Full Issue, Vol. 56 No. 3

Great Basin Naturalist

Volume 56 Number 3 Article 17

7-26-1996

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VOLUME 56 n2na 3 JULY 1996

BRIGHAM YOUNG university GREAT BASIN naturalist editor assistant editor RICHARD W BAUMANN NATHAN M SMITH 290 MLBM 190 MLBM PO box 20200 PO box 26879 brigham young university brigham young university provo UT 84602020084602 0200 provo UT 84602687984602 6879 8013785053801 378 5053 8013786688801 378 6688 FAX 8013783733801 378 3733 emailE mail nmshbll1byuedu

associate editors MICHAEL A BOWERS PAUL C MARSH blandy experimental farm university of center for environmental studies arizona virginia box 175 boyce VA 22620 state university tempe AZ 85287 J R CALLAHAN STANLEY D SMITH museum of southwestern biology university of department of biology new mexico albuquerque NM university of nevada las vegas mailing address box 3140 hemet CA 92546 las vegas NV 89154400489154 4004 JEFFREY J JOHANSEN PAUL T TUELLER department of biology john carroll university department of environmental resource sciences university heights OH 44118 university of nevada reno 1000 valley road reno NV 89512 BORIS C kondratieff department of entomology colorado state ROBERT C WHITMORE university fort collins CO 80523 division of forestry box 6125 west virginia university Morganmorgantowntown WV 26506612526506 6125

editorial board berranjerran T flinders chairman botany and range science duke S rogers zoology wilford M hess botany and range science richard R tolman zoology all are at brigham young university ex officio editorial board members include steven L taylor college of biology and agriculture H duane smith director monte L bean life science museum richard W baumann editor great basin naturalist the great basin naturalist founded in 1939 is published quarterly by brigham young university unpublished manuscripts that further our biological understanding of the great basinundbasin andund surrounding areas in western north america are accepted for publication subscriptions annual subscriptions to the great basin naturalist for 1996 are 25 for individual sub- scriscribersbers 30 outside the united states and 50 for institutions the price of single issues is 12 all back issues are in print and available for sale all matters pertaining to subscriptions back issues or other business should be directed to the editor great basin naturalist 290 MLBM PO box 20200 brigham young university provo UT 84602020084602 0200 scholarly exchanges libraries or other organizations interested in obtaining the great basin naturalist through a continuing exchange of scholarly publications should contact the exchange librarian 6385 HBLL PO box 26889 brigham young university provo UT 84602688984602 6889

editorial production staff joanne abel technical editor jan spencer assistant to the editor

copyright 0 1996 by brigham young university ISSN 001736140017 3614 official publication date 26 july 1996 7967 96 totso750 19016 the great basin naturalist PUBLISHED AT PROVO UTAH BY BRIGHAM YOUNG university

ISSN 001736140017 3614

VOLUME 56 31 JULY 1996 no 3

great basin naturalist 563 0 1996 appp 191 196 biogeographic significance OF LOW ELEVATION RECORDS FORporforneotomaNEOTOMA CINEREA FROM THE NORTHERN bonneville BASIN UTAH

donald K GraysoGraysongrassonnl1 stephanie D Livingston 2 eric Rickartrickartsrickart33 and monson W shaver 1114

ABSTRACT the existence of low elevation populations of neotoma cinerea in the northern bonneville basin shows either that these mammals can survive many thousands of years in xericxene habitats or that they can move across bencxericxenc lowlands far more readily than has been appreciated or both current models of great basin small mammal biogeography are far too static to encompass properly the interaction of the wide range of geographical and biological variability that has produced the modern distribution of those mammals that have for several decades been treated as montane within the great basin

key words great basin biogeography island biogeography neotoma cinerea mammals

ever since J H brown s insightful analyses coherence that has been assigned to it here of great basin small mammal biogeography we add to that growing body and call for a brown 1971 1978 see also lomolino et al more dynamic view of great basin small mam- 1989 geographersbiogeographersbio have treated the bushy mal historic biogeography tailed woodratwoodral neotoma cincinereaoincineredmineredered as a member of an assemblage of small mammals that is cur- NEOTOMA CINEREA ON rently isolated on great basin mountains the HOMESTEAD KNOLL UTAH composition of this assemblage is of particular importance because it has been used to gener- located a few km west and south of great ate and test hypotheses about the past and salt lake in north central utah the lakeside future of great basin montane mammals eg mountains are formed from a complex of north grayson 1987 1993 patterson 1990 cutler trending hills ridges knolls and small moun- 1991 mcdonald and brown 1992 murphy and tains fig 1 the northwestern most spur of weiss 1992 grayson and livingston 1993 this complex is homestead knoll a low maxi- however there is a growing body of data that mum elevation 1615 m rocky promontory suggests that this group of mammals lacks the that is devoid of active springs and permanent

iburkeaburkeurke memorial museum box 353010 university of washington seattle WA 98195 quaternary sciences center desert research institute box 60220 reno NV 89506 sutah museum of natural history university of utah salt lake city UT 84112 4uauutahh geological survey 2363 south fooothill drive salt lake city UT 84109

191 192 GREAT BASIN naturalist volume 56

M s F t

0 0 greatgreot

homestead CD cavecove saltsolt

lakeloke z

0 N A 35 cl KM IF fig 1 location of homestead cave within the northern bonneville basin

streams and that is separated from other parts homestead knoll is dotted by a number of of the lakeside group by valleys whose maxi- caseseavescaves one of which homestead cave sits on mum elevations do not exceed 1465 m the northwestern edge of the knoll at an eleva- the barren playa of pleistocene lake bon- tion of 1406 ra fig 2 approximately 11 m neville is located to the immediate west and wide and 6 m high at its mouth this 25 m northwest of homestead knoll vegetation of deep eavecavepavegave has since 1992 been the focus of the knoll is dominated by shrubs and grasses interdisciplinary paleopaleoecologicalecological work funded although there are a few scattered utah junipersjunijuniperuspers by the department of defense with D B juniperus osteosperma on its highest reaches madsen of the utah geological survey 3 most prominent among the shrubs are authors of this paper DKG SDL and MWS atriplex confertifoliaconfertifolia tetradymia spinosa and have been involved with the excavation and tetradymia glabrataglabrateglabrata artemisia dentatatritridentatatridentate is analysis of a deep sequence of vertebrate present along seasonally moist drainages while remains from this site to provide background artemisia spinescensspinescentspinescens chrysothamnus sp and data for the analysis of the mammalian compo- sarcobatus vermiculatus are present but un- nent of the excavated faunahaunafaunal we conducted a common above the flanks of the knoll artemisia brief 270 trap night small mammal survey in nova occurs on those flanks as does Ceratceratoidesoides the vicinity of homestead cave in june 1995 lanata while S venniculatusvermiculatus becomes increas- with 1 exception the results of this survey ingly abundant as the valley bottoms are were quite predictable trapping success was approached we made no attempt to identify low with 3 species dipodomys ordiiordia 3 indi- the grasses that form the understory beneath vidualsvi peromyscus maniculatus 11 individu- the shrubs but cheatcheatgrassgrass bromus tectoriumtectorumtectorum als and neotoma lepidacepida 6 individuals com- and perhaps B rubens is extremely abundant prising nearly the entire trapped assemblage on the flats beneath the knoll the 1 exception however was remarkable we 19961 low ELEVATION neotoma CINEREA 193

