuge and having no fossil record (Russell 1971, Taylor et al. 1963). Presently S. pilsbryi is considered extinct. It is unknown if S. pilsbryi evolved at Fish Springs after the desiccation of Lake Bonneville (Russell 1971). More studies of molluscan biogeographic distribution and habitat requirements in the Intermountain Region are needed to under- stand the evolution of the aquatic systems in the Great Basin (Yen 1951, Taylor et al. 1963, Taylor 1960, Russell 1971, Murray 1970). AMPHIBIANS Fourteen species of amphibians occur in Nevada and in Utah, with 12 species common to both states (Linsdale 1940, Banta 1965, La Rivers 1942, Tanner 1931). Most of these am- phibians can be placed into one of two groups. The first group is postulated as arriving into the Intermountain Region from the south and is largely confined to the Colorado River drainage in Utah and Nevada (Tanner 1978). These species include Bufo cognatus, B. mi- croscaphus, B. punctatus, B. woodhousei, Hyla arenicolor, Rana onca, and R. fisheri. R. onca and R. fisheri may be part of the Rana pipiens “complex.” HOVINGH: INTERMOUNTAIN BIOGEOGRAPHY 741 Bufo woodhousei is the only amphibian ar- riving from the south that penetrates into the Bonnevillle Basin and extends along the axis of Interstate 15 to and including portions of the Snake River drainage of Idaho. In the Bonneville Basin, B. woodhousei is found in the Sevier River drainage and in the isolated Snake Valley Basin in western Utah (Tanner 1931). Another species, Scaphiopus inter- montanus also came from the south because there was not much suitable habitat during the pluvial times for this amphibian to breed (Hovingh et al. 1985). Scaphiopus intermon- tanus is not dependent upon water for migra- tion as is the other species in the first group. The second group is postulated as arriving from the north, from the east, or from the west. This second group includes Ambystoma tigrinum, Hyla regilla, Pseudacris triseriata, Bufo boreas, Rana pretiosa, and R. pipiens. If one were to ask if any amphibians occurred in the Intermountain Region during the plu- vial and glacial era some 25,000 years ago, this second group would have the most likely can- didates. The Leopard Frog (R. pipiens), being found in many isolated springs throughout the Bonneville Basin and in numerous basins of Nevada, in mountainous habitat, and adapt- ing to flood plains of the White River (Uintah County), is one species that most likely occu- pied the region during the pluvial times. The Western Toad (Bufo boreas) largely inhabits the northwest United States and oc- cupies areas in northern Nevada and the mountain regions of Utah. There are some recognized subspecies in central and southern Nevada (Linsdale 1940). The Western Toad has not been found along the Utah-Nevada border (in particular, the Snake and Spring valleys). This toad breeds in the valley floors in Nevada and up to 3,050 m elevation in the mountains of Utah. When the species breeds in the valley floors of Utah, there may be some site competition with B. woodhousei. It seems that if the Western Toad occupied the Bonneville Basin during the pluvial times, it would presently occupy the valleys of the Utah-Nevada border. The Tiger Salamander (Ambystoma ti- grinum) and the Chorus Frog (Pseudacris triseriata) are very common in the aquatic systems of the Uinta Mountains in regions that formerly were occupied by glaciers. The Tiger Salamander is also common in the ponds 742 and reservoirs along the Wasatch Front and in the Wasatch Mountain lakes (Tanner 1931). The Chorus Frog is less abundant along the lower elevations of the Wasatch Front (Tan- ner 1931) and is scarce in the high mountain lakes of the Central Wasatch Mountains. Nei- ther species has been collected in Nevada, although the salamander is found in both the Raft River and Pine Valley mountains. It would seem that if the Tiger Salamander and Chorus Frog were present during Lake Bon- neville times, their distribution would occur in regions of eastern Nevada and western Bonneville Basin (Spring and Snake valleys). Their limited distribution might be explained by (1) relict populations during the pluvial times with no opportunity for extending their range into the Bonneville Basin, or (2) migra- tion into Utah in postpluvial and postglacial times along the Uinta Mountains and fanning out via the Wasatch Mountains to the Pine Valley Mountains in the south and the Raft River Mountains in the north. In the cases of the Tiger Salamander and the Chorus Frog (two eastern amphibians), one could postulate that between Lake Bonneville and the alpine glaciers there was not any habitat for these amphibians. The Pacific Treefrog (Hyla regilla) is com- mon in California and Oregon and in isolated populations in both the Colorado River drainage of Nevada and southwestern Utah, in central Nevada, and in the Raft River Mountains of Utah (Tanner 1931, Reynolds and Stephens 1984). This particular frog may have extended its range eastward during the pluvial times and now in the postpluvial era remains in isolated pockets. The Western Spotted Frog (Rana pretiosa) occurs in many relict populations from near Juneau, Alaska, southward throughout Wash- ington, Oregon, British Columbia, Idaho, western Montana, the upper Humboldt River drainage in Nevada, Yellowstone National Park, and the Big Horn Mountains in Wyo- ming and in isolated pockets of the Bonneville Basin and drainage system in Utah (Dunlap 1977, Turner and Dumas 1972, Morris and Tanner 1967, Tanner 1931). The museum records of the University of California (Berke- ley), Brigham Young University (Provo), Uni- versity of Utah (Salt Lake City), and the Uni- versity of Michigan (Ann Arbor) show the distribution in Utah to include the Snake Val- GREAT BASIN NATURALIST Vol. 46, No. 4 ley and Deep Creek drainage in the western Bonneville Basin and the Mono Lake (Juab County), the San Pitch River (a tributary of the Sevier River), and numerous locations in Salt Lake, Utah, Summit, and Wasatch coun- ties in the eastern Bonneville Basin. Western Spotted Frogs have not been found in the drainage of Thousand Springs Creek in north- eastern Nevada and northwestern Utah, in the Raft River Mountain region, and in the main Sevier River drainage. Thus, from the records, the distribution of the Western Spot- ted Frog in the Bonneville Basin is several isolated populations. In 1980 I found the Western Spotted Frog in Tule (White) Valley in Millard County, Utah. Tule Valley lies between Snake Valley on the west and the Sevier drainage basin on the east. Tule Valley has been isolated from the Lake Bonneville aquatic system for 14,000 years, and highly aquatic species such as the Western Spotted Frog could only occur in Tule Valley if it also occurred in the Lake Bonneville environs (assuming no human in- tervention). Thus, one may assume that the Western Spotted Frog along with the Leop- ard Frog occupied the Bonneville Basin at the time of Lake Bonneville. In examining the Tule Valley populations, one finds the Western Spotted Frog has adapted to a more saline environment than that found in other parts of its distribution. The wetlands lie between 1,347 and 1,350 m elevation on the valley floor, and most have warm water sources (25-19 C). The total dis- solved solids varies in the springs from 1,000 | to 1,400 mg/l (Stephens 1977) and the specific conductance varies between 1,000 and 3,000 umhos/cm at 25 C, depending on the location in the wetlands and the time of year. In Tule | Valley, the Western Spotted Frog breeds in | the cold water portion (the most distal from _ the spring source) of the warm water springs. Although amphibians are considered ter- restrial animals, the arid regions often limit | populations from extending their ranges. The distribution of these amphibians may be very local, endemic, and relict as are the Tule Val- | ley populations of the Western Spotted Frogs. The distribution of amphibians in the Inter- mountain Region can readily be the result of the pluvial and glacial eras and the subse- quent desiccation. However, one must pres- ently be careful in noting the presence or October 1986 absence of relict populations. These popula- tions may be a result of disturbances by natu- ral predators or humans and human-associ- ated animals. For instance, the Tiger Sala- mander replaced a viable breeding population of Leopard Frogs in an Emigration Canyon spring (Salt Lake County) over a period of 20 years (1963-1983). Leopard Frogs are known to eliminate Western Spotted Frogs (Dumas 1966). Spadefoot Toad (Scaphiopus) tadpoles are thought to eliminate other amphibian tad- poles (Creusere and Whitford 1976). Intro- duced bass, sunfish, and bullfrogs are highly destructive of native amphibian populations (Turner 1962, Licht 1974). Human utilization of scarce desert waters for agriculture and domestic uses deprives the amphibians of necessary breeding habitat (Turner 1962, Morris and Tanner 1969). It should be noted that many amphibian populations along the Wasatch Front in Utah and Salt Lake counties no longer exist; even though the breeding habitat still remains, it is surrounded by hous- ing developments or is adjacent to new high- ways. Certainly more fieldwork is needed in the biogeography of amphibians in the Inter- mountain Region. Particular attention must be applied to the Utah-Nevada border, the Raft River Mountains and Thousand Spring Creek, Goose Creek, and other drainages of the Snake River in the tristate region of Idaho, Utah, and Nevada. Breeding habitats must be identified and characterized. ACKNOWLEDGMENTS I would like to thank Dr. Donald J. Klemm (U.S. Environmental Protection Agency, En- vironmental Monitoring and Support Labora- tory, Cincinnati, Ohio 45268) for the identifi- cation of B. picta, H. stagnalis, N. obscura and E. punctata. Without his cooperation the review of the leeches would not have been written. LITERATURE CITED ON LEECHES Barnes, J. R., AND T. W. TOOLE. 1981. Macroinvertebrate and zooplankton communities of Utah Lake: a re- view of the literature. Pages 101-105 in Utah Lake Monograph. Great Basin Naturalist Mem. 5. BECK, D. E. 1954. Ecological and distributional notes on some Utah Hirudinea. Proc. Utah Acad. Sci., Arts, and Let. 31: 73-78. HOVINGH: INTERMOUNTAIN BIOGEOGRAPHY 743 Co ..ins, H. L., AND L. L. HOHMSTRAND. 1984a. Indicators of sexual maturity in the leech, Nephelopsis ob- scura (Annelida: Hirudinea). Amer. Midl. Nat. 112: 91-94. . 1984b. Early life history and growth of Nephelop- sis obscura Verrill, 1872 (Pharyngobdellida: Er- pobdellidae). J. Freshwater Ecology 2: 549-554. Davies, R. W., AND R. P. EVERETT. 1977. The life history, growth, and age structure of Nephelopsis obscura Verrill, 1872 (Hirudinoidea) in Alberta. Canadian J. Zool. 55: 620-627. Davies, R. W., AND T. B.REYNOLDSON. 1976. A compari- son of the life-cycle of Helobdella stagnalis (Linn. 1758) (Hirudinoidea) in two different geographical areas in Canada. J. Anim. Ecol. 45: 457—470. Davies, R. W., T. B. REYNOLDSON, AND R. P. EVERETT. 1977. Reproductive strategies of Erpobdella punc- tata (Hirudinoidea) in two temporary ponds. Oikos 29: 313-319. HERRMANN, S. J. 1970. Systematics, distribution, and ecology of Colorado Hirudinea. Amer. Midl. Nat. 83: 1-37. KLEMM, D. J. 1972. The leeches (Annelida: Hirudinea) of Michigan. Michigan Academician 4: 405—444. —__.. 1982. Leeches (Annelida: Hirudinea) of North America. U.S. Environmental Protection Agency, EPA-600/3-82-025. 177 pp. Murpny, P. M., AND M. A. LEARNER. 1982. The life history and production of the leech Helobdella stagnalis (Hirudinea: Glocciphonidae) in the River Ely, South Wales. Freshwater Biology 12: 321-329. PENNAK, R. W. 1968. Colorado semidrainage mountain lakes. Limnol. Oceanogr. 14: 720-725. PETERSON, D. L. 1983. Life cycle and reproduction of Nephelopsis obscura Verrill (Hirudinea: Erphob- dellidae) in permanent ponds of northwestern Minnesota. Freshwater Invertebrate Biol. 2: 165— WA. REYNOLDSON, T. B., AND R. W. Davies. 1980. A compara- tive study of weight regulation in Nephelopsis ob- scura and Erpobdella punctata (Hirudinoidea). Comp. Biochem. Physiol. 66A: 711-714. TILLMAN, D. L., ANDJ. R. BARNES. 1973. The reproductive biology of the leech Helobdella stagnalis (L.) in Utah Lake, Utah. Freshwater Biology 3: 137-145. LITERATURE CITED ON MOLLUSKS BalLy, J. L., JR, AND R. I. Batty. 1951-1952. Further ob- servations on the Mollusca of the relict lakes in the Great Basin. Nautilus 65: 46-53, 85-93. BaRNES, J. R., ANDT. W. TOOLE. 1981. Macroinvertebrate and zooplankton communities of Utah Lake: a re- view of the literature. Pages 101-106 in Great Basin Naturalist Mem. No. 5. BICKEL, D. 1977. A survey of locally endemic Mollusca of Utah, Colorado, Wyoming, Montana, North Da- kota and South Dakota. Department of Interior, Fish and Wildlife Service, Office of Endangered Species. Contract 14-16-0006-3030. CALL, R. E. 1884. On the Quaternary and Recent Mol- lusca of the Great Basin with descriptions of new forms. Bull. U.S. Geol. Surv. 11: 367—420. 744 CENTER FOR HEALTH AND ENVIRONMENTAL STUDIES. 1975. Environmental studies of: proposed Jor- danelle Reservoir site, Provo River, Utah Lake, Jordan River, proposed Lampton Reservoir site. Final Phase I report to the Bureau of Reclamation. Brigham Young University, Provo, Utah. CHAMBERLIN, R. V. 1933. Observations of Stagnicola kingi (Meek), living and extinct. Nautilus 46: 97-100. CHAMBERLIN, R. V., AND D. T. JoNEs. 1929. A descriptive catalog of the Mollusca of Utah. Bull. University of Utah 19: 1-203. HENDERSON, J. 1931. The problem of the Mollusca of Bear Lake and Utah Lake, Idaho-Utah. Nautilus 44: 109-113. JonEs, D. T. 1940a. Recent collections of Utah Mollusca, with extralimital records from certain Utah cabi- nets. Utah Acad. Sci., Arts, and Let. 17: 33-45. —__.. 1940b. Mollusks of the Oquirrh and Stansbury mountains. Nautilus 54: 27-29. Ke!TH, M. L., AND G. M. ANDERSON. 1963. Radiocarbon dating: fictitious results with mollusk shells. Sci- ence 141: 634-637. Murray, H. D. 1970. Discussion of Dr. Taylor's paper. Malacologia 10: 33-34. Riccs, A. C. 1984. Major carbon-14 deficiency in modern snail shells from southern Nevada springs. Science 141: 637. RvBIN, M., AND D. W. TayLor. 1963. Radiocarbon activity of shells from living clams and snails. Science 141: 637. RUSSELL, R. H. 1971. Mollusca of Fish Springs, Juab County, Utah: rediscovery of Stagnicola pilsbryi (Hemphill 1890). Great Basin Nat. 31: 223-236. SmakT, E. W. 1963. The peculiar mollusk populations of Bear Lake. Utah Acad. Sci., Arts, and Let. 40: 197-199. TayLor, D. W. 1960. Distribution of the freshwater clam Pisidium ultramontanum: a zoogeographic in- quiry. Amer. J. Sci. 258A: 325-334. ——___.. 1966. Summary of North American Blancan non- marine mollusks. Malacologia 4: 1-172. TayLor, D. W., H. J. WALTER, AND J. B. BurcH. 1963. Freshwater snails of the subgenus Hinkleyia (Lymnaeidae: Stagnicola) from the western United States. Malacologia 1: 237-281. WILLIAMS, J. S., A. D. WILLARD, AND V. PARKER. 1962. Recent history of Bear Lake Valley, Utah-Idaho. Amer. J. Sci. 260: 24-36. YEN, T. C. 1951. Fossil fresh-water mollusks and ecologi- cal interpretations. Bull. Geol. Soc. Amer. 62: 1375-1380. GREAT BASIN NATURALIST Vol. 46, No. 4 LITERATURE CITED ON AMPHIBIANS BantTA, B. H. 1965. A distribution checklist of recent am- phibians inhabiting the state of Nevada. Occ. Pap. Biol. Soc. Nevada 7: 1—4. CREUSERE, F. M., AND W. G. WHITFORD. 1976. Ecological relationships in a desert anuran community. Her- petologica 32: 7-18. Dun.ap, D. G. 1977. Wood and Western Spotted Frogs (Amphibia, Anura, Ranidae) in the Big Horn Mountains of Wyoming. J. Herp. 11: 85-87. HOVINGH, P., B. BENTON, AND D. BORNHOLDT. 1985. Aquatic parameters and life history observations of the Great Basin Spadefoot Toad in Utah. Great Basin Nat. 45: 22-30. La Rivers, I. 1942. Some new amphibian and reptile records for Nevada. J. Entom. & Zool. 34: 53-68. Licut, L. E. 1974. Survival of embryos, tadpoles, and adults of the frogs Rana aurora aurora and Rana pretiosa pretiosa sympatric in southwestern British Columbia. Canad. J. Zool. 52: 613-627. LINSDALE, J. M. 1940. Amphibians and reptiles in Ne- vada. Proc. Amer. Acad. Arts Sci. 73: 197-257. Morris, R. L., AND W. W. TANNER. 1969. The ecology of the western spotted from Rana pretiosa pretiosa Baird and Girard, a life history study. Great Basin Nat. 29: 45-81. REYNOLDS, T. D., AND T. D. STEPHENS. 1984. Multiple ectopic limbs in a wild population of Hyla regilla. _ Great Basin Nat. 44: 166-169. STEPHENS, J. C. 1977. Hydrologic reconnaissance of the Tule Valley drainage basin, Juab and Millard counties, Utah. Tech. Publ. Division of Water Rights No. 56. TANNER, V. M. 1931. A synoptical study of Utah amphibi- ans. Utah Acad. Sci., Arts, and Let. 8: 159-198. TANNER, W. W. 1978. Zoogeography of reptiles and am- _ phibians in the Intermountain Region. Pages — 43-53 in Intermountain biogeography: a sympo- | sium. Great Basin Naturalist Mem. 2. TURNER, F. D. 1962. An analysis of geographic variation | and distribution of Rana pretiosa. Amer. Phil. Soc. Yearbook 1962: 325-328. TURNER, F. D., AND P. C. Dumas. 1972. Rana pretiosa. | Pages 119.1-119.4 in American amphibians and | reptiles. HATCHING CHRONOLOGY OF BLUE GROUSE IN NORTHEASTERN OREGON John A. Crawford’, Walter Van Dyke’, Victor Coggins’, and Martin St. Louis? ABSTRACT. —Hatching chronology of Blue Grouse (Dendragapus obscurus) in northeastern Oregon was determined from 431 immatures examined from 1981 to 1985. Young hatched from 1 May through 8 July; median hatching dates for the five years ranged from 27 May to 5 June. Peak hatching in Oregon occurred from one to four weeks earlier than in most portions of the range of Blue Grouse but were similar to north central Washington and Idaho. Variations in hatching dates possibly were related to rainfall. Information concerning hatching chronol- ogy is essential for study of the breeding ecol- ogy (e.g., survival and recruitment) of bird populations and is useful for determining opti- mum times for population censusing. Bendell and Zwickel (1985) summarized hatching times of Blue Grouse from 25 locations throughout the range and noted that peak hatching dates, which ranged from 24 May to 11 July, were earliest near the center of distri- bution. Results of research on Vancouver Is- land, British Columbia (Bendell 1955, Zwickel and Bendell 1967, King 1971, Red- field 1975), southwestern Alberta (Boag 1965), and southcentral Montana (Mussehl 1960) indicated that peak hatching times of Blue Grouse were remarkably similar, mid- to late June, among these diverse locations. In contrast, Blue Grouse were reported to hatch from two to four weeks earlier in north- central Washington (Standing 1960, Hender- son 1960, Bauer 1962, Zwickel 1973) and western Idaho (Caswell 1954). Brown and Smith (1980) noted that most immatures were from six to eight weeks old at the end of Au- gust (indicating most hatched from early to mid-July) on their study area in eastern Ari- zona. Redfield (1975) and Zwickel (1977) found hatching dates of Blue Grouse differed among years on Vancouver Island and at- tributed these differences to annual variations in weather. Data are unavailable regarding hatching times from several portions of the range of this species, including Oregon. The purpose of this project was to determine the hatching chronology of Blue Grouse in north- eastern Oregon and to examine, evaluate, and assess annual variations in hatching times. From 1981 through 1985 wings and tails of 775 Blue Grouse taken by hunters from 29 August to 30 September each year were col- lected in Wallowa County in northeastern Or- egon. Blue Grouse in this region typically occupy coniferous forest, timbered draws, and adjacent grass-shrubland habitats at ele- vations ranging from 600 to 1,500 m. Age of birds was classified as adult or immature (young of the year) based on the condition of the outer two primaries (Bunnell et al. 1977) and the presence of juvenal feathers in the wing or tail. Sufficient information (date of kill and primary feathers present) was available to estimate hatching dates for 431 of 467 imma- tures examined. Dates of hatching were based on replacement rate of primary feathers (Zwickel and Lance 1966) and corrected for bias (Redfield and Zwickel 1976). Only 17 of the 431 immatures had com- pleted molting of the primaries and, in all instances, presence of sheathing material at the base of the eighth primary indicated the molt was recently finished. These birds were assigned the maximum age of 123 days. Me- dian (suggested by Redfield 1975) and mean hatching dates and periods of peak (14-day period in which greatest number of young hatched) and maximum (time interval during which > 70% of young hatched) hatching were based on young hatched/day. These data were summarized by weekly intervals, begin- ning with the earliest hatching date. One-way analysis of variance and the Student- Department of Fisheries and Wildlife, Oregon State University, Corvallis, Oregon 97331-3803. Oregon Department of Fish and Game, LaGrande, Oregon 97850. 3Oregon Department of Fish and Game, Enterprise, Oregon 97828. 745 746 GREAT BASIN NATURALIST Vol. 46, No. 4 TABLE 1. Percent of young Blue Grouse hatched during weekly intervals and median and range of hatching dates, northeastern Oregon, 1981-1985. 1981 1982 Category (n = 44) (n = 34) WEEK 1-7 May 2 0 8-14 May 2 3 15-21 May 23 15 22-28 May 16 24 29 May—4 June 27 32 5-11 June ll 21 12-18 June 9 6 19-25 June 5 0 26 June—2 July 5 0 3-9 July 0 0 HATCHING DATES Mean 30 May 30 May Median 29 May 31 May Range 1 May—29 June 14 May-18 June 1983 1984 1985 (n = 106) (n = 81) (n = 164) 1 0 1 5 1 6 8 6 15 AT 21 23 18 19 26 7 26 13 fT 14 11 8 ii 4 1 3 0 0 4 2 30 May 7 June 1 June 27 May 5 June 31 May 5 May-—30 June 13 May-8 July 5 May-8 July TABLE 2. Periods of peak (14-day period when most young hatched) and maximum (time during which > 70% of young hatched) hatching of Blue Grouse. Dates of peak Location hatching Dates Vancouver Island, 15-21 June’ 8—28 June British Columbia 17-30 June” 11 June-1 July Hardwicke Island, — 4—25 June British Columbia Southwestern 14-21 June — Alberta North central 22 May—4 June — Washington 10 May—25 May South central 14-21 June 9 June-3 July® Montana Western Idaho 25 May-7 June — Northeastern 20 May-2June 18 May-—7 June Oregon 1Some authors reported peak hatching periods of < 14 days. : Approximate mean dates for three years (two areas/year). 3Average dates for two years. man-Keuls mean separation test (Snedecor and Cochran 1967) were used to test for differ- ences in mean hatching dates among years. Precipitation and mean temperatures for March, April, and May of each year were obtained for the Enterprise weather station (Climatological Data Oregon, NOAA) and compared to median hatching dates. Hatching dates of Blue Grouse in north- eastern Oregon (Table 1) ranged from 1 May (1981) to 8 July (1984 and 1985). Median hatching dates (Table 1) ranged from 27 May Period of maximum hatching % young hatched Source 80 Bendeli 1955, King 1971 72 Zwickel and Bendell 1967 — Zwickel (personal communication) — Boag 1965 — Standing 1960, Bauer 1962 Zwickel 1973 — Zwickel (personal communication) 80 Mussehl 1960 — Caswell 1954 71 This study (1983) to 5 June (1984). Comparisons of mean | hatching dates, all of which were + 3 days of | respective median dates, indicated that hatching in 1984 (7 June) was significantly (P < 0.05) later than in the other four years | (range 30 May-—1 June); no other differences | were detected. Length of the hatching period. for all years combined, 10 weeks, was identi-' cal to that reported by Zwickel and Bendell. (1967) for Blue Grouse on Vancouver Island. | The mean annual length of the hatching pe-| riod in northeastern Oregon was 54 days, October 1986 however. In this study, hatching was concen- trated from mid-May to mid-June (Table 1). For all years combined, 54% of the young hatched in May, 45% in June, and 1% in July. Zwickel and Bendell (1967) noted peak hatch- ing lasted approximately two weeks on Van- couver Island, and the period of maximum hatching (> 70%) took place within three to four weeks; they found that 67% of the young hatched during the peak, 7% before, and 26% after the peak. For northeastern Oregon the peak of hatching (55% of the young) occurred from 20 May to 2 June (Table 2); 9% hatched before the peak and 36% after. Maximum hatching of young (71%) took place from 18 May to 7 June. Median hatching date for the five years was 31 May and the mean was | June. Dates of peak hatching of Blue Grouse in northeastern Oregon (Table 2) were from two to four weeks earlier than those reported for British Columbia, Alberta, Mon- tana, and Arizona (Bendell 1955, Mussehl 1960, Boag 1965, Zwickel and Bendell 1967, Brown and Smith 1980, Zwickel, personal communication) and one week earlier than hatching in northern California and northern Nevada (Zwickel, personal communication). Hatching dates were similar to those reported in north central Washington (Standing 1960, Henderson 1960, Bauer 1962, Zwickel 1973) and Idaho (Caswell) 1954. Factors affecting the timing of reproductive activities of Blue Grouse throughout their range are incompletely understood. King (1971) found that hatching times were related to elevation; Blue Grouse in subalpine areas hatched approximately 3.5 weeks later than those living at lower elevations. Marshall (1946) proposed that plant phenology in spring influenced the timing of migration of Blue Grouse, which in turn affected breeding times. Plant phenology also may directly in- fluence breeding times (Zwickel, personal communication). Plant phenology throughout the range of Blue Grouse is strongly influ- enced by elevation and latitude. Blue Grouse populations with which our data were com- pared (Table 2), except for birds in north cen- tral Washington and western Idaho, inhabited areas either farther north by > 4° (British Columbia and Alberta) or at higher elevations (= 1800 m in Montana and Arizona), which may account for earlier breeding in northeast- ern Oregon. The study site in Washington was CRAWFORD ET AL.: OREGON BLUE GROUSE 747 approximately 3° north of our area but was lower in elevation (450-900 m) and the study area of Caswell (1954) in Idaho bordered northeastern Oregon. Zwickel (1977) noted that temperature and precipitation partially accounted for annual differences in hatching chronology within populations; earlier hatching coincided with warm, dry conditions during April and May. Redfield (1975) suggested that annual differ- ences within populations were related to spring temperatures. In our study the median hatching date in 1984 was from 6 to 10 days later than in any of the other four years. Com- parisons of median hatching dates with mean monthly temperature and total monthly pre- cipitation during March, April, and May re- veal that precipitation during April/May (10.7 cm) and from March through May (15.4 cm) 1984 was the highest of the five years; mean values for the other four years were 9.2 cm and 13.4 cm, respectively. Temperature data for 1984 were similar to the other four years. No other trends were apparent from these data, and limited sample size (five years) pre- cluded statistical testing. We concluded that hatching times of Blue Grouse in northeastern Oregon were similar in most years. Mean hatching date differed only in 1984; all dates were within a 10-day interval. Latest hatching corresponded to the wettest spring of the five years. Hatching dates in northeastern Oregon were consistent with the observation of Bendell and Zwickel (1985) of early breeding within the central portion of the range of Blue Grouse. ACKNOWLEDGMENTS We would like to thank T. F. Haensly and S. M. Meyers for their assistance in determin- ing sex and age of birds. The manuscript was reviewed by F. C. Zwickel, E. C. Meslow, and R. L. Jarvis. Special appreciation is ex- pressed to F. C. Zwickel for generously sharing unpublished data. This is Technical Publication 7838 of the Oregon Agricultural Experiment Station. LITERATURE CITED BAUER, R. D. 1962. Ecology of Blue Grouse on summer range in north central Washington. Unpublished the- sis, Washington State University, Pullman. 81 pp. 748 BENDELL, J. F. 1955. Age, breeding behavior and migra- tion of Sooty Grouse, Dendragapus obscurus fuliginosus (Ridgway). Trans. N. Amer. Wildl. Conf. 20: 367-381. BENDELL, J. F., AND F. C. ZWICKEL. 1985. A survey of the biology, ecology, abundance and distribution of the Blue Grouse (Genus Dendragapus). Third Intern. Grouse Symp., World Pheasant Assoc. (in press). Boac, D. A. 1965. Indicators of sex, age and breeding phenology in Blue Grouse. J. Wildl. Manage. 29: 103-108. Brown, D. E., AND R. H. SmiTu. 1980. Winter-spring pre- cipitation and population levels of Blue Grouse in Arizona. Wildl. Soc. Bull. 8: 136-141. BUNNELL, S. D., J. A. RENSEL, J. F. KIMBALL, JR., AND M. L. WOLFE. 1977. Determination of age and sex of Dusky Blue Grouse. J. Wildl. Manage. 41: 662-666. CASWELL, E. B. 1954. A preliminary study of the life history and ecology of the Blue Grouse in west central Idaho. Unpublished thesis, University of Idaho, Moscow. 