1 1 111n41 22 1 1 14 rl51 1 L

C 0

aw7w N 3

v d W 4 17

4 ob

fig 2 location of homestead cave white arrow on homestead knoll the prominent terraces represent provo post provo regressive and stansbury beaches left by the waters of pleistocene lake bonneville

took a single neotoma cinerea from the back of lection at the utah museum of natural his- homestead cave itself tory university of utah for additional records because this individual was live trapped of this species from other low elevation set- and released we cannot report its age or sex or tings in the northern bonneville basin we provide standard measurements even though were quite successful in this search we do not have a voucher specimen we do a locomotive springs the only previously have an excellent videotape of the animal taken published low elevation record for neotoma by MWS and on file at the utah geological cinerea for the northern bonneville basin was survey and there is no doubt as to the identi- provided by durrant 1952348 UU 5048 as ficationfi of the individual having been taken in october 1947 from state- of vegetation in the immediate vicinity house locomotive springs 5500 ft 1676 in homestead cave departs from the homestead however we are unable to determine the loca- have knoll vegetation that we described in tion of statehouse and are otherwise hesitant only 1 the mouth of the cave sup- I major way to accept this record because of the substantial growth of ports a luxuriant ribes cereuecereum difference between the actual elevation of immediately beneath the triplinedriplinedripline it would be locomotive springs 1283 and the reported surprising if this shrub were not heavily uti- in elevation of statehouse 1676 given the lized by both neotoma cinerea taken at the in well nature of springs rear of the cave and neotoma lepidacepida taken at watered locomotive the front the record might be accurate but it is in need of verification locomotive springs is approxi- OTHER LOW ELEVATION NORTHERN mately 60 km north of homestead knoll an adult male neo- bonneville BASIN RECORDS b lakeside mountains FOR NEOTOMA CINEREA toma cinerea UU 14374 was collected 5 mi E lakeside 4600 ft 1402 in in june 1957 our discovery of neotoma cinerea on home- this distance and direction from lakeside how- stead knoll led us to search the mammal col ever describe a point in the great salt lake if 194 GREAT BASIN naturalist volume 56 the actual direction were southeast the speci- basin during those years we can however do men could have come from cave ridge on the much more than speculate about the history of eastern edge of the lakeside mountains N cinerea in the homestead knoll area approximately 10 km east of homestead knoll with a maximum elevation of 1615 m home- c newfoundland mountains A series of stead knoll was covered by the waters of pleis- three juvenile neotoma cinerea UU 9995 tocene lake bonneville 14500 years BPBE when 9996 9998 were collected in june 1951 from pleistocene lake bonneville was at its high an unspecified site at the north end of the see figure 2 obviously homestead knoll newfoundland mountains the collector s field must have received its woodratswoodrats after this time notes do not provide the elevation of the site but when this occurred is not clear between but do indicate that the specimens came from 14500 and at least 14200 years BPBE when lake an area of granite cliffs with a plant community bonneville stood at the provo level home- that included juniperus and tetradymia the stead knoll was an island of approximately 770 north end of the newfoundland mountains is acres not until lake bonneville fell to a local approximately 40 km west northwest of home- elevation of 1463 m did this island become stead knoll connected to the main body of the lakeside d cedar mountains there are records for mountains once this occurred homestead neotoma cinerea from 2 separate locations in knoll became part of the faunal mainland and the southern cedar mountains 4 from the would have been open to overland colonization cane springs area elevation 1768 m UU by terrestrial mammals 26340 27297 27299 and 273012730122 collected unfortunately we do not know when the between october 1952 and january 1953 and lake fell to this level however we do have I1 from the south end cedar mtnman 4850 ft direct evidence from homestead cave con- 1478 m this last specimen is reported to cerning the regional history of neotomaNeotorna cinerea have been caught in a garage suggesting that it excavations in this cave have provided a rich may have come from near dugway although stratified sequence of vertebrate remains the these specimens come from no closer than 95 mammals of which are being identified and km to the south of homestead cave we men- analyzed by one of the authors DKG to date tion them because they establish the likelihood a substantial sample of mammal specimens that neotoma cinerea occurs in suitable habitat from the 4 lowest homestead cave strata has throughout the cedar range been identified 37381 specimens all 4 assemblages contain both N cinerea biogeographic considerations and N lepidacepida but the ratio of N cinerea to N lepidacepida varies dramatically through time in although neotoma cinerea has frequently stratum I1 which dates to between ca 11300 been treated as being isolated on great basin and 10000 years BPBE bushy tailed woodratswoodrats mountains brown 1971 1978 grayson 1993 make up 993899.38 of the neotoma fauna in sub- these records demonstrate that bushy tailed sequent strata however they decline steadily woodratswoodrats can and do exist at low elevations in in abundance by stratum IV ca 8200 7200 andaridarld contexts within at least the northern bon- years BPBE N cinerea comprises only 4744.74 of neville basin how one must wonder did the neotoma assemblage fig 3 similarly N neotoma cinerea come to occupy such arid cinerea contributes 239723.97 of the total number low elevation settings as the newfoundland of identified mammalian specimens in stratum mountains maximum elevation 2130 m and I1 a number that declines to 1011.01 in stratum isolated knolls on the lakeside mountains IV fig 4 maximum elevation 2020 m the homestead cave fauna thus documents it is well established that during the late that N cinerea was present in the homestead pleistocene bushy tailed woodratswoodrats were far knoll area by 11300 years BPBE and remained a more widely distributed within the great basin common species in the small mammal fauna than they are today occupying low elevation through much of the early holocene after ca settings where they are no longer found gray- 8200 years BPBE however N lepidacepida became the son 1988 1993 As a result it is reasonable to overwhelmingly dominant member ofthe genus speculate that these animals were also wide- and N cinerea became locally rare since mam- spread in this part of the northern bonneville mals from later strata within homestead cave 199611996 LOW ELEVATION NEOTOMA CINEREA 195

N 1933 35 9938 loo100 N 1564 30 9- 08613 N 1912 2397 80 25 N 1347 1 70 2083 w 20 60 w N 564 z 04681 0 15 N 264 5- F Z 1109 40 w 10 w 30 w M 5- 8 N 208 20 N 4392 loi101 10- 0 474 0 III111 0 PIREIM liililill IV