105 pp. HENDERSON, U. B. 1960. A study of Blue Grouse on sum- mer range, north central Washington. Unpub- lished thesis, Washington State University, Pull- man. 96 pp. Kinc, D. G. 1971. The ecology and population dynamics of Blue Grouse in the subalpine. Unpublished thesis, University of British Columbia, Vancouver. 139 pp. GREAT BASIN NATURALIST Vol. 46, No. 4 MARSHALL, W. H. 1946. Cover preferences, seasonal movements, and food habitats of Richardson’s Grouse and Ruffed Grouse in southern Idaho. Wilson Bull. 58: 42-52. MUSSEHL, T. W. 1960. Blue Grouse production, move- ments, and populations in the Bridger Mountains, Montana. J. Wildl. Manage. 24: 60-68. REDFIELD, J. A. 1975. Comparative demography of in- creasing and stable populations of Blue Grouse (Dendragapus obscurus). Canadian J. Zool. 53: 1-11. REDFIELD, J. A., AND F. C. ZWICKEL. 1976. Determining the age of young Blue Grouse: a correction for bias. J. Wildl. Manage. 40: 349-351. SNEDECOR, G. W., AND W. G. Cocuran. 1967 Statistical methods. 6th ed. Iowa State University Press, Ames. 593 pp. ZWICKEL, F. C. 1973. Dispersion of female Blue Grouse during the brood season. Condor 75: 114-119. _____. 1977. Local variations in the time of breeding of female Blue Grouse. Condor 79: 185-191. ZWICKEL, F. C., AND J. F. BENDELL. 1967. Early mortality and the regulation of numbers in Blue Grouse. Canadian J. Zool. 45: 817-851. ZWICKEL, F. C., AND A. N. LANCE. 1966. Determining the age of young Blue Grouse. J. Wildl. Manage. 30: 712-717. NEW SOUTH AMERICAN LEAFHOPPERS IN THE GENUS DOCALIDIA, WITH A KEY TO 37 SPECIES (CICADELLIDAE: COELIDIINAE, TERULIINI) M. W. Nielson! ABSTRACT.—Ten new species of Docalidia are described and illustrated. These are pennyi, gracilitas, zanoli, triquetra, paracrista, convexa, setacea, and caterva from Brazil and vesica and vella from Peru. A key to males of 37 species described since the last revision of the genus is included. The number of known species is now 116, making Docalidia the largest teruliine genus. Since the treatment of the genus Docalidia in a revision of the tribe Teruliini (Nielson 1979), several new species in the genus have been described (Nielson 1982a, 1982b, 1982c). In this paper 10 additional new spe- cies are described and illustrated, bringing the total to 116 known species in this the largest of the teruliine genera. The richness of the fauna of this group in South America (only one species known in Central America and one in the West Indies) is staggering. The number of known species prior to 1979 was 21 and since then the num- ber has increased nearly 600%. Most of the new species described herein and in earlier papers were the result of collections made during the last 20 years. As new areas of tropi- cal America become more accessible and col- lections more extensive, many new species of Docalidia will be found. Docalidiine leafhoppers are small to medium-sized, robust species with short, broad heads. The crown is short, broad, de- pressed, but usually not carinate, and the pronotum is noticeably inflated in many spe- cies. A very well-developed median clypeal carina distinguishes this group from other similar appearing genera that usually have a weakly developed clypeal carina. The long, usually slender, aedeagus with or without a single subapical ventral spine, simple to or- nate style and 10th segment processes, and the broad plate will readily distinguish the group from all other teruliine genera. A key to males of 37 species not previously keyed is given to accommodate species de- scribed here and in my three earlier papers cited above. The remainder are keyed in my 1979 paper. Hosts and biology of these leafhoppers are poorly known. Paucity of populations ac- counts for the lack of knowlege of their bio- nomics and importance to agriculture and sil- viculture in the tropics of the new world. Key to Males of Docalidia iL, Style simple, without spines or setae....... 2 — Style ornate, with spines and/or setae PLESENt ies asians oleate See eae 6 Style filamentous in distal half ............ 3 — Style broad in distal half ................. 4 Pygofer with small caudoventral lobe (Fig. 1); aedeagus with subapical spine (Fig. Set edran el he mee mato GS ic oie one caterva, n. sp. — Pygofer with very long broad caudoventral process (Fig. 19, Nielson 1982b); aedeagus without subapical spine (Fig. 23, Nielson 1982.5) ey ones cen meneame exilis Nielson Aedeagus with subapical spine; pygofer with caudodorsal process bifurcate or rounded dis- tallyectoy NGA ee te Sena ah cee oer er recente 5 — Aedeagus without subapical spine (Fig. 83, Nielson 1982a); pygofer with caudodorsal pro- cess single and pointed (Fig. 79, Nielson MOS Qa re Mitte wes ores aerate: glabra Nielson Pygofer with caudoventral process very long and lobelike, apex rounded (Fig. 1, Nielson MOS Dea ee hv a RN oie lobata Nielson — Pygofer with caudoventral process short, nar- row, curved mesally and pointed apically (Fig. 31, Nielson 1982a)....... hansoni Nielson Style with 1-3 spines in distal halfto third .. 7 — Style with numerous spines and/or setae in distallhalfitojthind@aemanmeneeoeco eee 9 Aedeagus with subapical spine (Fig. 17, Niel- son 1982a); style with single, terminal spine (Fig. 14, Nielson 1982a).......... nuda Nielson ‘Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah 84602. 749 11(10). GREAT BASIN NATURALIST Aedeagus without subapical spine; style with lateralispineés®s a2 cc pytvsn ae ete 8 Pygofer with long caudoventral process (Fig. 37, Nielson 1982b); style with long subapical spine directed distally (Fig. 38, Nielson TIC 5355) 0} ipa is MO aM a he Sa hawsi Nielson Pygofer without such process (Fig. 49, Niel- son 1982b); style with long spine medially and two short subapical spines directed laterally (Fig. 50, Nielson 1982b) tuberculata Nielson Pygofer with distinctive long caudoventral DIOCESS cient sie mene Me an eR RS eReMantn aioe 10 Pygofer without such process (small lobe of- tenipresent)eyigis lee oe eaters toe te DP) Segment 10 without well-developmed, orna- mental ventral processes (often much re- ducedibutiapparent)s seen eer eee 11 Segment 10 with well-developed, ornamental VeEnitralj prOCeSSeS Meas ela) ae eee hee 12 Aedeagus with subapical spine (Fig. 64, Niel- son 1982a), style with spines in distal half (Fig. 62, Nielson 1982a)........ paragracilis Nielson Aedeagus without such spine (Fig. 35, Niel- son 1982b); style with spines in distal third ie Ne TR eR RG a a geet crista Nielson Stata thn suse ei eha Oe 13 Style with spines and setae combined ...... 21 Style with spines or setae . Style with spines only, not densely packed.. 14 Style with setae only, densely packed appear- ingiaswelvet (Higa s) meester vella, n. sp. . Style with one to two large medial spines (or near middle) and numerous shorter spines belowanidistallhalffe amen ceede see ae 15 Style without medial spines, numerous spines in distal third . Style with shaft narrow in distal half........ 16 Style with shaft expanded in distal third (Fig. PAU) etterser ces ake. ear teteey Panereeere eae zanoli, n. sp. . Style with spines below medial spine(s) long and on inner lateral margin............... We Style with spines below medial spine very short (spiculated) and on dorsal margin (Fig. 74, Nielson 1982a)........... corneola Nielson . Segment 10 with very long ventral processes (Fig. 55, Nielson 1982a); style with single medial spine (Fig. 57, Nielson 1982a) Fo Re eater ieee Cee ate rare dorsti Nielson Segment 10 with short ventral processes (Fig. 67, Nielson 1982a);style with two medial spines (Fig. 68, Nielson 1982a) Be 3 tN cy cr er he bispinata Nielson . Pygofex with distally enlarged, caudally trun- Sra eRe mee reheat ce 19 Pygofer with bladelike caudodorsal process . 20 cate, caudodorsal process . Pygofer with caudodorsal process with teeth in dorsal half (Fig. 7, Nielson 1982c); style with large teeth on inner lateral margin and without vesica in distal third (Fig. 8, Nielson 19820) cee eee eee multidentata Nielson 20(18). 21(12). 25(24). 26(23). Vol. 46, No. 4 Pygofer with caudodorsal process with fine serrations on caudal margin (Fig. 23); style with fine teeth or serrations on inner lateral margin and with large bladder or vesica in distal one-eighth (Fig. 27)........ vesica, n. sp. Pygofer with caudal processes close together (Fig. 30); style with long spines near dorsal Thaneinl (Riga 34) ee pennyi, n. sp. Pygofer with caudal processes widely sepa- rated (Fig. 25, Nielson 1982a); style with short serrations on inner lateral margin (Fig. 26, Nielson 1982a) taylori Nielson Segment 10 with long ventral process extend- ing beyond segment 10 (Fig. 1, Nielson 1982c); plate long and narrow (Fig. 6, Nielson T9826) aoa Noe a eee breddini Nielson Segment 10 with short ventral process with short, bluntly pointed, subapical secondary process (Fig. 7, Nielson 1982a); plate long and broad (Fig. 12, Nielson 1982a) Dee a Ra S863 0°0 0 robertsi Nielson Aedeagus with very short, subapical ventral Spineior spine absent... eee 23 Aedeagus with long, subapical ventral spine. 27 Aedeagus with ventral spine not more than three times as long as wide ............... 24 Aedeagus with ventral spine absent........ 26 Segment 10 with well-developed ventral pro- cesses; style with 1-2 prominent lateral spines on middle of inner lateral margin .... 25 Segment 10 with poorly developed ventral processes (Fig. 19, Nielson 1982a); style with- out such spines (Fig. 20, Nielson 1982a) eo eo ves ue patula Nielson Segment 10 with long ventral process nearly reaching to apex of segment 10, process ter- minating with short blunt point (Fig. 1, Niel- son 1982b); style with two medial spines be- fore dentate inner margin below (Fig. 2, Nielson 982.b))h saan eeeEoeoceee rema Nielson Segment 10 with shorter ventral process, pro- cess reaching to middle of segment 10, pro- cess narrowed at distal half and curved dor- sally (Fig. 37); style with one medial spine before dentate inner margin below (Fig. Ct ee IG ela Gb co's 6 triquetra, n. sp. Style with spines and setae on distal one-fifth, spines on inner lateral margin, setae arranged in one large tuft dorsally (Fig. 26, Nielson 19825) oceania toe Oe thola Nielson Style with setae only on distal third, setae on inner and outer lateral margins (Fig. 44, Niel- Sons! 982) hanna bipenicula Nielson Style with distal half to third with one to two prominent spines and with lateral setae on MAFPINS 6. ee ses co oe Oe eee 28 Style with distal half to third with lateral setae only or with lateral spines only on margins .. 32 27). Segment 10 with poorly developed ventral processes; pygofer with straight or nearly straight caudodorsal processes ............ 29 | October 1986 NIELSON: NEW AMERICAN LEAFHOPPERS 751 Figs. 1-8. Docalidia caterva, n. sp.: 1, Male pygofer and segment 10, lateral view. 2, Segment 10 and pygofer processes, ventral view. 3, Aedeagus, lateral view. 4, Aedeagus, ventral view. 5, Connective and right style, dorsal view. 6, Style, lateral view. 7, Plate, ventral view. 8, Female venter, ventral view. 32(27). Segment 10 with large well-developed ven- tral process (Fig. 7, Nielson 1982b); pygofer with caudodorsal process decurved apically (Fig. 7, Nielson 1982b) .......... unca Nielson Style with setae occupying distal half, setae longdandimostly/stoutsano enone oe. Style with setae occupying distal third, setae short and fine Style with prominent medial lobe on inner lateral margin (Fig. 48), aedeagus with broad Shatta(BiowAd) penne ee setacea, n. sp. Style without such lobe (Fig. 55), aedeagus with narrow shaft (Fig. 54)..... gracilitas, n. sp. Style narrow throughout with short, stout spine subapically in dorsal view (Fig. 38, Niel- SOnGLOS 2a) apres nies digitata Nielson Style triangulate in distal third with long nar- row spine distad of middle in lateral view (Fig. 51, Nielson 1982a) ............ hirsuta Nielson Style with spines only 33(32). Style with setae only Style with convex lobe medially on inner lat- eral margin, spines confined to medial lobe (Big G4) ceca tiara tana convexa, n. sp. Style not as above, spines occupying entire distal third (Fig. 74)......... paracrista, n. sp. Style with distal half narrowed throughout .. 35 Style with distal half to third broadly ex- JOY: 110-10 bene rer re Mec rte meee atimita asi elon ri ra 36 . Style with setae on inner lateral margin from distal three-fourths to apex (Fig. 44, Nielson 1982a) permagna Nielson Style with setae on dorsal margin basal of apex (Fig. 19, Nielson 1982c)...... subcrista Nielson . Style with distal half expanded (Fig. 14, Niel- son 1982c); segment 10 without ventral pro- cess (Fig. 13, Nielson 1982c). . . lateralis Nielson Style with distal third expanded, triangulate (Fig. 14, Nielson 1982b); segment 10 with short ventral process (Fig. 13, Nielson 1982b) triangulata Nielson 752 GREAT BASIN NATURALIST Vol. 46, No. 4 > | \ ‘ Figs. 9-15. Docalidia vella, n. sp.: 9, Male pygofer and segment 10, lateral view. 10, Segment 10 and pygofer processes, ventral view. 11, Aedeagus, lateral view. 12, Aedeagus, ventral view. 13, Connective and right style, dorsal view. 14, Style, lateral view. 15, Plate, ventral view. Docalidia caterva, n. sp. Figs. 1-8 LENGTH.—Male, 5.30 mm, female, 6.40 mm. Small, robust species. General color black with two narrow, broken, pale, ochraceous transverse bands on forewings (wider in fe- male) and numerous pale to ochre spots on veins and cells, apex pale to ochraceous; crown ochre; eyes reddish brown; pronotum and scutellum black in male, brown in female; face black in male, light brown in female. Head large, broad, nearly as wide as prono- tum; crown short, narrow, much narrower than width of eyes, depressed, lateral margin convergent basally; eyes large, elongate ovoid; pronotum short, slightly longer than crown, inflated; scutellum large, much longer than pronotum, inflated anteriorly; forewings long and broad; clypeus narrow, median clypeal carina well developed; clypellus nar- row, lateral margins flared distally. MALE.—Pygofer in lateral view with very small caudoventral lobe, caudodorsal margin with long narrow process (Figs. 1, 2); segment 10 without ventral processes (Figs. 1, 2); aedeagus asymmetrical, long, nearly tubular, broadly curved in lateral view, with long sub- apical spine on lateroventral margin and toothed medially on one side of dorsolateral margin (Figs. 3, 4); gonopore very large on lateroventral margin (Fig. 4); style long, very narrow and needlelike in distal half, reaching to about middle of aedeagal shaft (Figs. 5, 6); plate long, narrow, slightly constricted medi- ally, enlarged subapically, and narrowed dis- tally to rounded apex (Fig. 7). FEMALE.