I1 STRATUM iliIII111 IV STRATUM fig 4 changing contribution of neotoma cinerea to the fig 3 changing contribution of N cinerea to the total number of identified mammalian specimens NISP neotoma N cinerea plus N lepidacepida fauna homestead per stratum at homestead cave strata I1 IV cave strata I1 IV N total number of neotoma specimens identified to the species level including those identified as N cf cinerea and N cf lepidacepida implications have not yet been completely identified we do the discovery of neotoma cinerea on home- not know whether N cinerea survived the very stead knoll does not simply represent an unex- xeric middle holocene ca 7500 5000 years pected natural historical tidbit our discovery BEBY here documents either that populations of neotoma currently there are 2 options for explaining cinerea within the great basin can find suffi- the modern existence ofofnoanN cinerea on home- cient refuge in low elevation xenexeric habitats to stead knoll first animals living here today survive for many thousands of years or that may be direct descendants of the initial woodratwoodral this species can move across xericxene lowlands far colonizers of the knoll colonizers that arrived more readily than has been appreciated or sometime between 14500 and 11300 years BEBY both indeed insofar as bushy tailed woodratswoodrats if so the population has survived even though are more effective colonizers than has been its numbers dropped precipitously toward the realized an effective parallel may exist in the end of the early holocene ca 8200 7200 years yellow nosed cotton rat sigmodonSigsigmodontmodon ochrogna BPBE and presumably fell even further during thus a montane mammal of the southwest the heart of the middle holocene assuming that has apparently expanded its range across that N cinerea does not now survive in the val- low elevation valleys during the past 50 years leys that separate homestead knoll from nearby davis and dunford 1987 see also davis and uplands and that it has not been able to sur- callahan 1992 on microtus mexicanusmexicanus vive in those valleys since at least 7000 years elsewhere grayson and livingston 1993 BEBY then this population has existed on an iso- have noted that sylvilagusSylvilagus nuttalnuttalhinuttafflihi can cross lated upland a few thousand acres in extent for valley bottoms in at least parts of the great a minimum of some 7 millennia basin now it seems that N cinerea can sur- the other and certainly more likely option vive in habitats that are anything but montane is that neotoma cinerea has not been isolated this fact leads us to suggest that the nested on homestead knoll for this entire period of ness of great basin mammal faunalfaunas gensusensu pat- time that populations on the knoll have been terson and atmar 1986 patterson 1987 1990 augmented by immigrants from elsewhere and might reflect a combination of extinction histo- that any local extinctextinctionsions of N cinerea on the riesrles and colonization abilities in addition the knoll have been followed by recolonizationsrecolomzations homestead knoll record for N cinerea takes from nearby populations indeed it is even its place alongside other recent data docu- possible that the current representatives of the menting that current models of great basin species colonized homestead knoll during the small mammal biogeography are far too static mid 1980s a time of extraordinarily high pre- to encompass properly the wide range of geo- cipitation in the northern great basin arnow graphical and biological variability that has and stephens 1990 produced the modern distribution of those 196 GREAT BASIN naturalist volume 56 mammals that for several decades have been flying mammals in the american southwest ameri- treated as montane within the great basin can naturalist 129 398 406 craysongrayson 1993 grayson and DURRANTDURBANT S D 1952 mammals of utah taxonomy and eg livingston distribution university of kansas publications 1993 lawlor 1995 rickart 1995 in the south- museum of natural history 6 west modern montane mammal distributions GRAYSON D K 1987 the biogeographic history of small have clearly been determined by a complex mammals in the great basin observations on the last combination of holocene extinctextinctionsions and 20000 years journal of mammalogy 68 359 375 1988 danger cave last supper cave hanging colonizations eg davis and dunford 1987 rock shelter the faunalfaunas american museum of nat- lomolino et al 1989 davis and callahan 1992 ural history anthropological papers 66 it now appears that the situation in the great 1993 the deserts past a natural prehistory of the great press basin is quite similar basin smithsonianSmith soman institution washington DC GRAYSON D K AND S D livingston 1993 missing acknowledgments mammals on great basin mountains holocene extincextine tionseions and inadequate knowledge conservation biol- the research reported here was supported ogy 7 527 532 by a grant homkomfromheom the US department of defense LAWLOR T 1995 biogeography of great basin mammals program paradigm lost unpublished manuscript on file at the legacy project 0304843028x728 department of biological sciences humboldt state environmentalPaleopaleoenvironmental change on hill air force university arcata CA base and dugway proving grounds our LOMOLINO M V J H BROWN AND R DAVIS 1989 island thanks to D B madsen for assistance at all biogeography of montane forest mammals in the american southwest ecology 70 180 194 stages of this project and to R S thompson for MCDONALD K A AND J H BROWN 1992 using mon- confirming identifications we also plant thank tane mammals to model extinctionsextinctions due to global R davis D B madsen B D patterson and change conservation biology 6 409 415 T A vaughan for helpful comments on a draft MURPHY D D AND WEISS S B 1992 effects of climate of this paper change on biological diversity in western north america species losses and mechanisms pages 355 368 in R L peters and T E lovejoy editors literature CITED global warming and biological diversity yale university press new haven CT ARNOW 1 AND D SILPIILNSSTEPHENS 1990 hydrologic character- PATTERSONPAITEKSON B D 1987 the principle of nested subsets istics of the cleatcreatgleatgreat salt lake utah 1847 1986 US and its implications for biological conservation con- geological survey watelwater supply paper 2332 servation biology 1 323 334 BROWN J H 1971 mammals on mountaintopsmountain tops nonequibonequi 1990 on the temporal development of nested librilibnumlinnumlibriurnurn insular biogeography american naturalist subset patterns of species composition oikosbikos 59 105467105 467 478 330 342 1978 the theory of insular biogeography and the PATTERSON B D AND W ATMAR 1986 nested subsets distribution of boreal birds and mammals pages and the structure of insular mammalian faunalfaunas and 209 227 in K T harper and J L reveal editors archipelagosarchipelagoes biological journal of the linnean soci- intermountainintelinter mountain biogeography a symposium great ety 28 65 82 basin naturalist memoirs 2 RICKART E A 1995 elevational diversity gradients bio- CUILLR A 1991 nested faunalfaunas and extinction in frag- geography and the structure of montane mammal mented habitats conservation biology 5 496 505 communities in the intermountain region unpub- DAVIS R AND J R CALLAHAN 1992 post pleistocene dis- lished manuscript on file at the utah museum of persal in the mexican vole microtus mexicanusmexic anus an natural history university of utah salt lake city example of an apparent trend in the distribution of southwestern mammals great basin naturalist 52 received 15 november 1995 262 268 accepted 20 march 1996 DAVIS R AND C DUNFORD 1987 an example of contem- porary colonization of montane islands by small non great basin naturalist 563 0 1996 appp 197 204

SYNOPSIS OF THE MOSSES OF WYOMING

PPMA eckell

ABSTRACT A revised list of the mosses of the state of wyoming is presented recorded are 315 species and varieties

key words bryophytes wyoming porter rocky mountains checklist flora

publication of the mosses of wyoming began I1 was able to locate and borrow for study some with aven nelson 1900 listing 119 species of mr smiley s specimens several were deter- his collections were made essentially in the mined by R S williams laramie and medicine bow ranges of carbon no checklist focusing on the moss flora of and albany counties and his specimens are yellowstone national park has as yet been pub- now at the rocky mountain herbarium RM lished the 357 or so moss specimens auratedcurated neiNelnelsonneisonsonss specimens were determined in large at YELLO all derive from the park the fol- part by professor john M holzinger of the lowing are collectors and dates of collecting state normal school at winona minnesota activity dwight smiley 1932 H S conard aven nelsonneison s and elias neisnelsonneisonNelsolsoyss wyoming col- 1948 winona welch 1951 eula whitehouse lections were distributed under printed labels 1951 elva lawton 1953 no biographical data as plants of wyoming from the rocky moun- exist on mr smiley at YELLO whipple per- tain herbariurrherbarium and plants of yellowstone sonal communication I1 have no other record national park from the rocky mountain herb- that these collectors worked in wyoming but arium citing holzinger and a few others as my use of various herbaria for this checklist is determiners however because they were not exhaustive incidentally there are 11 pack- issued without serial numbers showing only ets of liverwortsliverworts at YELLO and no representa- the collectors numbers together with other tion of sphagnum data they do not constitute true exsiccatae recent collectors in the state include the sayre 1971 late fredrick hermann holmes rolston and in 1935 cedric lambert porter published a william weber all of whose wyoming speci- valuable checklist of the mosses of the state of mens are distributed in various herbaria espe- wyoming citing 215 species and varieties in cially COLO steven churchill 1979 1982 this publication porter mentioned a paper by john spence 1985 and alvin L medina 1994 dwight C smiley on the mosses of yellow- collected additional taxa two taxa listed here stone national park which porter cited again as new to the state were recently collected by two years later as A key to the mosses of yel- joseph elliott scorpidium scorpioscorpioidesscorpioideaides and lowstonelowstone national park unpublished porter cinclidium stygiumstygium general references to ex- 1937 porter included smiley s names but tensive collections made in the state are given apparently did not examine his specimens by lawton 1971 the substance of a dissertation written by the purpose of this paper is to present an porter at the university of washington in 1937 up to date list of the mosses of the state of was the development of a useful key to the wyoming incorporating reports published hepatic and bryophyte taxa of wyoming it since porter s manuscripts as well as additional includes additional county records a few addi- unpublished information there are 315 species tional species and references to yellowstone and varieties in the present list some idea of national park again apparently citing smiley s the degree of representation of the flora of unpublished material on a recent visit to the wyoming comprising this list may be inferred yellowstone national park herbarium YELLO from a glance at similar checklists for other