—Seventh sternum large, about _ two to three times as long as preceding seg- { \ ! i I) October 1986 ment, caudal margin produced at middle, with shallow narrow concavity medially and short blunt projection on either side of middle (Fig. 8). HOLOTYPE (male).—BRAZIL: Am. [Ama- zonas|, Manaus, INPA, 1.V.1976, E. Castel- lon B. (MZUSP). Allotype female, same data as holotype, except 28.1V.1976 (MZUSP). REMARKS.—This species is similar in gen- eral habitus to limpidosparsa (Stal) but is eas- ily separated by the very small caudoventral lobe on the pygofer, by the needlelike style, and by the dentations on the aedeagal shaft. \ Docalidia vella, n. sp. Figs. 9-15 LENGTH.—Male, 6.40 mm. Small, robust species. General color black except for pale anterior area of crown and narrow, ochre apical margin of forewings. Head short, broad, much narrower than pronotum; crown broad, about as wide as eye, lateral margins parallel except convergent near base; pronotum short, about as long as crown, inflated; scutellum large, median length greater than median length of prono- tum; forewing long and broad; clypeus long and broad; median clypeal carina well devel- oped; clypellus narrow; lateral margins nearly parallel. MALE.—Pygofer in lateral view with long, narrow, apically decurved caudoventral pro- cess (Figs. 9, 10); caudodorsal margin with process as in vesica (Fig. 9); segment 10 with ventral processes as in vesica (Figs. 9, 10, 23); aedeagus asymmetrical; configuration as in vesica except without subapical spine (Figs. 11, 12, 25); style long, nearly as long as aedea- gus, basal half narrow, distal half very broad, inner dorsal margin lobed distally, with longi- tudinal fold along entire inner lateral margin in distal half, this margin densely covered with very short setae giving an appearance of velvet, apex with few weak membranous spines (Figs. 13, 14); plate long and very broad, apex with short macrosetae (Fig. 15). FEMALE.— Unknown. HOLOTYPE (male).—PERU: Madre de Dios, Rio Tambopata Res., 30 air km SW Pto. Maldonado, 290 m, subtropical moist forest, 11-15. X1.1979, J. W. Heppner (USNM). REMARKS.—This species is similar in gen- NIELSON: NEW AMERICAN LEAFHOPPERS 753 eral habitus and some male genital characters to vesica but can be distinguished by the ab- sence of the subapical aedeagal spine and by the velvetlike inner lateral margin in the en- larged distal half of the style. Docalidia zanoli, n. sp. Figs. 16-22 LENGTH.—Male, 6.75 mm. Small robust species. General color dark brown with broad, nearly complete, trans- verse translucent band on middle _ of forewings, veins black with pale spots; crown pale; eyes light brown; pronotum and scutel- lum black with bullae on pronotum pale; face black. Head short, very broad, slightly narrower than pronotum; crown broad about as wide as eye, depressed, lateral margins as in gracili- tas; eyes large, semiglobular; pronotum and scutellum as in gracilitas except more deeply marked; forewing long and broad; clypeus and clypellus configuration as in gracilitas. MALE.—Pygofer in lateral view with long, narrow, distally curved caudoventral process and with long, narrow caudodorsal process, caudodorsal process longer than caudoventral process (Figs. 16, 17); segment 10 with ventral process very large, broad at basal half, nar- rowed at distal half, curved caudodorsally and extending to apex of segment 10 (Figs. 16, 17); aedeagus asymmetrical, long, narrow along dorsal margin, broad along ventral margin, slightly curved throughout in lateral view with long, narrow subapical spine on ventral margin (Figs. 18, 19); gonopore basad of mid- dle on ventral margin; style long, reaching to about apex of aedeagus, with medial process on inner lateral margin, process asymmetri- cally toothed distally, distal third of style ex- panded, with inner lateral margin serrated and submarginally sclerotized, apex rounded with apical teeth (Figs. 20, 21); plate long and broad, tapered toward bluntly rounded apex (Fig. 22). FEMALE.— Unknown. Ho.otyPe (male).—BRAZIL: Cruzeiro do Sul, ACRE, .___.II.1963, M. Alvarenga (UFP). REMARKS.—This species is similar in male genital characters to dentatula (Metcalf), but it can easily be separated by the presence of a 754 17 GREAT BASIN NATURALIST Vol. 46, No. 4 Figs. 16-22. Docalidia zanoli, n. sp.: 15, Male pygofer and segment 10, lateral view. 17, Segment 10 and pygofer processes, ventral view. 18, Aedeagus, lateral view. 19, Aedeagus, ventral view. 20, Connective and right style, dorsal view. 21, Style, lateral view. 22, Plate, ventral view. long medial process on the inner lateral mar- gin of the style. I name this species for Keti Maria Rocha Zanol, Universidade Federal Do Parana, who has kindly sent me material for study and who is working on the Delto- cephalinae of Brazil. Docalidia vesica, n. sp. Figs. 23-29 LENGTH.—Male, 6.40 mm. Medium-sized, robust species. General color black throughout with narrow ochra- ceous band distally on forewings. Head short, broad, much narrower than pronotum; crown short, broad, about as wide as eye, depressed on either side of middle, lateral margins parallel except convergent | basally; pronotum short, about as long as crown, slightly inflated; scutellum large, me- dian length greater than median length of | pronotum; forewing long and broad; clypeus long and broad, median clypeal carina well developed; clypellus narrow, lateral margins | slightly expanded distally. MALE.—Pygofer in lateral view with long, very narrow, sinuate caudoventral process, | caudodorsal margin with long, rather stout | process, process enalrged distally, caudal — margin subtruncate and finely serrate (Figs. | 23, 24): segment 10 with well-developed ven- | tral processes, process very broad in basal | half, with narrow projection curved dorsally | (Figs. 23, 24); aedeagus asymmetrical, long | { |} October 1986 NIELSON: NEW AMERICAN LEAFHOPPERS 755 Figs. 23-29. Docalidia vesica, n. sp.: 23, Male pygofer and segment 10, lateral view. 24, Segment 10 and pygofer processes, ventral view. 25, Aedeagus, lateral view. 26, Aedeagus, ventral view. 27, Connective and right style, dorsal view. 28, Style, lateral view. 29, Plate, ventral view. and nearly needlelike throughout, broadly curved in lateral view, with long subapical spine on ventral margin (Figs. 25, 26); gonopore basad of middle on lateral margin; style very long, extending beyond apex of aedeagus, broad throughout except for con- striction along middle, finely toothed in distal third of inner lateral margin and with large, saclike membranous bladder dorsally in distal one-eighth (Figs. 27, 28); plate long and very broad with microsetae distally (Fig. 29). FEMALE.— Unknown. HOLOTYPE (male).—PERU: Monson Val- ley, Tingo Maria, 8.X.1954, E. L. Schlinger and E. S. Ross (CAS). REMARKS.—Docalidia vesica has similar style to patula Nielson and similar aedeagus and segment 10 processes to unca Nielson but can be distinguished from both species by the long, stout caudodorsal process of the pygofer, which is enlarged distally and trun- cate and serrate on caudal margin, and by the long sinuate caudoventral pygofer process. It lacks the spines on the middle of the inner margin of style in wnca and has a much longer subapical aedeagal spine than patula. Docalidia pennyi, n. sp. Figs. 30-36 LENGTH.—Male, 7.75 mm. Large, robust species. General color brown. Crown deep tan with dark markings; eyes reddish brown; pronotum and scutellum black; forewing brown with broad, deep tan transverse band near middle and along apex. Head short and very broad, narrower than pronotum; crown short, slightly exceeding anterior margin of eyes, broad, about as wide as eyes; pronotum and _ scutellum large; forewing broad; clypeus broad with well- 756 GREAT BASIN NATURALIST Vol. 46, No. 4 Figs. 30-36. Docalidia pennyi, n. sp.: 30, Male pygofer and segment 10, lateral view. 31, Segment 10 and pygofer processes, ventral view. 32, Aedeagus, lateral view. 33, Aedeagus, ventral view. 34, Connective and right style, dorsal view. 35, Style, lateral view. 36, Plate, ventral view. veloped median longitudinal carina; clypellus narrow, lateral margins expanded distally. MALE.—Pygofer in lateral view with two long bladelike processes close together on caudodorsal margin, ventral process narrower (Fig. 30); segment 10 with pair of very long, narrow ventral processes that extend beyond apex of segment 10, minutely dentate on dor- sal margin and at apex (Figs. 30, 31); aedeagus asymmetrical, long, with longitudinal trough medially on ventral margin, apex curved dor- sally in lateral view and with moderately long subapical ventral spine directed basally, spine dentate; gonopore medial on ventral margin (Figs. 32, 33); style very long, nearly reaching apex of aedeagus, narrow, broadly sinuate in lateral view, with numerous spines near dor- sal margin on distal one-fourth (Figs. 34, 35); plate long and broad, with numerous, long, microsetae along inner margin and at apex (Fig. 36). FEMALE.—Unknown. HOLOTYPE (male).—BRAZIL: LeBrea, Amazonas, 27.V.1963. Antonio Carqueira, 2518 (MZUSP). REMARKS.—This species is near dentatula (Metcalf) and can be distinguished by the nar- row style with numerous spines on its distal third. I take pleasure in naming this species for Dr. Norman D. Penny, California Academy of Sciences, who has collected sev- eral new species of coelidiine leafhoppers de- scribed in this and other papers. Docalidia triquetra, n. sp. Figs. 37-43 LENGTH.—Male, 6.15 mm. October 1986 NIELSON: NEW AMERICAN LEAFHOPPERS 757 Figs. 37-43. Docalidia triquetra, n. sp.: 37, Male pygofer and segment 10, lateral view. 38, Segment 10 and pygofer processes, ventral view. 39, Aedeagus, lateral view. 40, Aedeagus, ventral view. 41, Connective and right style, dorsal view. 42, Style, lateral view. 43, Plate, ventral view. Small, robust species. General color brown with large, triangular ochre area on middle of forewing; crown pale; eyes reddish brown; pronotum black with ochraceous bullae; scutellum black; veins of forewings with ochre spots; face dark brown. Head short and very broad, narrower than pronotum; crown short and wide, nearly as wide as eyes, depressed, lateral margins con- vergent basally; eyes large, semiglobular; pronotum short, about as long medially as crown; scutellum large, median length greater than median length of pronotum; forewing long and broad; clypellus long and broad, with prominent median longitudinal carina; clypellus narrow, lateral margins par- allel except for expansion distally. MALE.—Pygofer in lateral view with very long process on caudodorsal margin, process reaching apex of segment 10, asymmetrically flanged laterally on distal third of ventral mar- gin (Figs. 37, 38); segment 10 with short, basally broad ventral process, process gradu- ally narrowed toward apex, apex curved dor- sally to a blunt point (Figs. 37, 38); aedeagus slightly asymmetrical, long, very narrow, compressed laterally at distal two-thirds, broadly curved in lateral view, with very small spine subapically on ventral margin (Figs. 39, 40); gonopore basad of middle on ventral mar- gin; style very long, nearly reaching to apex of aedeagus, ornate at distal half with long lateral process on middle of inner margin, process directed distally and curved dorsally at distal half, distal third of style subtriangular, with lateral teeth on inner margin from base to apex, apex clefted medially, inner part sharply pointed and sclerotized, outer part 758 45 GREAT BASIN NATURALIST Vol. 46, No. 4 50 Figs. 44-50. Docalidia setacea, n. sp.: 44, Male pygofer and segment 10, lateral view. 45, Segment 10 and pygofer processes, ventral view. 46, Aedeagus, lateral view. 47, Aedeagus, ventral view. 48, Connective and right style, dorsal view. 49, Style, lateral view. 50, Plate, ventral view. narrowly rounded and translucent (Figs. 41, 42): plate long and broad with short, microse- tae distally (Fig. 43). FEMALE.—Unknown. HOLOTYPE (male).—BRAZIL: Ro [Rondo- nia], Porto Velho, 11.1X.1965, Epitacio, DPTO Zool., UF Parana (UFP). REMARKS.—This species is similar to fer- riplena (Walker) in male genital characters but can be distinguished by the flanged cau- dodorsal process of the pygofer and by the distal third of the style, which is clefted api- cally with inner side sharply pointed and outer one rounded. Docalidia setacea, n. sp. Figs. 44-50 LENGTH.—Male, 8.10—8.30 mm. Large, robust species. General color deep tan to dark brown throughout; pronotum and scutellum black with numerous small tannish spots; veins of forewings with small suffused spots; face brown to blackish. Head short, very broad, narrower than pronotum; crown short, broad, about as wide as eye, disk depressed in either side of mid- dle, lateral margins parallel except conver- gent basally; pronotum short, little longer me- dially than crown; pronotum large, length greater than length of pronotum; forewing long and broad; clypeus long, narrow, median clypeal carina well developed; clypellus nar- row, lateral margins broad distally. MALE.—Pygofer in lateral view with very small caudoventral lobe, caudodorsal margin with long, narrow process (Fig. 44); segment 10 with poorly developed. ventral process (Figs. 44, 45); aedeagus asymmetrical, long, slightly curved dorsally at distal fourth in lat- eral view, flanged medially on dorsal margin in dorsal view, with long, very narrow ventral spine on middle of shaft (Figs. 46, 47); October 1986 NIELSON: NEW AMERICAN LEAFHOPPERS 759 52 Figs. 51-57. Docalidia gracilitas, n. sp.:51, Male pygofer and segment 10, lateral view. 52, Segment 10 and pygofer processes, ventral view. 53, Aedeagus, lateral view. 54, Aedeagus, ventral view. 55, Connective and right style, dorsal view. 56, Style, lateral view. 57, Plate, ventral view. gonopore basad of middle on ventral margin; style long, not reaching apex of aedeagus, or- nate at distal half, inner lateral margin with broad, distally rounded lobe on middle, di- rected anterio-mesally, with numerous stout setae on inner lateral margin from lobe to subapex of style, apex with narrow, blunt spine directed laterally in lateral view and covered with long microsetae (Figs. 48, 49); plate long and very broad, with few microse- tae on distal third (Fig. 50). FEMALE.—Unknown. HOLOTYPE (male).—BRAZIL: Rondonia, Porto Velho, 1.VI.1979, J. Campbell (MZUSP). Paratype. One male, same data as holotype except 15.III.1979, D. Need (au- thor’s collection). REMARKS.