clintonlclinton herbarium buffalo museum of science buffalo NY 14214

197 198 GREAT BASIN naturalist volume 56 areas it is probably to be expected that the calhergoncalliergonCalhCallihrgonergon cordicordifoliumcordifohumfolium hedw kindbkinda TENN andaridarld intermountain states such as utah and calliergonCalliergon gigantgiganteumeum schimp kindbkinda NY CalhCalli will calliergoncalhergonergon richardsonchardsonnrichardsoniarichardsoniinii mitt kindbkinda vnin earnstwarnst lawton nevada have a more depauperate flora 1971 spence 1988 recorded 342 species for the calliergoncalhergonCalhCalliergon stramineum brid kindbkinda COLO RM entire intermountain west the following calliergoncalhergonCalliCalhergon tnfanumtriforiumtrifariumtrifarium web & mohr kindakindb cooper and counts are for species and varieties except andrus1994andrus 1994 Callier VVTU utah which is species only arizona 381 calliergonellagonella cuspidatecuspidatacuspidata hedw loeske WTU campyliumCampyca puliumpyliumlium chrysophylluchrysophyllum brid J lange bueBUEBUF RM johnsen no date colorado 292 weber 1973 Campycampyliumlium hispidulumhispidulum brid mitt NY idaho 257 mccleary and green 1971 mon- Campycampyliumlium polygampolygamumum schimp in BSG C jens RM tana 358 eversman and sharp 1980 nevada Campycampyliumlium stellatumstellatum hedw C jens RM 165 lawton 1958 lavin 1981 oregon 441 Conardia compactscompactacompacta C muell robins NY WTU gratoneuronfilicinum hedw spruce NY RM al 1982 cratoneuronfilicinum christy et utah 256 flowers 1973 drepanocladus aduncusaruncus hedw earnstwamstwarnst var aduncusaruncus RM there appears to be no checklist for the state var kneifkneifffilii schimp in BSG moenk RM of washington new york state which is said var polycarpuspolycarpmpolypolycarpouscarpus bland ex voit roth COLO RM to have a diverse moss flora has 503 species drepanocladus capillifoliuscapillifolius Warn st earnstwarnst RM drepanocladus crassicostatuscrcissicostatus janssens CANM and varieties ketchledge 1980 A strikingstrike1119 lilg drepanocladus sendtnersendtnensendtnefisendtnennefineninenn schimp earnstwarnst BING comparison is to the oceanic island of new- hygroamblystegium noterophilum sull & lesqfesq in sull foundlandfoundland which boasts a moss flora of 445 earnstwarnst RM species brassard 1983 and which is geophisgeophys hygroamblystegium tanaxtenax hedw jenn RM hygrohypnum bestilbestnbesan ren & ren WTU iographically rather plain compared to the geo- bryhn in broth hygrohypnum dunusculumduriusculum de not jamieson CSU morphic extremes and diversity of wyoming in hygrohypnum lundumluridumalundum hedw jenn NY RM the central rocky mountains hygrohypnum molle hedw loeske RM WTU the following checklist is based largely on a hygrohypnuhygrohypnum ochraceum turn ex wils loeske BUF review of specimens housed in the rocky CSU RM hygrohypnum smithnsmithiismithia sw in liljli1jhilj broth RM mountain herbarium additional herbaria were leptodictyum cumilehumile P beaucbeauv ochyraoshyra RM contacted in instances of taxa reported in the leptodictyum ripariripariumum hedw earnstwarnst NY RM literature but with no representation at RM and limpnchtialimptichtia revolversrevolrevolvensvens sw loeske BING BUF numerous new records have been added from palwtriellapalustnella commutatecommutata brid oshyraochyra RM WTU palustnellapalustriella decidecipiensdecvpienspiens not ochyraoshyra RM WTU field collections by myself and others I1 have de pseudocalliergon turgescentturgesturgescenscens T jens loeske COLO RM attempted to cite at least one reliable specimen Sasanioniasaniomasanconianionia uncinata hedw loeske NY RM of each taxon by giving the abbreviation of the Sarmentsannenthypnumsarmenthypnumhypnum sannensannentosumsarmentosumsarmenttosumosum wahlenewahlenb tuom &cc I1 kop herbarium at which the specimen is located BING COLO RM scorpidium scorpioideascorpioidesides additional taxa are added from porter s 1937 scorpio hedw limarlimpr BUF wamstorfiawarnstorfia exannulata schimp in BSG loeske dissertation if they did not occur in his previ- var exannulata CSU RM ous publication if a herbarium designation is wamstorfiawarnstorflafluitansflwtans hedw loeske COLO RM noted below for a taxon no further reference to the literature is given although several ref- andreaeaceae to the same species throughout the lit- erences andreaea rupestrisrupestnsrupestris hedw YELLO erature may have been cited only one citation is presented for names for which specimens aulacomniaceaeaufacomniaceae have not been seen nomenclature other than sphagnum follows aulacomnium androgynusandrogynum hedw schwaegrSchwaschwaegeregr TENN aulacomnium palustre hedw Schwaschwaegrschwaegeregr BUF RM anderson et al 1990 sphagnum nomencla- TENN ture follows anderson 1990 families are given alphabetically genera bartramiaceae alphabetically within families and alpha- species par beticallycally anaceliaanacolia menziemenziemmenziesiimenziesiasiisilsll turn WTU beti within genera bartramia ithyphylla brid RM philonotisfontanaPhilophilonotisnotisnotts fontana hedw budbrid varfontanavarvan fontana bueBVFBUF RM CHECKLIST OF MOSSES OF WYOMING var americanaamenamencanacanaoanacuna cismdism flow BUF RM var caespitosa jur schimp bueBUEBUF RM amblystegiaceae var pumila turn brid RM amblystegium serpens hedw schimp var serpens NY brachytheciaceae RM varjuratzkanum schimp rau & heahem RM brachythecium acutumscutum mitt sull RM amblystegium cariumvatiumvarium hedw lindblinda NY RM brachythecium albicans hedw schimp in BSG RM 199611996 