—Docalidia setacea is similar to multispiculata Nielson in setal pattern and arrangement on the style but can be distin- guished by the presence of a broad process on the middle of the inner margin of the style, by the narrow, blunt spine apically on the style and by the narrower aedeagus. Docalidia gracilitas, n. sp. Figs. 51-57 LENGTH.—Male, 7.20 mm, female, 7.30—7.50 mm. Moderate-sized, robust species. General color light brown; pronotum and scutellum dark brown; eyes reddish brown; face brown with black markings. Head short and very broad, slightly nar- rower than pronotum, broadly rounded ante- riorly; crown short, broad, about as wide as eye, depressed, lateral margins sinuate and converging basally; eyes large, semiglobular; pronotum about as long as crown; scutellum large, median length greater than median length of pronotum; forewing long and broad; clypeus long and broad with prominent me- 760 GREAT BASIN NATURALIST Vol. 46, No. 4 63 Figs. 58-66. Docalidia convexa, n. sp.: 58, Male pygofer and segment 10, lateral view. 59, Segment 10 and pygofer processes, ventral view. 60, Aedeagus, distal portion, lateral view. 61, Aedeagus, distal portion, ventral view. 62, Aedeagus, basal portion, lateral view. 63, Aedeagus, basal portion, ventral view. 64, Style, dorsal view. 65, Style, lateral view. 66, Plate, ventral view. dian longitudinal carina; clypellus narrow, lat- eral margins sinuate. MALE.—Pygofer in lateral view with small, narrow caudoventral lobe, caudodorsal mar- gin produced to long, attenuated process (Fig. 51); segment 10 long and narrow with short, inconspicuous ventral lobe (Fig. 51, 52); aedeagus long, very narrow throughout, somewhat compressed laterally, broadly curved in lateral view, with long, very slen- der, subapical process (Figs. 53, 54); gonopore basad of middle on lateral margin; style long, narrowed medially, expanded at distal half on inner lateral margin on dorsal view, with row of setae from middle of inner margin to apex, setae longer, larger, and wider apart at middle, becoming shorter, nar- rower, and closer together toward apex, apex with a short, sharp spine (Figs. 55, 56); plate long and broad as in gracilis Nielson, with some short microsetae near apex (Fig. 57). FEMALE.—Seventh sternum large, about three times as long as preceding segment, caudal margin sinuate, lateral margins with submarginal trough. HOLOTYPE (male).—BRAZIL: Mn. Am. 10.11.1967, Varios 2526 (MZUSP). Allotype female, Amazonas, Manaus, INPA, 30. XII.1977, A. Soares (MZUSP). Paratype, one female, same data as allotype, except 14.V.1979, J. Arias (author's collection). REMARKS.—From gracilis, to which it is similar in male genital characters, gracilitas can be distinguished by the narrow caudoven- tral lobe of the pygofer, by the row of spines on the style that are restricted to the inner October 1986 NIELSON: NEW AMERICAN LEAFHOPPERS 761 Figs. 67-75. Docalidia paracrista, n. sp.: 67, Head, pronotum and scutellum, dorsal view. 68, Face, ventral view. 69, Plate pygofer and segment 10, lateral view. 70, Segment 10 and pygofer processes, ventral view. 71, Aedeagus, lateral view. 72, Aedeagus, ventral view. 73, Connective and right style, dorsal view. 74, Style, lateral view. 75, Plate, ventral view. lateral margin, and by the pointed style with a short apical spine. Docalidia convexa, n. sp. Figs. 58-66 LENGTH.—Male 6.00 mm. Small, slender species. General color deep brown with broad, pale band medially and narrower pale band apically in forewing; head dark brown; pronotum and scutellum nearly black with small, pale spots on surface; veins of forewings with small, pale spots; face brown. Head large, narrower than pronotum; crown narrow, width less than width of eye, not depressed, lateral margins convergent basally; pronotum and scutellum apparently short (pin thrust through them); forewing long, narrow, clypeus narrow, median clypeal carina well developed; clypellus narrow, lat- eral margins expanded distally. MALE.—Pygofer in lateral view with very small caudoventral lobe, caudodorsal margin with long, narrow, bluntly rounded process (Figs. 58, 59); segment 10 with poorly devel- oped ventral process or flange (Figs. 58, 59); aedeagus asymmetrical (broken subbasally), long, narrow with long subapical spine on ventral margin Figs. 60, 61, 62, 63); gonopore medial on lateral margin; style long, about as long as aedeagus, inner lateral margin convex medially with numerous long spines on mid- 762 dle portion, apical third narrowed to rounded apex (Figs. 64, 65); plate long, narrow, with few microsetae distally (Fig. 66). FEMALE.— Unknown. HOLOTYPE (male).—BRAZIL: Mato Grosso, Reserva Humboldt, 10° 11’ S 59° 48’ O, 22.1I1.1977. Norman D. Penny (MZUSP). REMARKS.—This species is near loricata (Osborn) but can be differentiated by the poorly developed ventral processes of seg- ment 10, by the convexity on the middle of the inner lateral margin of the style, and by the lack of spines on the apex of the style. Docalidia paracrista, n. sp. Figs. 67-75 LENGTH.—Male, 7.25 mm. Moderate-sized slender species. General color orange with dark markings on costa and subapical areas of forewings, apical margins bordered with brown; crown pale, eyes translucent pale orange; pronotum and scutel- lum orange; face pale orange. Head large, distinctly narrower than prono- tum (atypical), produced (Fig. 67); crown long, produced, narrower than transocular width, depressed, lateral margins parallel ex- cept convergent near base; eyes large, elon- gate ovoid; pronotum short, median length less than median length of crown; scutellum large, median length greater than length of pronotum; forewing long and narrow with atypical obtusely truncate apex; venation atypical, bases of anteapical cells on same transverse line; clypeus long and narrow, with prominent median longitudinal carina (Fig. 68); clypellus narrow, lateral margins parallel. MALE.—Pygofer in lateral view with small, elongate caudoventral lobe (Figs. 69, 70); cau- dodorsal margin with long, narrow process; segment 10 with long, ventrally depressed flange or process (Figs. 69, 70); aedeagus asymmetrical, long, somewhat tubular and sinuate in lateral view, with long subapical spine arising laterally (Figs. 71, 72); gonopore near middle of shaft on ventral margin; style long, slender, nearly reaching apex of aedea- gus, broad subdistally with numerous, very long spines on inner lateral margin to nar- rowed apex, spines of equal length and closely appressed (Figs. 73, 74); plate long and nar- GREAT BASIN NATURALIST Vol. 46, No. 4 row with short microsetae distally (Fig. 75). FEMALE.—Unknown. HOLOTYPE (male).—BRAZIL: Rondonia, Vilhena, 02.VIII.1983. Norman Penny (MZUSP). REMARKS.—Docalidia paracrista is similar to crista Nielson in male genital characteris- tics but can be distinguished from it by the general habitus characters and by the small, elongate caudoventral lobe of the pygofer and by the presence of a long subapical spine on the aedeagus. ACKNOWLEDGMENTS The material described in this paper, for which I am most grateful, was kindly fur- nished by the following individuals: Dr. Paul Arnaud, California Academy of Sciences, San Francisco (CAS); Dr. James P. Kramer, U.S. National Museum, Washington, D.C. (USNM); Dr. Norman D. Penny, California Academy of Sciences, San Francisco, who col- lected specimens while working in Manaus, Brazil, and who sent material from the Insti- tuto Nacional de Pesquisas da Amazonia (the types of specimens from this institute have been sent to the Museu de Zoologia, Univer- sidade de Sao Paulo [MZUSP]); and Keti Maria Rocha Zanol, Universidade Federal do Parana, Curitiba (UFP). I appreciate very much the review of the paper by Mr. Ray Gill, California Department of Food and Agriculture, Sacramento, and Dr. Bruce Triplehorn, Liberty University, Lynchburg, Virginia, whose suggestions ma- terially improved its contents. I also thank Jean Stanger for the excellent illustrations. LITERATURE CITED NIELSON, M. W. 1979. A revision of the subfamily Coelidi- inae (Homoptera: Cicadellidae) III. Tribe Teruli- ini. Pacific Insects Monograph 35. 329 pp. 1282 figs. ____. 1982a. New species of Brazilian leafhoppers in the genus Docalidia (Cicadellidae: Coelidiinae: Teruliini). Entomography 1: 237—256. . 1982b. New species of leafhoppers of Docalidia from Peru (Cicadellidae: Coelidinae: Teruliini). Entomography 1: 289-302. . 1982c. Some additional new leafhopper species of Docalidia from South America (Cicadellidae: Coelidiinae: Teruliini). Entomography 1:439-445. FOSSIL BIRDS OF THE OREANA LOCAL FAUNA (BLANCAN), OWYHEE COUNTY, IDAHO Jonathan J. Becker’ ABSTRACT.—The Oreana local fauna (Blancan) occurs in expesures of the Glenns Ferry Formation in Owyhee County, Idaho. Fossil birds present include Phalacrocorax cf. P. idahensis, Pelecanus cf. P. halieus, an indeterminate anatid, an indeterminate falconid, two species of Otus, and a species of Colaptes larger than modern C. auratus that provides the earliest record of a colaptine woodpecker. The Oreana local fauna is a Blancan (= Pliocene) assemblage of vertebrate fossils from two localities in southwestern Idaho near the town of Oreana, Owyhee County (IMNH 74001 in Sec 25 and IMNH 74004 (= IMNH 78031) in Sec 1, T4S, R1W; 43 degrees 02’ N Lat., 116 degrees 24’ W Long., Oreana Quad- rangle, U.S. Geologic Survey 7.5 minute se- ries topographical map, 1949). Fossils from both localities come from exposures of the Glenns Ferry Formation (Malde et al. 1963) (= Oreana Formation of Anderson 1965) and correlate with the Hagerman local fauna, ap- proximately 75 miles to the east (Conrad 1980). Smith et al. (1982) discuss the bio- stratigraphy of fishes in this formation. IMNH 74001 has produced many thou- sands of complete, disarticulated skeletal ele- ments of fish, along with a few mammal and bird remains from thick lenses of fine sand interbedded with clays. This locality possibly represents a shoreline with swash accumula- tions (Schaeffer 1972). Vertebrate fossil re- mains are much more rare from IMNH 74004. More detailed information on each locality is available from IMNH upon request. MATERIALS AND METHODS Comparative material of modern species examined is in collections of the American Museum of Natural History (AMNH), the Idaho Museum of Natural History (IMNH), Pierce Brodkorb (PB), and the National Mu- seum of Natural History, Smithsonian Institu- tion (USNM). All fossil specimens from the Oreana local fauna are in the vertebrate pale- ontology collections of the IMNH. Measurements were made with Kanon dial calipers, accurate to 0.05 mm and rounded to the nearest 0.1 mm. BMDP statistical soft- ware program BMDPI1D was used to calculate simple descriptive statistics (Dixon 1981). Computations were made at the Northeast Regional Data Center (NERDC) at the Uni- versity of Florida, Gainesville. Anatomical terminology follows Baumel et al. (1979). SYSTEMATIC PALEONTOLOGY Order Pelecaniformes Sharpe, 1891 Family Phalacrocoracidae (Bonaparte, 1853) Genus Phalacrocorax Brisson, 1760 Phalacrocorax cf P. idahensis (Marsh, 1870) MATERIAL.—IMNH 74001/26527, complete left carpometacarpus; IMNH 74004/30221, proximal end of left ulna; 74004/30223, proxi- mal end of right ulna. Tentatively referred. — 74004/30224, partial upper mandible; 74004/ 30222, proximal end of left ulna; 74001/30217, right scapula. REMARKS.—Although originally described from the Hemphillian Chalk Hills Formation (Marsh 1870), this species is better known from the Blancan Hagerman local fauna (Wet- more 1933, Brodkorb 1958, Murray 1970). The referred upper mandible is short and heavy, having a concave dorsal surface, char- acteristic of the subgenus Phalacrocorax (Howard 1946). The ulnae are within the range of P. idahensis or are slightly larger (Murray 1970). The complete carpometacarpus is larger than any reported by Murray (1970), eliminat- Department of Zoology, University of Florida, Gainesville, Florida 32611. Present address: Division of Birds, National Museum of Natural History, | Smithsonian Institution, Washington, D.C. 20560. 763 764 GREAT BASIN NATURALIST Vol. 46, No. 4 TABLE 1. Measurements of humeri of Otus asio. Data are mean + standard deviation (number) and observed range. Measurements are TWSHAFT, transverse width humeral shaft; DSHAFT, depth of humeral shaft; TWDIST trans- verse width of distal end of humerus; DDIST, depth of distal end of humerus. Otus asio Measurement Males Females IMNH 28411 TWSHAFT 3.19 + 0.25 (7) 2 BLO Se (0,45) (8) 3.3 2.8-3.6 3.1-3.6 DSHAFT 2.76+ 0.22 (7) 2.87 + 0.25 (3) 2.9 2.4-3.1 2.6-3.1 TWDIST 8.66 + 0.59 (7) 9.17 + 0.67 (3) 8.6 7.9-9.6 8.4-9.6 DDIST 4.2 + 0.22(7) 4.90 + 0.70 (3) 4.1 3.9-4.5 4.4-5.7 ing the possibility of its being referable to either of the other cormorants reported from the Glenns Ferry Formation, P. macer and P. auritus , which are smaller. It differs from the carpometacarpus of P. macropus by the square shape of the process of metacarpal I (Murray 1970) and by the intermetacarpal tuberosity being in line with the proximal metacarpal symphysis whereas this is more distad in P. macropus. Family Pelecanidae Vigors, 1825 Pelecanus Linnaeus, 1758 Pelecanus cf P. halieus Wetmore, 1933 MATERIAL. —74001/26528, nearly complete left femur, lacking medial condyle and caudal portion of lateral condyle. 74004/30225, proxi-. mal end of left femur. DESCRIPTION.—IMNH 26528 is from an immature individual and is abraded. Size in- termediate between P. erythrorhynchos and P. occidentalis. Shaft more robust than in P. occidentalis. Neck more distinct, popliteal fossa deeper, undercutting internal condyle, and head angles more proximally than in ei- ther P. occidentalis or P. erythrorhynchos. REMARKS.—This species was named by Wetmore (1933) from the proximal end of a radius. On the basis of this element it was said to be slightly smaller than P. 0. occidentalis but probably closely related to P. ery- throrhynchos. Even with the present mate- rial, the systematic position of this poorly known species cannot be clarified. Order Anseriformes (Wagler, 1831) Family Anatidae Vigors, 1825 Subfamily Anatinae (Vigors, 1825) Genus and species indeterminate MATERIAL. —74001/30504, complete left radius. REMARKS.—This anatid specimen is similar to the radii in females of Aythya collaris but is slightly shorter and more robust. Because the radius is not a diagnostic element in the Anati- dae (Woolfenden 1961:2), I have not identi- fied this specimen beyond the level of subfam- ily. Order Falconiformes cf Falconidae Vigors, 1824 Genus and species indeterminate MATERIAL. —74001/30219, caudal portion of neurocranium. i REMARKS.—Neurocranium preserved from the caudal wall of the orbit caudally, basisphe- noid plate missing. Brain case is somewhat bulbous and prominentia cerebellaris is well developed. This skull fragment shows the greatest similarity to the Falconiformes (ab- sence of trabecular bone precludes assign- ment to the Strigiformes), specifically to the Falconidae. It is similar in size to that of Falco peregrinus. Order Strigiformes (Wagler, 1830) Family Strigidae Vigor, 1825 Subfamily Striginae (Vigors, 1825) Genus Otus Pennant, 1769 Otus cf O. asio (Linnaeus, 1758) MATERIAL.—IMNH _ 74001/28411. Distal end of right humerus. DESCRIPTION.—Similar in size to a male of O. asio naevius. Shape of fossa m. brachialis and the shape and development of the epi- condylus dorsalis within the range of variation of modern populations of O. asio. IMNH 28411 differs from all modern specimens of O. | asio examined in having a shallower fossa ole- | crani. See Table 1 for measurements. | REMARKS.—Ford and Murray (1967) re- ported an indeterminate owl the size of Otus | October 1986 asio from the Hagerman local fauna. The above specimen may represent the same spe- cies. Otus sp. (Kaup, 1852) MATERIAL.—IMNH 74001/30216, nearly complete right tarsometatarsus with caudal portion of trochlea IV missing. DESCRIPTION.—Similar in morphology to female O. flammeolus (USNM 554125) but much smaller (skeletons of males of O. flammeolus unavailable). Caudal projection of process on trochlea IT more developed in fos- sil. Calcaneal ridge not as inclined laterally. REMARKS.—The paucity of skeletons of modern species of small owls makes it impos- sible to determine the exact systematic rela- tionships of this fossil specimen. Order Piciformes (Meyer and Wolf, 1810) Suborder Pici Meyer and Wolf, 1810 Family Picidae Vigors, 1825 Subfamily Picinae (Vigors, 1825) Tribe Colaptini Genus Colaptes Vigors, 1826 GENERIC DIAGNOSIS.—The skull of Co- laptes may be distinguished from other gen- era of New World Picinae by the following combination of characters: (1) Interorbital septum completely ossified (similar to Sphy- rapicus, Campethera, Piculus, Celeus, and Dinopium; incompletely ossified or perforate in Xiphidiopicus, Dendrocopus, Picoides, Ve- niliornis, Dryocopus, Campe-philus, Picus, and Chrysocolaptes; variable in species of Melanerpes); (2) dorsal surface of brain case slightly dimpled (heavily dimpled in Dryoco- pus, Campephilus, Picus, and Chrysoco- laptes; smooth in Melanerpes, Sphyrapicus , and Xiphidiopicus, slightly dimpled in other genera examined); (3) supraorbital ridge present (absent to slightly developed in Melanerpes, Campethera, Dendrocopus, Pi- cus, and Dinopium; present in other genera examined); (4) groove for hyoids present (simi- lar to Campethera, Piculus, Dryocopus, Campephilus, Picus, and Dinopium; absent to slightly developed in other genera examined); (5) frontals flat to concave (similar to Melaner- pes, Piculus, Celeus, Dryocopus, Camp- ephilus, and Chrysocolaptes ; inflated and ex- panded to varying degrees in other genera ) examined, producing a distinct, midsagittal crest in Xiphidiopicus, Campethera, Picus, BECKER: IDAHO FOSSIL BIRDS 765 Din-opium, and some species of Picoides ); (6) interorbital constriction narrow (similar to Celeus and Veniliornis ; wide in Campephilus, Dendrocopus, and Sphyrapicus; intermedi- ate in other genera examined); (7) narrow width between nares (similar to Melanerpes and Dinopium, wide in Sphyrapicus, Den- drocopus, Picoides, Veniliornis, Camp- ephilus, and Chrysocolaptes ; intermediate in other genera examined); (8) basisphenoid re- gion inflated (not inflated in Veniliornis, Dry- ocopus, Campephilus, or Chrysocolaptes , in- flated in other genera examined); and (9) otic region inflated (not inflated in Veniliornis, Campephilus or Chrysocolaptes ); inflated to slightly inflated in other genera. Colaptes sp. MATERIAL. —IMNH 74001/30218, cranium lacking entire upper jaw, pterygoids, and quadrates. DEscCrRIPTION.—IMNH 30218 is distin- guished from Colaptes auratus and C. melanochloros by larger size; from C. campes- tris by having a more developed postorbital process and a deeper, well-defined temporal fossa; from C. pitius by having a more bulbous prominentia cerebellaris and a more caudo- rostrally oriented temporal fossa; and from C. rupicola by being smaller and having more distinct hyoid grooves. Colaptes (=Neso- celeus) fernandinae is very distinct from all other species of Colaptes. In this species the dorsum of the skull is dimpled, hyoid grooves deep, prominentia cerebellaris poorly devel- oped, and the nuchal crest is well developed. See Table 2 for measurements. REMARKS.—North American Neogene woodpeckers include Palaeonerpes shorti Cracraft and Morony 1969, based on a single distal end of a tibiotarsus from deposits equiv- alent to the upper part of the Valentine For- mation (early Clarendonian). Cracraft and Morony (1969) suggest that the affinities of Palaeonerpes are likely to be with the melanerpine woodpeckers rather than with genera such as Dendrocopos, Dryocopus , or Colaptes. Pliopicus brodkorbi Feduccia and Wilson 1967, based on a single distal tarsometatarsus, is from the mid- to late Clarendonian Wa- keeney local fauna (late Miocene) from the Ogallala Formation, Kansas. Feduccia and Wilson (1967) consider Pliopicus to be allied 766 GREAT BASIN NATURALIST Vol. 46, No. 4 TABLE2. Measurements of the crania of species of Colaptes. Data are mean, standard deviation, number measured, and observed range. LENGTH, greatest length from the caudal portion of the supraoccipital (Prominentia cerebellaris ) to the nasofrontal suture, measured on the midsagittal plane (Planum medianum),; DEPTH, depth of skull from dorsal groove for the hyoid to the slight, anterioposteriorly oriented groove in the basitemporal, measured on the midsagittal plane; WIDTH, greatest transverse width brain case; W-TEMPORAL, transverse width of brain case, measured in the temporal fossa immediately caudal to the postorbital process; W-POSTORB, transverse width between postorbital processes; IORB-CONST, narrowest interorbital constriction; W-L[ORBSEPT, transverse width interorbital septum; L-FMAG, anteroposterior diameter of foramen magnum, measured from the caudad projection of the occipital condyle to the rostral surface with the caudal border of the foramen; W-FMAG, greatest transverse width of foramen magnum; W-BULLAE, distance between bullae (i.e., between medial surfaces of O. exoccipitale ala tympanica); COND-SR, occipital condyle to sphenoidal rostrum, measured from caudad portion of occipital condyle to the rostralmost extension of the sphenoidal rostrum; EUSTACIAN, distance between openings of the eustacian tubes; W-BASITEMP, greatest transverse width of basitemporal plate; SR-FO, sphenoidal rostrum to ventral border of foramen opticum; FO-PC, foramen opticum to prominentia cerebellaris. () = specimen damaged. Measurement auratus campestris pitius melanochloros rupicola fernandinae IMNH 30218 LENGTH 29.33 + 0.93 (28) 31.0 31.7 29.0 32.5; 29.7 SIE 34.1 27.5-31.1 DEPTH 17.89 + 0.42 (28) 19.2 19.1 17.2 19.1; 19.2 18.3 19.6 17.15-19.15 WIDTH 22.39 + 0.63 (28) 22.8 23.9 21.1 24.3; 24.0 — 23.6 21.3-23.7 W-TEMPORAL 20.75 + 0.76 (20) 20.2 21.4 18.6 21.6; 21.1 (20.6) 21.4 18.9-22.2 W-POSTORB 21.86 + 0.79 (28) 22:9 23.4 21.0 23.4; 22.9 21.8 2.7 20.4-22.95 Iors-Const _ 8.71 + 0.61 (30) TH 9.2 9.2 Gene Weil (6.7) 7.25-9.8 W-IORBSEPT 1.12 + 0.13 (19) (1.5) (1.7) (1.6) 1.8; 1.7 1.4 1.45 0.95-1.55 L-FMAG 3.94 + 0.16 (28) 4.1 4.4 3.8 4.2: 3.7 — 37 3.9 3.6-4.3 W-FMAG 5.76 + 0.28 (29) 5.9 5.4 5.5 6.5; 6.4 6.0 5.65 5.35-6.3 W-BULLAE 10.77 + 0.44 (29) 12.1 11.4 10.5 11.3; 11.0 11.5 11.35 10.2-12.0 CoND-SR 11.93 + 0.66 (26) 11.4 12.4 10.45 11.9; 12.1 11.6 12.95 10.55-13.3 E,USTACIAN 3.94 + 0.30 (27) 4.5 4.4 4.8 4.6; 3.2 4.5 4.0 3.25—4.75 W-BasireMp _—‘13.91 + 0.51(27) _—(13.0)_~—«(1.9) Gis) CORO) ai.) (14.85) 13.2-14.95 SR-FO 3.84 + 0.27 (27) 4.5 4.2 4.3 3.9; 3.9 4.0 4.55 3.25—-4.55 FO-PC 16.16 + 0.53 (27) 16.5 Wee 15.7 18.0; 17.3 12.4 17.6 15.3-17.3 to Melanerpes. Whereas Cracraft and Morony This specimen may represent either a distinct (1969) reject this suggestion, they could not ally Pliopicus with any other particular group of woodpeckers. Brodkorb (1970) described a fossil species of ivory-billed woodpecker, Campephilus dalquesti, based on a single dis- tal tarsometatarsus, from the Blancan (Pliocene) Beck Ranch local fauna, Texas, and Feduccia (1975) mentioned the occurrence of Colaptes sp. from the Rexroad local faunas, Kansas. With only a single specimen known, the exact systematic relationships of Colaptes sp. from the Oreana local fauna remain uncertain. species or merely a Blancan population of the living Colaptes auratus lineage that was larger. Additional specimens of this species are needed to choose between these alterna- tives. In either case, this record is the earliest known occurrence of a colaptine woodpecker. DISCUSSION The Blancan North American Land Mam- mal Age (= Pliocene) has a diverse fossil avi- fauna, with approximately 90 avian species reported from 16 localities. Localities in the October 1986 Glenns Ferry Formation have produced some 30 of these species (Feduccia 1975 and refer- ences therein, Miller 1944, this study). There are also several unstudied collections of fossil birds from this formation (Becker in prepara- tion). The systematics of many of these species are still poorly known. New species, often only known from fragmentary material, were proposed more on the basis of a presumed difference in geologic time than on quantifi- able differences in morphology. Sexual and geographic variation in the osteology of living species was rarely quantified. Many Blancan species of birds should be critically reexam- ined before being accepted as valid, extinct taxa. ACKNOWLEDGMENTS For the loan of material or for access to collections I thank G. Barrowclough and F. Vuilleumier, American Museum of Natural History; J. A. White, Idaho Museum of Natu- ral History; P. Brodkorb, University of Flor- ida; and S. L. Olson, National Museum of Natural History. I thank P. Brodkorb, S. L. Olson, and D. W. Steadman for their com- ments on this manuscript. LITERATURE CITED ANDERSON, N. R. 1965. Upper Cenozoic stratigraphy of the Oreana 15 minute quadrangle, Idaho. Unpub- lished dissertation, University of Utah, Salt Lake City, BAUMEL, J. J., A.S. Kinc, A. M. Lucas, J. E. BREAZILE, AND H. E. EVANS, (EDS.). 1979. Nomina anatomica avium. Academic Press, New York. 637 pp. BRODKORB, P. 1958. Fossil birds from Idaho. Wilson Bull. 70: 237-242. _ ____. 1970. The paleospecies of woodpeckers. Quart. J. Florida Acad. Sci. 33: 132-136 (published 23 Feb- ruary 1971). ConraD, G. S. 1980. The biostratigraphy and mammalian paleontology of the Glenns Ferry Formation from Hammett to Oreana, Idaho. Unpublished disser- tation, Idaho State University, Pocatello. 351 pp. Dixon, W. J. 1981. BMDP statistical software. University of California Press, Berkeley. 725 pp. Fepuccia, A. 1975. Professor Hibbard’s fossil birds. Univ. Michigan Mus. Paleont., Pap. Paleont. 12: 67-70. FEDUCCIA, A., AND R. L. WILSON. 1967. Avian fossils from the lower Pliocene of Kansas. Occ. Pap. Mus. Zool., Univ. Michigan 655: 1-6. Forp, N. L., AND B. G. Murray, JR. 1967. Fossil owls from the Hagerman local fauna (Upper Pliocene) of Idaho. Auk 84: 115-117. BECKER: IDAHO FOSSIL BIRDS 767 Howarp, H. 1946. A review. of the Pleistocene birds of Fossil Lake, Oregon. Carnegie Instit. Wash., Publ. 551: 142-195, 2 pls. MALDE, H. E., H. A. POWERS, AND C. H. MARSHALL. 1963. Reconnaissance geologic map of west-central Snake River Plain, Idaho: U. S. Geol. Survey Mise. Inv. Map IJ-373. MarsH, O. C. 1870. Notice of some fossil birds, from the Cretaceous and Tertiary formations of the United States. Amer. J. Sci. (2nd Series) 49(146): 205-217. MILLER, L. 1944. Some Pliocene birds from Oregon and Idaho. Condor 46: 25-32. Murray, B. 1970. A redescription of two Pliocene cor- morants. Condor 72: 293-298. SHAEFFER, W. 1972. Ecology and palaeoecology of marine environments. Univ. Chicago Press. 568 pp. SHort, L. L., JR. 1965. Hybridization in the Flickers (Co- laptes) of North America. Bull. Amer. Mus. Nat. Hist. 129(4): 307-428. SMITH, G. R., K. Swirypczuk, P. G. KIMMEL, AND B. H. WILKINSON. 1982. Fish biostratigraphy of late Miocene to Pleistocene sediments of the western Snake River Plain, Idaho. Pages 519-541 in B. Bonnichsen and R. M. Breckenridge, eds., Ceno- zoic geology of Idaho. Idaho Bureau of Mines and Geol. Bull. 26. WETMORE, A. 1933. Pliocene bird remains from Idaho. Smithsonian Misc. Coll. 87(20): 1-12. WOOLFENDEN, G. E. 1961. Postcranial osteology of the waterfowl. Bull. Florida State Mus., Biol. Sci. 6: 1-129. APPENDIX Skulls of the following recent woodpeckers were examined to develop the generic diagno- sis. Melanerpes lewis (1), M. erythrocephalus (8), M. formicivorus (10), M. cruenateus (2), M. pucherani (1), M. chrysogens (2), M. (Chryserpes) striatus (1), M. (Centurus) hy- popolius (1), M. radiolatus (3), M. rubricapil- lus (6), M. uropygialis (4), M. aurifrons (12), M. carolinus (12), M. caymanensis (= super- cilliaris, 3); Sphyrapicus varius (9), S. nuchalis (2), S. thyroideus (2); Xiphidiopicus percussus (1); Campethera bennettii (1), C. abingoni (2), C. taeniolaema (1); Dendropicus fuscescens (3); Picoides arizonae (1), P. minor (1), P. major (1), P. scalaris (4), P. nuttallii (1), P. pubescens (11), P. borealis (9), P. villosus (8), P. trydactylus (1), P. arcticus (2); Venilior- nis fumigatus (2), V. sanguineus (2), V. cassini (2): Piculus flavigula (2), P. rubiginosus (2), P. auricularis (1); Colaptes auratus (31; auratus group—14, chrysocaulosus group—3, cafer group—12, chrysoides group—1; species groups after Short, 1965), C. campestris (1), C. pitius (1), C. rupicola (2), C. melano- chloros (1), C. fernandinae (1), C. (Chrysop- tilus) punctigula (1); Celeus undulatus (2), C. 768 GREAT BASIN NATURALIST Vol. 46, No. 4 castaneus (1), C. elegans (1), C. flavus (1); C. rubricollis (1), Picus canus (1), P. viridis Dryocopus lineatus (3), D. pileatus (10), D. (2); Dinopium javenense (2), D. benghalense martius (1); Campephilus guate-malensis (1), (2); Chrysocolaptes lucidus (1). INDEX TO VOLUME 46 The genera, species, and other taxa described as new to science in this volume appear in bold type in this index. Abronia nana var. harrisii. p. 258. Advertisement call variation in the Arizona tree frog, Hyla wrightorum Taylor, 1938, p. 378. Agathoxylon lemonii sp. nov., from the