MOSSES OF WYOMING 199 brachythecium collinumcollinum schielSchleischleichcb ex C muell schimp buxbaumiaceaeBUXBAUMIA CEAF in BSGBSC bueBUEBUF NY RM brachythecium erythrorrhizon schimp in BSG RM buxbaumia aphyllaaphylly hedw WTU brachytheciumfendletbrachythecium fendlenbendlen sull jaegcaeg RM buxbaumia vindis DC moug & nesti WTU brachytheciumbrachytheciumfrigidumfngidum C muell besch NY US brachythecium leiicgii16l roiirgii grout NY RM US climaciaceae brachythecium nelsoniinelsonelsonianii croutgroutgroutlrgrouterI1 SU NY RM brachythecium oedipodium mitt taeg COLO RM US climacium americanumameriamerlamencanumamencanum brid BING BUF brachythecium rivulanvularerivularere schimp in BSG NY RM US climacium dendroiddendroidesdendroideaes hedw web & mohr COLO brachythecium rutarutabulumbulum hedw schimp in BSG NY CSU RM brachythecium salebrosum web & mohr schimp in climacium kindbergkindbergiiii ren & card grout porter 1935 BSG bueBUEBUF RM US brachythecium starkelstarkei brid schimp in BSG porter 1935 dicranaceae brachythecium turgiturgidumdum CJC J hartmharam kindbkmdbkinda COLO NY RM US Campycampylopuslopus frasrafragihsfragiasfragilisgilis brid bruch & schimp in BSG brachythecium velutinum hedw schimp in BSG WTU var velutinum spence 1985 cynodontium alpestrealpestre wahlenbwahlene milde lawton 1971 var venushimvenustumvenushimnimtum de not arc bueBUEBUF RM cynodontium polycarpon hedw schimp porter 1935 cimphyllumcirriphyllum cirrosum schwaegrSchwaschwaegeregr in schultes grout NY cynodontium schistischiesti web & mohr lindalindb porter 1935 eurhynchium origanumoreganumoreganum sull jaegcaeg spence 1985 dichodontium olympicumolympwumolymp icumwummum ren & card RM eurhynchium pulchellumpulchellum hedw jennjerm RM dichodontium pellucidpellucidumum he1 1 1waww schimp RM homalothecium aeneum mitt lawt RM US dicranellaDicranella schrebeschrebenanaschreberianaschreschfebenanatianariana hedw hilf ex crum & ander- homalothecium nevanevadensedense fesqlesq ren & card US son RM homalothecium pinnatifidpinnatifidumpmnatifidumum sull & lesqfesq lawt buebunBUEBUF dicranellaDicranella subulata hedw schimp RM RM dicranoweisia cirratacarrata hedw lindalindb ex milde RM tomentypnum miens hedw loeske COLO RM dicranoweisia crispula hedw lindalindb ex milde COLO RM BRYACEAE dicranum bonjeanbonjeaniibonjeanubonjeanuii de not in lisa porter 1937 dicranum muehlenbeckmuehlenbeckiimuehlenbeckiaii bruch & schimpsebimp inm BSG COLO bryum algovicum sendtnsendan ex C muell buebuhBUFBUE RM US RM bryum alpinumalpmum hudsbuds ex with porter 1935 dicranum polysetumpolysetum sw wynne 1943 ireland 1982 bryum amblyodon C muell spence 1985 dicranum rhabdocarpum sull RM bryum arcticumarcticum R br bruch & schimp in BSG lawton dicranum scopallumscopanumscopallumpanum hedw RM 1971 dicranum spadiceum zett COLO RM bryum argenteum hedw RM dicranum tauricumtaurtauncumicum sapehinsapehmsapshin RM bryum caespiticium hedw RM US oncophorus birensvirens hedw brid RM bryum camillarecapcapillareillare hedw RM US oncophorus wahlenbergii brid COLO RM bryum cyclophyllum schwaegrSchwaschwaegeregr bruch & schimp in BSGBSC leucobryumParaparaleucobryum enereeenerve thed in hartmharam loeske COLO RM WTU RM bryum dichotomum hedw spence 1985 bryum hsaelisaekisae de not var cuspidatum bruch & schimp in dltrichaceaeditrichaceae BSG marg RM bryum pallescenspallescentpallescens schleich ex schwaegrSchwaschwaegeregr RM Ceraceratodontodon purpurpurpureuspurpureouseus hedw brid bueBUEBUF RM bryum pseudotriquetrumpseudotnquetrumpseudo triquetrum hedw gaertn meyer & scherb distichumDistichum capillaceum hedw bruch & schimp inm BSG RM BUF RM bryum turbiturbinatumnatum hedw turn NY RM distichumDistichum inclinatum hedw bruch & schimp inm BSG bryum uliginosum brid bruch & schimp in BSG spence NY RM 1985 ditrichumflexicauleditnchum flexicaule schwaegrSchwaschwaegeregr hamp RM bryum weigweigellweigelnweigeliieliieilieill spreng in biehler RM Saelania glaucescentglaucescens hedw broth in bomanss & broth leptobryumLeptobryum pynpytifonnepynformeforme hedw wils RM WTU dohliapohlia andalusicaandalusica hoehn broth shaw 1981 dohliapohlia bobolanboianlandendetideriderl sull broth var senata shaw CSU RM encalyptaceaeencalypraceae dohliapohlia camptotrachela ren & card broth shaw 1981 dohliapohlia cruda hedw lindblmdhlindbe BUF RM encalypta alpina sm porter 1937 dohliapohlia drummondrummonddrummondndrummondiidrummondsdnii C muell andr WTU encalypta ciliata hedw porter 1937 horton 1983 dohliapohlia elongataelongateelongata hedw porter 1935 encalypta procelaprocera bruch porter 1937 dohliapohlia lescunanalescuriana sull croutgloutgrout porter 1935 encalypta rhabdocarpa schwaegrSchwaschwaegeregr RM dohliapohlia longicollalongicolla hedw lindalindb RM encalypta vulgaris hedw RM dohliapohlia ludwigludwignii schwaegrSchwaschwaegeregr ex schwaegrSchwaschwaegeregr broth RM dohliapohlia hutansnutans hedw lindalindb buebukBUFBUE CSU RM flssidentaceaefissidentaceae dohliapohlia obtusifoliaobtusifiblia brid L koch COLO RM dohliapohlia proliproligeragera kindbkmdbkinda ex brid lindblinda ex arnell BUF fissidens adianthoides hedw WTU dohliapohlia tundraeaundrae shaw BUF RM fissidens bryoidesbry oides hedw RM WTU dohliapohlia wahlenbergii web & mohr andrews RM fissidens grandtgrandigrandifronsgrandifironsfrons brid bueBUEBUF RM WTU roelliajoellia roellnroelliikoelln broth in roell andrews ex crum RM fissidens obtusifoliusobtusifolious wils var margimarginatusnatus flow NY WTU fissidens osmundioides hedw WTU 200 GREAT BASIN naturalist volume 56

fontinalaceae var resupinatum tayl schimp in spruce porter 1935 dichelymafalcatum hedw myr COLO RM hypnum lindberg ii mitt RM dichelyma uncinatum mitt WTU hypnum pallescentpallescenspalles cens hedw P beaucbeauv RM fontinalis antipyretica hedw var antipyretica RM WTU hypnum revolutumrevolutum mitt lindalindb RM varvan giggigantesgiganteaantea sull sull RM hypnum rauchenvauvauchenvauchericheri lesqfesq LAF var oregonensis ren & card RM WTU isopterygiopsis pulpulchellachella hedw iwatsiwais RM fontinalis hypnoideshypnoides hartmharam var hypnoideshypnoides RM platydietyajungennannioidesplatydictyajungermanmoides brid crum RM var dunaeiduriaeidunaev schimp husn RM ptiliumptilinum crista castrensis hedw de not WTU fontinalis neomexicana sulsuisullsulisuii & lesqfesq RM pylaisiella polyantha hedw grout porter 1935

funariaceae leskeaceae eunana flavicantflavicansflavicans michxmicha porter 1935 funariaflavicansfunana Pseudopseudoleskealeskea incurincurvatiincurvatavata hedw loeske CSU RM hygrornetncahygromettica funaria hedw RM Pseudopseudoleskealeskea patens lindblinda kindbkinda RM physcomithumphyscomitnum hookenbooken hampe TENN fseudoleskeaPseudopseudoleskealeskea radiradicoseradicosacosa mitt macoun & kindbkinda RM var compactscompactacomp acta best lawton 1971 grimmiaceae var mallidapallidapalltda best crum steere & anderson lawton 1971 coscinodonCoscino don calyptracalyptratustus hook drumm C jens ex in Pseudopseudoleskealeskea stenophyllastenophylla ren & spence kindbkinda RM card in roell 1985 grimmia affiasaffimsaffini8 hoppe & hornschhoensch ex hornschhoensch RM pseudoleskeella tectoriumtectorumtectorum funck ex brid kindbkinda in broth grimmia anodon bruch & schimp BSG NY RM in in NY RM WTU grimmia anomala hampe ex schimp lawton 1971 grimmia dondonnianadonmanadonciananiana sm wynne 1943 meesiaceae grimmia elatiorelation bruch ex bals & de not RM grimmia laevigatalaevigata brid brid polporterpoi ter 1935 amblyodon dealdealbatusbatus hedw bruch & schimp BSG grimmia montana bruch & schimp BSG RM in in WTU gemmiagnmmiagrimmia avalisovalis hedw lindb RM linda meesiameecia uliginoseuliginosa hedw RM grimmia plagiopodia hedw RM grimmia pulpulvinatepulvinatavinata hedw sm RM MNIACEAE grimmia telemmatenenimatenemmatenehimanimahlma ren & card RM gemmiagnmmiagfimmia torquatetorquatatorquata hornschhoensch grev CSU in cinclidium stygium sw schrad BUFBUE NYS grimmia hichohichophyllatnchophyllaphyliaphylla grev 1971 spence 1985 in bue lawton mnium ambiguum H muell RM jaffueliobryum wrighwnghtiiwrightiiwrightiiitiitil sull in gray wynne 1943 in ther mnium arizonicumarizoanzomcumnicum amann CSU RM racomitnumracomitriumrhacomitrium canescentcanescenscanescens hedw brid COLO RM WTU mnium blyttii bruch & schimp BSG RM RacoracomithumracomitnummitHum lanuginosum hedw brid crum and ander- in mnium marginatumnatum with brid ex P beaucbeauv RM son 1981 margi mnium spinuspinulosumspmulosumlosum bruch & schimp BSG porter 1935 rhacomitrium sudeticumsudetic & in racomitnumracomitrium um funck bruch schimp in BSG spence 1985 COLO WTU mnium thomsonii schimp RM schistidium agassiziiagassiznagassianii sull & sullsulisuii RM agassiz lesqfesq in sulsui plagiommumplagiomnium ciliareciliary C muell T kop RM schistidium apocarpumum hedw bruch & schimp in BSG apocarp in plagiomniumplagiommum cuspidatum hedw T kop RM NY RM plagiomniumplagiommum drummondrummond dnii bruch & schimp T kop RM schistidium occidentale churchill 1971 lawt lawton plagiomnium ellipticelhpticumellipticumum brid T kop RM RM plagiomniumplagiommum medium bruch & schimp in B S G T schistidium nvularere brid podppoap var nvularerivularere RM rivularedula rivula kop bueBUFBUERMRM var latifolium zett crum & BUF latifolium anderson plagiomniumplagiommum rostratum schrad T kop WTU schistidium tenenimtenerum zett nyhbyh RM rhizomnium magnimagnifoliummagnifohumfolium honkhorik T kop RM scoulenascouleriaScoulenauleria aquaticaaquaticalaquatica hook in drumm RM in rhizomnium pseudopunctatum bruch & schimp T kop RM hedwigiaceae rhizomnium punctatum hedw T kop porter 1935 spence 1985 hedwigishedwigiaHedw igia ciliata hedw P beaucbeauv BUF RM helodiaceae neckeraceae heiHelhelodiumhelodtumodiumodlum blandowblandoioublandowiiblandoblancobiancoiouidutonii web & mohr earnstwarnst BUF RM neckersneckera jennatapennata hedw RM hylocomiaceae orthotrichaceae hylocomium splendentsplendens hedw schimp in BSG RM amphibiumamphidium lapponicumlappomcumlapponicumnicom hedw schimp CSU pleurozium schreschrebenberlberibeyl budbrid mitt WTU orthotrichum affine brid porter 1935 rhytidiadelphusrhiftidiadelphus thquetrustnquetms hedw earnstwarnst RM orthotnchumorthotrichum alpestrealpestre hornschhoensch in BSG RM orthotrichum anoanomalummalum hedw NY HYPNACEAEHYPNACEAL orthotrichumorthotnchum cupulatumcupulatum brid NY RM orthotrichum hailu sull & lesqfesq in sull BUF RM hypnum cupressiforme hedw var cupressiforme BUF orthotrichumorthotnchum holzmgenholzholzingeriingeri ren & card in holz RM NY RM orthotrichumorthotnchum laevigatum zett COLO RM 199611996 MOSSES OF WYOMING 201 orthotnchumorthotrichum obtusifolium brid MO Stegonia latifolialatifolia schwaegrscbwaegrSchwaschwaegeregr in schultes vent ex broth orthotrichumorthotnchum pellucidpellucidumum lindblinda bunbukbupBUE NY RM lawton 1971 orthotnchumorthotrichum praemorsum vent in roell porter 1935 vitt tortellafragilistortella frafragilisfragulisgilis hook & wils in drumm limarlimpr buebukBUEBUF 1973 LAFLAE NY RM WTU orthotnchumorthotrichum pylaisiipylaisn brid vitt 1973 tortella humilis hedw jenn spence 1985 orthotnchumorthotfichum rivulanvularerivularere turn lawton 1971 tortella tortelloidestortellotdestortelloides S greene robins bueBVFBUF WTU orthotnchumorthotrichum rupestrerupestre schleich ex schwaegrSchwaschwaegeregr RM toitortellaTottelladeliadeila tortutortuosatortuoseosa hedw limarlimpr bukBUFBUE RM tortula canicaninerviscanmerviscannervismerdismervis mitt broth BUFBUE NY RM plagiotheciaceae tortula ineinermisinermiaamsnms brid mont porter 1935 tortula mucronifoliamucromfolia Schwaschwaegrschwaegeregr bukBUFBUE NY RM plagiothecium denticulatum hedw schimp in BSG tortula norvegicanorve gica web wahlenbwahlene ex lindblinda bueBUFBUE RM WTU tortula papillosissima copp broth bueBUFBUE RM plagiothecium piliferumpihferumpiliferum sw ex hartmharam schimp in BSG tortula ruralis hedw gaertn meyer & scherb BUF spence 1985 RM tnchostomopsistrichostomopsis australasianaustralasiae grev & hook robins polytrichaceae BING BUF

Atratnchumatrichumichum selwynselwynnii aust RM US pterygynandraceaeptfrygynandraceae attiAtiqatnchumatiqchumchum undulatum hedw P beaucbeauv spence 1985 polytnchastrumpolytrichastrum alpinum hedw G L sm var alpinum heterocladium dimordimorphumphum andbndbrid schimp in BSG BUF COLO RM myurellajulacea schwaegrschwaegerSchwaegr schimp in BSG RM polytnchumpolytiqchum commune hedw RM polytnchumformosumpolytrichumformosum hedw ireland 1982 rhytidiaceae polytnchumpolytrichumjuniperinumjumpennum hedw RM polytrichum longisetumlongisetum budbrid RM rhytidium rugosum hedw kindakindb WTU polytrichum lyalinlyallnlyallii mitt kindbkmdbkinda bukBUFBUE RM polytrichum piliferumpiliferum hedw RM seligeriaceae polytrichum sexangulare brid RM polytrichum strictstrictumstriatumstnctumsanctumum brid RM alindiablindia abutaacuta hedw bruch & schimp in BSG COLO CSU RM pottiaceae seiseligenaseligeriaSelseiiseilSeligeriagerlaigena campylopoda kindbkinda in macoun & kindbkinda ALTA BUFBUE RM barbula convoluta hedw var convoluta RM barbula unguiculata hedw bueBUEBUF LAFILAF sphagnaceae bryoerythrophyllum recurvirostrarecurvirostre hedw chen BUF RM WTU sphagnum angustifolium C jens ex russ C jens in toll COLO RM desmatodonDesmatodon cerneuscernuus huebbueb bruch & schimp in bsrbarBSG RM WTU sphagnum annulatum H lindblinda ex earnstwarnst BING RM