Da- kota Formation, Utah, p. 559. Agathoxylon lemonii, p. 561. Allred, Darin B., Clive D. Jorgensen, and Richard L. Westcott, article by, p. 173. Ambrosiodmus ferus, p. 269. Ambrosiodmus paucus, p. 269. Andersen, Ferron L., Lauritz A. Jensen, H. Dennis McCurdy, and Craig R. Nichols, article by, p. 208. Anderson, Stanley H., and Kevin J. Gutz- willer, article by, p. 358. Andersen, William R., and Craig E. Cole- man, article by, p. 573. Apple maggot (Rhagoletis pomonella) adapta- tion for cherries in Utah, p. 173. Aquilegia formosa var. fosteri, p. 259. Arabis vivariensis, p. 263. Arthur, W. John, Timothy D. Reynolds, John W. Connelly, and Douglas K. Halford, arti- cle by, p. 513. Astragalus eremiticus var. ampullarioides, p. 262. Astragalus limnocharis var. tabulaeus, p. 261. Astragalus preussii var. cutleri, p. 256. Barn owl diet includes mammal species new to the island fauna of the Great Salt Lake, p. 307. Barnes, James R., and David Ng, article by, p- 310. Barnes, James R., J. V. McArthur, and C. E. Cushing, article by, p. 204. Barr, William F., Michael P. Stafford, and James B. Johnson, article by, p. 287. Bartmann, Richard M.., article by, p. 245. Baugh, Thomas M., John W. Pedretti, and James E. Deacon, article by, p. 441. Baumann, R. W., B. P. Stark, and S. W. Szezytko, article by, p. 383. Beatty, Joseph J., John O. Whitaker, Jr., Chris Maser, and Robert M. Storm, article by, p. 228. Becker, Jonathan J., article by, p. 763. Behan, Barbara, and Bruce L. Welch, article by, p. 161. Biogeographic aspects of leeches, mollusks, and amphibians in the Intermountain Re- gion, p. 736. Biology of Red-necked Phalaropes (Phalaro- pus lobatus) at the western edge of the Great Basin in fall migration, p. 185. Black, Richard D., Jack D. Brotherson, and Lars L. Rasmussen, article by, p. 348. Blake, Elizabeth A., and Michael R. Wagner, article by, p. 169. Blake, Elizabeth A., Robert L. Mathiasen, and Carleton B. Edminster, article by, p. 277. Blockage and recovery of nitrification in soils exposed to acetylene, p. 316. Breeding recores for Clark's Grebe in Colo- rado and Nevada, p. 581. Bright, Donald E., articles by, 641, 679. Brotherson, Jack D., C. Morden, and Bruce N. Smith, article by, p. 140. Brotherson, Jack D., and Lars L. Rasmussen, article by, p. 148. Brotherson, Jack D., Lars L. Rasmussen, and Richard D. Black, article by, p. 348. Brotherson, Jack D., J. Skousen, and J. N. Davis, article by, p. 508. Brotherson, Jack D., and Von Winkel, article by, p. 535. Brown, Barbara A., John O. Whitaker, Jr., Thomas W. French, and Chris Maser, arti- cle by, p. 421. Brown, John A., James A. Young, Raymond A. Evans, and Bruce A. Roundy, article by, D> dks Brusven, Merlyn A., and C. Evan Hornig, article by, p. 33. Bunderson, E. D., D. J. Weber, and D. L. Nelson, article by, p. 427. 769 770 Bunn, Richard L.., article by, p. 581. Burns, Steven J., and Richard E. Terry, arti- cle by, p. 316. Caldwell, Bruce A., C. Y. Li, Chris Maser, and Zane Maser, article by, p. 411. Camissonia atwoodii, p. 258. Canids from the late Pleistocene of Utah, p. 415. Carpenter, Alan T., and Elizabeth E. Neely, article by, p. 728. Carphoborus bicornis, p. 269. Carter, James L., Harry V. Leland, Steven V. Fend, and Albert D. Mahood, article by, p 595. Carter, John, Vincent Lamarra, and Chuck Liff, article by, p. 690. Chaetophloeus pouteriae, p. 269. Characteristics of mule deer beds, p. 542. Chloroplast ultrastructure in the desert shrub Chrysothamnus nauseosus ssp. Albicaulis , p. 973. Cleomella palmerana var. goodrichii, p. 263. Cnemonyx euphorbiae, p. 270. Coggins, Victor, John A. Crawford, Walter Van Dyke, and Martin St. Louis, article by, p. 745. Coleman, Craig E., and William R. An- dersen, article by, p. 573. Coleoptera of the Idaho National Engineering Laboratory: an annotated checklist, p. 287. Comparative habitat and community relation- ships of Atriplex confertifolia and Sarcoba- tus vermiculatus in central Utah, p. 348. Comparison of insects from burned and un- burned areas after a range fire, p. 721. Comparison of vegetation patterns resulting from bulldozing and two-way chaining on a Utah pinyon-juniper big game range, p. 508. Composition and abundance of periphyton and aquatic insects in a Sierra Nevada, Cali- fornia stream, p. 595. Connelly, John W., Timothy D. Reynolds, Douglas K. Halford, and W. John Arthur, article by, p. 513. Corthylus convexifrons, p. 270. Corthylus senticosus, p. 271. Corthylus sentosus, p. 271. Crawford, John A., Walter Van Dyke, Victor Coggins, and Martin St. Louis, article by, p. 745. Crawford, John A., Walter Van Dyke, S. Mark Meyers, and Thomas F. Haensly, ar- ticle by, p. 123. GREAT BASIN NATURALIST Vol. 46, No. 4 Cryptantha cinerea var. arenicola, p. 255. Cryptocarenus pubescens, p. 271. Cryptocarenus spatulatus, p. 272. Cryptogamic soil crusts: recovery from graz- ing near Camp Floyd State Park, Utah, USA, p. 632. Cushing, C. E., James R. Barnes, and J. V. McArthur, article by, p. 204. Cymopterus acaulis var. parvus, p. 79. Dam-raised fawns, an alternative to bottle feeding, p. 217. Davis, J. N., J. Skousen, and Jack D. Brother- son, article by, p. 508. Deacon, James E., Thomas M. Baugh, and John W. Pedretti, article by, p. 441. Deacon, James E., Richard A. Heckmann, and Paul D. Greger, article by, p. 662. Deacon, James E., and Jack E. Williams, arti- cle by, p. 220. Dendrocranulus mexicanus, p. 272. Denning habitat and diet of the swift fox in western South Dakota, p. 249. Diatom flora of Cowboy Hot Spring, Mono County, California, p. 612. Diseases associated with Juniperus os- teosperma and a model for predicting their occurrence with environmental site factors, p. 427. Distributional study of the Zion Snail, Physa zionis , Zion National Park, Utah, p. 310. Docalidia caterva, p. 752. Docalidia convexa, p. 761. Docalidia gracilitas, p. 759. Docalidia paracrista, p. 762. Docalidia pennyi, p. 755. Docalidia setacea, p. 758. Docalidia triquetra, p. 756. Docalidia vella, p. 753. Docalidia vesica, p. 754. Docalidia zanoli, p. 753. Draba kassii, p. 264. Drewes, Henry G., Timothy Modde, and Mark A. Rumble, article by, p. 39. Dunn, Peter O., and Ronald A. Ryder, article by, p. 651. Dynamic landforms and plant communities in a pluvial lake basin, p. 1. Ecological differences of C; and C, plant spe- cies from central Utah in habitats and min- eral composition, p. 140. Edminster, Carleton B., Robert L. Mathi-| asen, aun Jelbezalbetiy A. ‘Blake. article by, p PAT Oc October 1986 Edminster, Carleton B., Robert L. Mathi- asen, and Frank B. Hawksworth, article by, p. 685. Effect of excluding shredders on leaf litter decomposition in two streams, p. 204. Effects of dwarf mistletoe on spruce in the White Mountains, Arizona, p. 685. Effects of suspended sediment on leaf pro- cessing by Hesperophylax occidentalis (Tri- choptera: Limnephilidae) and Pteronarcys californica (Plecoptera: Pteronarcidae), p. 33. Effects of watershed alteration on the brook trout population of a small Black Hills stream, p. 39. Ekins, Laura, and Samuel R. Rushforth, arti- cle by, p. 612. Energy and protein content of coyote prey in southeastern Idaho, p. 274. Erigeron zothecinus, p. 262. Estimates of site potential for Douglas-fir based on site index for several southwest- ern habitat types, p. 277. Evans, Raymond A., James A. Young, Bruce A. Roundy, and John A. Brown, article by, (D5 Ihe Everett, Richard L., article by, p. 706. Fall diet of blue grouse in Oregon, p. 123. Fay, Harlan, C. Y. Li, and Chris Maser, arti- cle by, p. 646. Feduccia, Alan, and Charles G. Oviatt, article by, p. 547. Feeding habits of metamorphosed Am- bystoma tigrinum melanostictum in ponds of high pH (>9), p. 299. Fend, Steven V., Harry V. Leland, James L. Carter, and Albert D. Mahood, article by, p. 595. Findholt, Scott L., article by, p. 128. Floristic analysis of the southwestern United States, p. 46. Foliage age as a factor in food utilization by the western spruce budworm, Choristoneura occidentalis, p. 169. Food habits of clouded salamanders (Aneides ferreus) in Curry County, Oregon (Am- phibia: Caudata: Plethodontidae), p. 228. Fossil birds of the Oreana local fauna (Blan- can), Owyhee County, Idaho, p. 763. Freeman, D. Carl, E. Durant McArthur, and Stewart C. Sanderson, article by, p. 157. | French, Thomas W., Barbara A. Brown, John O. Whitaker, Jr., and Chris Maser, article by, p. 421. INDEX 771 Genetic variation of woodrats (Neotoma cinerea) and deer mice (Peromyscus manic- ulatus) on montane habitat islands in the Great Basin, p. 577. Genter, David L., article by, p. 241. Genus Paralidia with descriptions of new spe- cies (Homoptera: Cicadellidae: Coelidi- inae), p. 329. Germano, David J., and David N. Lawhead, article by, p. 711. Gnatharus, p. 463. Gnatharus tibetensis, p. 463. Goodrich, Sherel, articles by, pp. 66, 366. Gordon, Christine C., article by, p. 166. Grant, C. Val, article by, p. 469. Grazing and passerine breeding birds in a Great Basin low-shrub desert, p. 567. Greger, Paul D., Richard A. Heckmann, and James E. Deacon, article by, p. 662. Groves, Craig B., and Barry L. Keller, article by, p. 404. Growth rates of mule deer fetuses under dif- ferent winter conditions, p. 245. Gutzwiller, Kevin J., and Stanley H. Ander- son, article by, p. 358. Habenaria zothecina, p. 259. Habitat relationships of saltcedar (Tamarix ramosissima) in central Utah, p. 535. Haensly, Thomas F., John A. Crawford, Wal- ter Van Dyke, and S. Mark Meyers, article by, p. 123. Halford, Douglas K., Timothy D. Reynolds, John W. Connelly, and W. John Arthur, article by, p. 513. Hansen, James D., article by, p. 721. Hansen, Richard M., and James G. Mac- Cracken, article by, p. 274. Hatching chronology of Blue Grouse in north- eastern Oregon, p. 745. Hawksworth, Frank G., Robert L. Mathi- asen, and Carleton B. Edminster, article by, p. 685. Heckmann, Richard A., James E. Deacon, and Paul D. Greger, article by, p. 662. Heil, Ken, and Stanley L. Welsh, article by, p. 677. History of fish hatchery development in the Great Basin states of Utah and Nevada, p. 083. Hornig, C. Evan, and Merlyn A. Brusven, article by, p. 33. Hovingh, Peter, article by, p. 736. Huang, Fu-sheng, and Stephen L. Wood, ar- ticle by p. 465. (12 Hydrology of Bear Lake Basin and its impact on the trophic state of Bear Lake, Utah- Idaho, p. 690. Hylesinus caseariae, p. 272. Infection of young Douglas-firs and spruces by dwarf mistletoes in the Southwest, p. 528. Initial survey of acetylene reduction and se- lected microorganisms in the feces of 19 species of mammals, p. 646. Inventory of Utah crayfish with notes on cur- rent distribution, p. 625. Ips orientalis, p. 461. Tris pariensis, p. 256. Isozymes of an autopolyploid shrub, Atriplex canescens (Chenopodiaceae), p. 157. Jehl, Joseph R., Jr., article by, p. 185. Jenkins, Stephen H., and William T. Mewaldt, article by, p. 577. Jennings, William, Loraine Yeatts, and Velma Richards, article by, p. 175. Jensen, Lauritz A., Ferron L. Anderson, H. Dennis McCurdy, and Craig R. Nichols, article by, p. 208. Johansen, Jeffrey R., Samuel R. Rushforth, and Lorin E. Squires, article by, p. 398. Johansen, Jeffrey R., and Larry L. St. Clair, article by, p. 632. Johnson, James B., Michael P. Stafford, and William F. Barr, article by, p. 287. Johnson, James E., article by, p. 625. Jorgensen, Clive D., Darin B. Allred, and Richard L. Westcott, article by, p. 173. Keller, Barry L., and Craig R. Groves, article by, p. 404. Knowles, Craig J., article by, p. 198. Koniak, Susan, article by, p. 178. Lamarra, Vincent, Chuck Liff, and John Carter, article by, p. 690. Larsen, John H., Jr., and Brian T. Miller, article by, p. 299. Lawhead, David N., and David J. Germano, article by, p. 711. Leland, Harry V., Steven V. Fend, James L. Carter, and Albert D. Mahood, article by, p. 595. Lesica, Peter, article by, p. 22. Li, C. Y., Chris Maser, and Harlan Fay, arti- cle by, p. 646. Li, C. Y., Chris Maser, Zane Maser, and Bruce A. Caldwell, article by, p. 411. Life strategies in the evolution of the Colo- rado squawfish (Ptychocheilus lucius), p. 656. GREAT BASIN NATURALIST Vol. 46, No. 4 Liff, Chuck, Vincent Lamarra, and John Carter, article by, p. 690. Lomatium scabrum var. tripinnatum, p. 99. Lupinus argenteus var. moabensis, p. 262. MacCracken, James G., and Richard M. Hansen, article by, p. 274. Madsen, James H., Jr., and Michael E. Nel- son, article by, p. 415. Mahood, Albert D., Harry V. Leland, Steven V. Fend, and James L. Carter, article by, p. 595. Marti, Carl D., article by, p. 307. Maser, Chris, Barbara A. Brown, John O. Whitaker, Jr., and Thomas W. French, ar- ticle by, p. 421. Maser, Chris, C. Y. Li, and Harlan Fay, arti- cle by, p. 646. Maser, Chris, C. Y. Li, Zane Maser, and Bruce A. Caldwell, article by, p. 411. Maser, Chris, John O. Whitaker, Jr., Robert M. Storm, and Joseph J. Beatty, article by, pee: Maser, Zane, C. Y. Li, Chris Maser, and Bruce A. Caldwell, article by, p. 411. Mata, S. A., J. M. Schmid, and J. C. Mitchell, articles by, pp. 445, 449. Mathiasen, Robert L., article by, p. 528. Mathiasen, Robert L., Elizabeth A. Blake, and Carleton B. Edminster, article by, p. OT Mathiasen, Robert L., Frank G. Hawks- worth, and Carleton B. Edminster, article by, p. 685. McArthur, E. Durant, Stewart C. Sanderson, and D. Carl Freeman, article by, p. 157. McArthur, E. Durant, and Bruce L. Welch, article by, p. 281. McArthur, J. V., James R. Barnes, and C. E. Cushing, article by, p. 204. McCurdy, H. Dennis, Ferron L. Andersen, Lauritz A. Jensen, and Craig R. Nichols, article by, p. 208. McLaughlin, Steven P., article by, p. 46. Medin, Dean E., article by, p. 567. Medlyn, David A., and William D. Tidwell, article by, p. 452. Mentzelia cronquistii, p. 550. Mentzelia multicaulis var. librina, p. 556. Mentzelia pumila var. lagrosa, p. 558. Mentzelia shultziorum, p. 361. | Mewaldt, William T., and Stephen H. Jenkins, article by, p. 577. Meyers, S. Mark, John A. Crawford, Walter Van Dyke, and Thomas F. Haensly, article byznpeal 23: October 1986 Microhabitat affinities of Gambel oak seedlings, p. 294. Miller, Brain T., and John H. Larsen, Jr., article by, p. 299. Mitchell, J. C., J. M. Schmid, and S. A. Mata, articles by, pp. 445, 449. Modde, Timothy, Henry G. Drewes, and Mark A. Rumble, article by, p. 39. Montane insular butterfly biogeography: fauna of Ball Mountain, Siskiyou County, California, p. 336. Morden, C., Jack D. Brotherson, and Bruce N. Smith, article by, p. 140. Movements by small mammals on a radioac- tive waste disposal area in southeastern Idaho, p. 404. Neely, Elizabeth E., and Alan T. Carpenter, article by, p. 728. Neese, Elizabeth C., article