I1 sphagnum contorcontortumtum schultz BING desmatodonDesmatodon guepinguepiniiguepimiii bruch & schimp in BSG bukBUFBUE RRMM sphagnumfimbriatumsphagnum desmatodonDesmatodon heianhelmliheimn hedw mitt bukBUFBUE RM fimbriatum wils in wils & hook f COLO RM Desmadesmatodontodon latifoliuslatifolius hedw brid TENN RM WTU sphagnumfuseumsphagnum schimp RM Desmadesmatodontodon leucostomaleucostoma abrR br berggr WTU luscumfuscum Klingklmggrgr rbr sphagnum platyphyllum desmatodonDesmatodon obtusifoliusobtusifolious schwaegrSchwaschwaegeregr schimp BUF RM lindalindb ex braithwBraithw sull ex earnstwarnst RM desmatodonDesmatodon plinthobiusplmthobius sull & lesqfesq in sull medina 1994 sphagnum russowiirussowii earnstwarnst RM sphagnum rosum crome RM Desmadesmatodontodon porten james in aust porter 1937 squarrosumsquar sphagnum desmatodonDesmatodon syssystyliussystyloustylius schimp BUF RM subsecundum nees in sturm BING sphagnum teres schimp aongstr didymodonDidymodon asperiaspenaspenfoliusasperifoliusasperifoliousfolius mitt crum steere & anderson in hartmharam BING COLO RM sphagnum warnstorfii russ RM COLO didymodonfallaxdidymodonDidymodon fallaxfallad Ifhedwedw zand varvan RM varfallaxfallaxfallad splachnaceae var reflexus brid zand crumgrumcrum and anderson 1981 didymodonDidymodon ngirigingidulusrigidulusrigidulousdulus hedw var ngidulusrigidulusrigidulousdulus BUF ngirigi splachnum sphaericumsphaencum hedw YELLO var gracilis schleich ex lookhookI & grev zand BUF taylonatayloriathyloriananThynayloriaioria acuminateacuminataacuminata hornschhoensch crum and anderson 1981 NY RM taylonataylotqatayrona ligulaligulataligulateta dicks lindblinda COLO RM var icmadophilusienwdophilus schimp ex C muell zand BUF taylonatayloriathyloriatayThyloria serrata hedw bruch & schimp in BSG NY didymodonDidymodon vinealisneails andbndbrid zander vaivar vinealis bukBUFBUE RM var flaccidflaccidusus bruch & schimp in schimp zand BUF tetraphidaceae var sch spreng lundus Hornhornschhoensch in zand as didy tetraphisTetraphis pelpellucidalucida hedw RM modon fariusinformstritrl wynne 1943 var nicholsonicholsoniimcholsominicholsoniiiniinil culm zand RM thuidiaceae var rubiginosus mitt zand BUF sm gymnostomum aeruginosum bukBUFBUE RM WTU abietinellaAbie tinella abiablabietinaabietmaetina hedw fleisch bueBUFBUE RM hymenostylium recurvirostrarecurvirostre hedw dix RM WTU Paludella squarrosesquarrosasquarrosa hedw brid lawton 1971 tlmmiaceaetimmiaceae pterygoneurum kvatumovatum hedw dix bunBUFBUE NY RM pterygoneurum subsessile brid jur bukBUFBUE NY RM timmiadimmia austriacoaustriacaaustriaca hedw bueBUEBUF RM 202 GREAT BASIN naturalist volume 56 timmiatirnrmadimmia megapolitana hedw var megapolitanamegapohtana NY RM brotherellaBrotherella recurvantrecurrecurvansvans michamichx fleisch var barancabavancabavaricabavarica hessidesslhessl endbrid BUF RM lincoln gulch albany county aven nelson 2628 the material is scanty and prof problematic TAXA holzinger who identified it expressed a doubt as to the correctness of the determination brachythecium campestre C muell porter 1935 the specimen with holzinger s schimp inm BSG old faithful yellowstone opinion is at RM national park smiley according to porter bryum canaricanarienseense brid porter 1935 this 1934 1935 porter s citation is apparently not is a species of the west coast and not likely to based on a specimen but on dwight smiley s occur in wyoming the specimen cited by checklist since this is primarily a taxon of the porter nelson 7814 curatedaurated at RM and US eastern united states it should probably be seems to be bryum caespiticium hedw excluded from the state flora specimen not encalypta streptocarpa hedw porter 1930 seen 1935 excluded from north america ander- brachythecium oxycladon brid jaegcaeg this son et al 1990 taxon typical of the eastern united states is cymnostomumgymnostomum calcareumcalcacalcaneumreum nees & hornschhoenschHornsch based by porter 1934 1935 on a citation by in nees et al porter 1937 washakie co smiley no corresponding specimens were seen the specimen from WTU of porter sept 9 funatiaflavicansfunanaeunana flaviflavicantflavicansglavocans michamichx no specimens were 1935 no 2094 on limestone boulders in a seen of this taxon reported by porter 1935 As shady canyonscanyon from the ten sleep canyon in it appears to be a species of the eastern region the big horn mts washakie co has been of the united states crum and anderson determined to be gymnostomum aeruginosum 1981 it is of doubtful occurrence in wyoming by R zander the specimen in section shows a mnium hornum hedw porter 1935 this is ventral costal epidermis two stereidsteroid bands a a taxon of the eastern montane region of north central strand in the stem the capsules were america and the piedmont crum and ander- young and so rather ovoid son 1981 and not likely to occur inm wyoming homalothecium lutescentlutescenslutes cens hedw robins no specimen seen based on a citation by porter 1937 and proba- platydictya confervoidconfervoideses brid crum cited bly the specimen yellowstone national park by porter 1935 as a doubtful determination it nelson no 6041 RM appears to me to be cannot be located in the herbaria consulted homalothecium aeneum instead racomitilumfasciculareracomitnumfasciculare hedw brid porter hypnum callichroumcallichroum funck ex brid evan- 1935 did not see a specimen of this species ston uinta county aven nelson 4128 in part reported by nelson 1900 spence s citation the identity of this plant is doubtful porter for teton county 1985 refers to porter s 1935 uinta co porter 1937 the specimen doubtful citation no specimens were seen by nelson 4128 appears to be hypnum lindbergii the present author Macromacrocomacoma sullivantiisullivantii C muell grout BUF this record is due to a labeling error EXCLUDED TAXA vitt 1981 D vitt in litt in the orthotri chaceae boreali americanaeAmericanae exsiccatae fas- brachythecium calcareumcalcacalcaneumreum kindakindb A specimen ciculus iliIII111 nos 21 30 the label issued with ofofsmileysmiley s ofofbrachytheciumbrachytheciumflexicauleflexicaule now B this species name number 30 should be num- calcacalcareumcalcaneumreum from yellowstone national park and ber 27 orthotrichum rupestrerup estre the macro cited by porter 1935 was determined as a coma specimen originated in north carolina depauperate specimen of B frigifngidumfrigidumdum the orthotrichum from yellowstone national brachythecium glareosumglareosum br B &as&sS lake park J A christie in littliftiltz yellowstone national park smiley porter meesiameecia triquetra richt aongstr reported 1934 1935 specimens of smiley and other by cooper and andrus 1994 is oncophorus collectors at YELLO were variously brachyabrachy wahlenbergwahlenbergiiwahlenbergiaii thecium salebrosum B leibergleibergnleibergiiii B albicans orthotrichum speciosumspeciosum nees in sturm and B fiigidumfngidum porter 1935 specimens at YELLO and RM Breutelia mohnanamohtianajohnanamohTiana C muell broth car- were determinable as 0 laevigatum bon co porter 1937 excluded from north orthotrichum pallenspollens bruch ex brid var ameiamelamericaicaiealea anderson et al 1990 pardumparvum vent yellowstone national park 199611996 MOSSES OF WYOMING 203