by, p. 459. Neilson, Ronald P., and L. H. Wullstein, arti- cle by, p. 294. Nelson, D. L., E. D. Bunderson, and D. J. Weber, article by, p. 427. Nelson, Michael E., and James H. Madsen, Jr., article by, p. 415. New genus and species of leafhopper in the tribe Tinobregmini (Homoptera: Cicadelli- dae: Coelidiinae), p. 134. New genus of Scolytidae (Coleoptera) from Asia, p. 465. New oriental genus of leafhoppers in the tribe Coelidiini with descriptions of new species (Homoptera: Cicadellidae: Coelidiinae), p. Sie New records for Montropa hypopithys (Eri- caceae) from Colorado, p. 175. New Pseudoxylechinus (Coleoptera: Scolyti- dae) from India, p. 468. New Sclerocactus (Cactaceae) from Nevada, p. 677. New South American leafhoppers in the genus Docalidia, with a key to 37 species (Cicadelli- dae: Coelidiinae, Teruliini), p. 749. New species and a new combination of Mentzelia section Bartonia (Loasaceae) from the Colorado Plateau, p. 549. New species and new records of North Ameri- can Pityophthorus (Coleoptera: Scolyti- dae), Part VI. The Lautus group, p. 641. New species and new records of North Ameri- can Pityophthorus (Coleoptera: Scolyti- dae), Part VII, p. 679. New species of Mentzelia (Loasaceae) from Grant County, Utah, p. 361. INDEX 773 New species of Protocedroxylon from the Up- per Jurassic of British Columbia, Canada, p. 452. New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), Part XI, p. 265. New taxa and combinations in Utah flora, p. 254. New taxa and nomenclatural changes in Utah Penstemon (Scrophulariaceae), p. 459. New taxa in miscellaneous families from Utah, p. 261. : New thagriine leafhoppers from the Oriental Region, with a key to 30 species (Ho- moptera: Cicadellidae: Coelidiinae), p. 321. New variety of Mentzelia multicaulis (Loasa- ceae) from the Book Cliffs of Utah, p. 555. New variety of Mentzelia pumila (Loasaceae) from Utah, p. 557. Ng, David, and James R. Barnes, article by, p. 310. Nichols, Craig R., Ferron L. Andersen, Lau- ritz A. Jensen, and H. Dennis McCurdy, article by, p. 208. Nielson, M. W., articles by, pp. 134, 1387, 321, 329, 749. : North American stoneflies (Plecoptera): sys- tematics, distribution, and taxonomic refer- ences, p. 383. Note on food habits of the Screech Owl and the Burrowing Owl of southeastern Ore- gon, p. 421. Notes on the birds of Cold Spring Mountain, northwestern Colorado, p. 651. Notes on the Swainson’s Hawk in central Utah: insectivory, premigratory aggrega- tions, and kleptoparasitism, p. 302. Number and condition of seeds in ponderosa pine cones in central Arizona, p. 449. Olson-Rutz, Kathrin M., Philip J. Urness, and Laura A. Urness, article by, p. 217. Oveson, Mark C., H. Duane Smith, and Clyde L. Pritchett, article by, p. 542. Oviatt, Charles G., and Alan Feduccia, article by, p. 547. Paralidia bispinosa, p. 335. Paralidia denticulata, p. 334. Paralidia retrorsa, p. 332. Paralidia singularis, p. 331. Paralidia spinata, p. 329. Parasites of the woundfin minnow, Plagop- terus argentissimus, and other endemic fishes from the Virgin River, Utah, p. 662. 774 Pediomelum aromaticum var. tuhyi, p. 257. Pedretti, John W., Thomas M. Baugh, and James E. Deacon, article by, p. 441. Penstemon angustifolius var. dulcis, p. 459. Penstemon leonardii var. higginsii, p. 459. Penstemon scariosus var. cyanomontanus, p. 460. Penstemon thompsoniae var. desperatus, p. 460. Peterson, Steven R., and Dean F. Stauffer, article by, p. 117. Phacelia demissa var. minor, p. 256. Physaria chambersii var. sobolifera, p. 255. Pityophthorus bravoi, p. 679. Pityophthorus conscriptus, p. 680. Pityophthorus indefessus, p. 641. Pityophthorus inhabilis, p. 642. Pityophthorus levis, p. 273. Pityophthorus ostryacolens, p. 681. Pityophthorus tutulus, p. 643. Pityophthorus vegrandis, p. 643. Ponderosa pine conelet and cone mortality in central Arizona, p. 445. Prigge, Barry A., article by, p. 361. Prigge, B., and H. Thompson, article by, p. 549. Pritchett, Clyde L., H. Duane Smith, and Mark C. Oveson, article by, p. 542. Protocedroxylon macgregorii, p. 452. Pseudopityophthorus peregrinus, p. 462. Pseudoxylechinus, p. 465. Pseudoxylechinus indicus, p. 468. Pseudoxylechinus rugatus, p. 467. Pseudoxylechinus sinensis, p. 467. Pseudoxylechinus tibetensis, p. 466. Pseudoxylechinus uniformis, p. 465. Pseudoxylechinus variegatus, p. 466. Quercus (Fagaceae) in the Utah flora, p. 107. Quercus gambelii var. bonina, p. 108. Quercus havardii, var. tuckeri, p. 109. Rasmussen, Lars L., and Jack D. Brotherson, article by, p. 148. Rasmussen, Lars L., Jack D. Brotherson, and Richard D. Black, article by, p. 348. Relict occurrence of three “American” Scoly- tidae (Coleoptera) in Asia, p. 461. Response of winterfat (Ceratoides lanata) communities to release from grazing pres- sure, p. 148. Reynolds, Timothy D., John W. Connelly, Douglas K. Halford, and W. John Arthur, article by, p. 513. Richards, Velma, William Jennings, and Lo- raine Yeatts, article by, p. 175. GREAT BASIN NATURALIST Vol. 46, No. 4 Roll of three rodents in forest nitrogen fixation in western Oregon: another aspect of mam- mal—mycorrhizal fungus—tree mutualism, p. 411. Roundy, Bruce A., James A. Young, Ray- mond A. Evans, and John A. Brown, article bysapele Rumble, Mark A., Timothy Modde, and Henry G. Drewes, article by, p. 39. Rushforth, Samuel R., and Laura Ekins, arti- cle by, p. 612. Rushforth, Samuel R., Lorin E. Squires, and Jeffrey R. Johansen, article by, p. 398. Ryder, Ronald A., and Peter O. Dunn, article by, p. 651. St. Clair, Larry L., and Jeffrey R. Johansen, article by, p. 632. St. Louis, Martin, John A. Crawford, Walter Van Dyke, and Victor Coggins, article by, p. 745. Sanderson, Stewart C., E. Durant McArthur, and D. Carl Freeman, article by, p. 157. Schmid, J. M., J. C. Mitchell, and S. A. Mata, articles by, pp. 445, 449. Sclerocactus schlesseri, p. 677. Seasonal microhabitat relationships of blue grouse in southeastern Idaho, p. 117. Seasonal phenology and possible migration of the Mourning Cloak butterfly, Nymphalis antiopa (Lepidoptera: Nymphalidae) in California, p. 112. Shapiro, Arthur M., articles by, pp. 112, 336. Sharps, Jon C., and Daniel W. Uresk, article by, p. 249. Sigler, J. W., and W. F. Sigler, article by, p. 583. Sigler, W. F., and J. W. Sigler, article by, p. 583. Sites, Jack W., Jr., and Pamela Thompson, article by, p. 224. Size, structure, and habitat characteristics of populations of Braya humilis var. humilis (Brassicaceae): an alpine disjunct from Col- orado, p. 728. Skousen, J., J. N. Davis, and Jack D. Brother- son, article by, p. 508. Smith, Bruce N., C. Morden, and Jack D. Brotherson, article by, p. 140. Smith, Frank J., and Kaye H. Thorne, article by, p. 555. Smith, H. Duane, Mark C. Oveson, and Clyde L. Pritchett, article by, p. 542. Some relationships of black-tailed prairie dogs _ to livestock grazing, p. 198. ' | October 1986 Species diversity and habitat complexity: does vegetation organize vertebrate communi- ties in the Great Basin?, p. 711. Squires, Lorin E., Samuel R. Rushforth and Jeffrey R. Johansen, article by, p. 398. Stafford, Michael P., William F. Barr, and James B. Johnson, article by, p. 287. Stark, B. P., S. W. Szczytko, and R. W. Bau- mann, article by, p. 383. Status and distribution of California Gull nest- ing colonies in Wyoming, p. 128. Status and distribution of the Fish Creek Springs tui chub, p. 441. Stauffer, Dean F., and Steven R. Peterson, article by, p. 117. Stenolidia, p. 134. Stenolidia magna, p. 134. Storm, Robert M., John O. Whitaker, Jr., Chris Maser, and Joseph J. Beatty, article by, p. 228. | Stylolidia, p. 137. Stylolidia cristata, p. 138. ' Stylolidia pectinata, p. 137. _Subspecific identity of the Amargosa pupfish, Cyprinodon nevadensis, from Crystal Spring, Ash Meadows, Nevada, p. 220. | Sullivan, Brian K., article by, p. 378. | Szezytko, S. W., B. P. Stark, and R. W Bau- | mann, article by, p. 383. | Taylor, Daniel M.., article by, p. 305. |Terry, Richard E., and Steven J. Burns, arti- | cle by, p. 316. Thagria bifida, p. 322. Thagria insolentis, p. 325. | Thagria marissae, p. 326. Thagria melichari, p. 323. Thagria unidentata, p. 327. | Thayn, Gregory F., and William D. Tidwell, | atticle by, p. 559. |Thompson, H., and B. Prigge, article by, p. 549. - |Thompson, Pamela, and Jack W. Sites, Jr., | article by, p. 224. | Thorne, Kaye H., article by, p. 557. | Thorne, Kaye H., and Frank J. Smith, article iby, p. Opp: Three new records for diatoms from the Great Basin, USA, p. 398. | Three-year surveillance for cestode infections in sheep dogs in central Utah, p. 208. Tidwell, William D., and David A. Medlyn, article by, p. 452. Tidwell, William D., and Gregory F. Thayn, article by, p. 559. INDEX 779 Tree densities on pinyon-juniper woodland sites in Nevada and California, p. 178. Trees used simultaneously and sequentially by breeding cavity-nesting birds, p. 358. Trischidias exigua, p. 273. Trumpeter Swan (Cygnus buccinator) from the Pleistocene of Utah, p. 547. Turkey vultures decline at a traditional roost- ing site, p. 305. Two aberrant karyotypes in the sagebrush lizard (Sceloporus graciosus): triploidy and a “supernumerary oddity, p. 224. Tyus, Harold M., article by, p. 656. Understory seed rain in harvested pinyon-ju- niper woodlands, p. 706. Uresk, Daniel W., and Jon C. Sharps, article by, p. 249. Urness, Laura A., Kathrin M. Olson-Rutz, and Philip J. Urness, article by, p. 217. Urness, Philip J., Kathrin M. Olson-Rutz, and Laura A. Urness, article by, p. 217. Utah flora: Apiaceae (Umbelliferae), p. 66. Utah flora: Juncaceae, p. 366. Van Dyke, Walter, John A. Crawford, Victor Coggins, and Martin St. Louis, article by, p. 745. Van Dyke, Walter, John A. Crawford, S. Mark Meyers, and Thomas F. Haensly, ar- ticle by, p. 123. Vegetation and flora of Pine Butte Fen, Teton County, Montana, p. 22. Vertebrate fauna of the Idaho National Envi- ronmental Research Park, p. 513. Wagner, Michael R., and Elizabeth A. Blake, article by, p. 169. Weber, D. J., E. D. Bunderson, and D. L. Nelson, article by, p. 427. Welch, Bruce L., and Barbara Behan, article by, p. 161. Welch, Bruce L., and E. Durant McArthur, article by, p. 281. Welsh, Stanley L., articles by, pp. 107, 254, 261. Welsh, Stanley L., and Ken Heil, article by, Os O07. Westcott, Richard L., Clive D. Jorgensen, and Darin B. Allred, article by, p. 173. Whitaker, John O., Jr., Barbara A. Brown, Thomas W. French, and Chris Maser, arti- cle by, p. 421. Whitaker, John O., Jr., Chris Maser, Robert M. Storm, and Joseph J. Beatty, article by, p. 228. 776 Wildlife distribution and abundance on the Utah Oil Shale Tracts, 1975-1984, p. 469. Williams, Jack E., and James E. Deacon, arti- cle by, p. 220. Winkel, Von, and Jack D. Brotherson, article byapaose: Winter food habits of the pine marten in Colo- rado, p. 166. Wintering bats of the upper Snake River Plain: occurrence in lava-tube caves, p. 241. Wintering mule deer preference for 21 acces- sions of big sagebrush, p. 281. Winter nutritive content of black sagebrush (Artemisia nova) grown in a uniform gar- den, p. 161. GREAT BASIN NATURALIST Vol. 46, No. 4 Woffinden, Neil D., article by, p. 302. Wood, Stephen L., articles by, pp. 265, 468. Wood, Stephen L., and Fu-sheng Huang, ar- ticle by, p. 465. Wood, Stephen L., and Hui-fen Yin, article by, p. 461. Wullstein, L. H., and Ronald P. Neilson, arti- cle by, p. 294. Xenophthorus, p. 462. Yeatts, Loraine, William Jennings, and Velma Richards, article by, p. 175. Yin, Hui-fen, and Stephen L. 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Carter, and Alberta Malroued antes 4 0), chy i Teh velo Waits acat emer ary cle wc eteaen ati Nae One CA i Diatom flora of Cowboy Hot Spring, Mono County, California. Laura Ekins and Samuel Re rshifortlnss eins ee) LS On ML AES ADAG uu Na ee Inventory of Utah crayfish with notes on current distribution. James E. Johnson.... . Cryptogamic soil crusts: recovery from grazing near Camp Floyd State Park, Utah, USA. Jeffrey R. Johansen and'Larry L. St. Clair... 6227000202 2 ee?) New species and new records of North American Pityophthorus (Coleoptera: Scolyti- dae), Part VI. The Lautus group. Donald E. Bright....................... Initial survey of acetylene reduction and selected microorganisms in the feces of 19 species of mammals. C. Y. Li, Chris Maser, and Harlan Fay................ Notes on the birds of Cold Spring Mountain, northwestern Colorado. Peter O. Dunn ang onal dvAvany dense sie en al MOON aie cele nine eae irl ra eae MRT Bam Life strategies in the evolution of the Colorado squawfish (Ptychocheilus lucius). Efaro latte Wyaisei eo: cs in Sis hy eee etek Cue ELEN Sara em Oia aan cba era Parasites of the woundfin minnow, Plagopterus argentissimus, and other endemic fishes from the Virgin River, Utah. Richard A. Heckmann, James E. Deacon, and Pauli) Gregend oonccus soles cg cute lai bra fentimies fahg lag nails aan ares eC a New Sclerocactus (Cactaceae) from Nevada. Ken Heil and Stanley L. Welsh........ New species and new records of North American Pityophthorus (Coleoptera: Scolyti- dae):Part Vill Donaldil Brights vans ccna seis ciauaco ior ayretiwa eae seme ae Effects of dwarf mistletoe on spruce in the White Mountains, Arizona. Robert L. Mathiasen, Frank G. Hawksworth, and Carleton B. Edminster............. Hydrology of Bear Lake Basin and its impact on the trophic state of Bear Lake, Utah-Idaho. Vincent Lamarra, Chuck Liff, and John Carter................ Understory seed rain in harvested pinyon-juniper woodlands. Richard L. Everett. . . Species diversity and habitat complexity: does vegetation organize vertebrate commu- nities in the Great Basin? David J. Germano and David N. Lawhead. ....... Comparison of insects from burned and unburned areas after a range fire. Jena D. FATS OT eau pe nes eee AS ose ocr MN ave Un MAL ela Aaa Ole Sa a eG a 0) co cae ce Size, structure, and habitat characteristics of populations of Braya humilis var. hantelis (Brassicaceae): an alpine disjunct from Colorado. Elizabeth E. Neely ae uBE TE Carpenter icicle heer anait oe aden vee Var ial ire Ra UO a tN tes an NLRC TS eae Biogeographic aspects of leeches, mollusks, and amphibians in the Hiroe ier Region Peters El ovaim clays oni eis cy esis ooianeae ne weMean ey glad ea mentor aN ace iS cate La ae Hatching chronology of Blue Grouse in northeastern Oregon. John A. Crawford, Walter Van Dyke, Victor Coggins, and Martin St. Louis... -.6/../...0.02.. New South American leafhoppers in the genus Docalidia, with a key to 37 species (Cicadellidae: Coelidiinae, Teruliini). M. W. Nielson. .................--. 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