flowers 1973 excluded from north america and access to field collections and jennifer anderson et al 1990 wipple of yellowstone is acknowledged taxo- physcomitrium pyriforme hedw hampe nomic assistance was provided by richard cited by porter 1937 for crook co is proba- andrus terry mclntoshmcintoshmeintosh howard crum john bly a specimen collected by marion ownbey spence ronald pursell norton miller dale from that county no 556a TENN and deter- vitt and richard zander the treatment of mined by porter as P turbiturbinatumnatum but which sphagnum was considerably improved by infor- upon examination is P hookerihookershookeri mation provided by dr richard andrus valu- plagiomnium affine bland ex funck T kop able specimens were sent to me by janice PE porter 1935 excluded from north america mckee nancy kastning culp william buck anderson et al 1990 holmes rolston william reese and joseph plagiothecium cavifoliumcavifolium brid iwatsiwais elliott this project was sponsored by the sav- porter 1935 based this name on a specimen of age fund of the buffalo museum of science elias nelson 5242 which is isopterygiopsis and is dedicated to the memory of the late pulpulchellachella elva lawton sphagnum capillicapillifoliumfolium earhehrh hedw porter 1935 porter s specimens 1198 and literature CITED 1199 collected in 1932 identified as S capilli had ANDERSON L E 1990 A checklist of sphagnum in north folium been redetermined by R andrus as america north of mexico bryologist 93 500 501 S russowiirussowii andrus in littlift ANDERSON L E H A CRUM AND W R BUCK 1990 list sphagnum majus russ C jens porter of the mosses of north america north of mexico 1935 crum 1984 taxa collected from wyom- bryologist 93 448 499 BRASSARD GUY R 1983 checklist of the mosses of the ing and identified as S majus have all been S island of newfoundland canada bryologist 86 546354 63 annulatum according to andrus in liftfitthittbitt who CHRISTY J A J H LYFORD AND D H WAGNER 1982 states that the nearest sites would be in british checklist of oregon mosses bryologist 85 22 36 columbia central alberta and minnesota CHURCHILL S P 1979 mosses of the great plains illiliIII111 additions nebraska and the sphagnum palustre L RM to black hills of south COLO this dakota and wyoming bryologist 82 72 75 species has been found only along the west 1982 mosses of the great plains VIII additions coast by andrus in littlift bryologist 85 218 221 sphagnum fecurecutrecurecurimmrecurrecurvumdumrimmfimmvum P beaucbeauv COLO RM COOPER DAVID J AND RICHARD E ANDRUS 1994 pat- material of this species from the interior of the terns of vegetation and water chemistry in peatpeatlandslands of the west central wind river range wyoming united states is referable to S angustifolium USA canadian journal of botany 72 1586 1597 according to andrus in littlift sphagnum recur CRUM H A 1984 sphagnopsidaspbagnopsida sphagnaceae north vum is an eastern coastal plain species american flora seriesserles II11 part 11 new york botanical garden tortula princeps not reports by porter 180 appp de CRUM H A AND L ANDERSON mosses from E 1981 of east- carbon and crook counties 1935 were ern north america columbia university press NY based on nelson 2818 and 5034 at RM and a EVERSMAN S AND A J SHARP 1980 first checklist of specimen RM by hermann no 17844 which montana mosses proceedings of the montana acad- were redetermined by R zander as tortula emy of science 39 12 24 FFLOWERSWERS S A HOLMGREN EDITOR 1973 mosses utah ruralis and the west brigham young university press wtissiaweissia controversacontcontroversyroversa hedw porter 1935 all provo UT citations for this species appear to be based on HORTON D G 1983 A revision of the encalyptaceae II11 specimens of dwight C smiley deposited at journal of the hattonhattori botanical laboratory 54 353 532 YELLO 3 seen all specimens were dicianodicrano IRELANDLAND R R 1982 moss flora of the maritime provinces reisiaweisia crispula national museums of canada ottawa JOHNSEN ARDITH B undated keys to the mosses of ari- acknowledgments zona research paper 14 museum of northern arizona flagstaff ketchledge E H 1980 revised checklist of the author gratefully mosses the acknowledges the help of new york state in R S mitchell editor flora of of the late mason hale for his early support of new york state new york state museum bulletin this project holmes rolston and the late fred 440 albany NY of LAVIN M 1981 new records for the moss flora of nevada hermann the assistance ronald hartman bryologist of 84 93 94 the rocky mountain herbarium who pro- LAWTON E 1958 mosses of nevada bryologist 61 vided me with a copy of porter s dissertation 314 334 204 GREAT BASIN naturalist volume 56

1971 moss flora of the pacific northwest journal sijansijawHAW A J 1981 A taxonomic revision of the propagulifer of the hattonhattori botanical laboratory nichinanNichinan japan ous species dohliapohliaofofpohha musci inm north america jour- mccleary J A AND V V GREEN 1971 A checklist of nal of the hattonhattori botanical laboratory 50 1 8811 idaho mosses bryologistbiyologist7474 175 180 SPENCEPENCE J R 1985 checklist of the mosses of grand teton MEDINA ALVIN L 1994 lichens and bryophytes of the national park and teton county wyoming great rochelle hills campbell county wyoming evansiaevangia basin naturalist 45 124 126 1 121 130 1988 checklist of the mosses of the intermoun- NELSON AVEN 1900 the cryptogamscryptocryptogramsgams of wyoming 10th tain west USA great basin naturalist 48 394 401 annual leportreport of the wyoming experiment station VITTITT D H 1973 A revision of the genus orthotnchumorthotrichum laramie 38 appp bryophytorum bibliotheca J cramer verlag vaduz PORTER C L 1930 fruiting plants of encalypta confortaconcontortatorta EDITOR 1981 orthotnchaceaeorthotrichaceae boreali ameriamerlamen bryologist 34 93 canabcanae exsiccatae fasciculus III111 nos 21 30 univer- 1934 the moss genus brachythecium in wyoming sity of albertaofalberta edmonton canada university of wyoming publications in science 1991 rediscovery of orthotnchumorthotrichum lolholboiholzingerholzingeriholzingenholzingerizingen its botany 19 235 241 morphology and habitat in western north america 1935 bryophytes of wyoming part II11 hepaticae bryologistbryologist9494 77 79 concluded and musci bryologist 38 101 114 WEBEREBER W 1973 guide to the mosses of colorado insti- 1937 the bryophytes of wyoming unpublished tute of arctic and alpine research occasional paper doctoral dissertation university of washington 6 university of colorado denver seattle WYNNEYNNE F E 1943 range extensions of mosses in western SAYRESAYRL G 1971 Cryptogamcryptogamaeae exsiccatae an annotated bib- north america bryologist 46 149 155 liographylio of published exsiccatae of algae lichenes hepaticae and musci memoirs of the new york received 21 december 1995 botanical garden 192 175 176 214 215 accepted 2277 march 1996 great basin naturalist 563 C 1996 appp 205 210

VARIATION IN bitterbrushBITTERBRUSH PURSHIA tridentatatridentateDENTATATRI PURSH CRUDE PROTEIN IN southwestern MONTANA

carl L WamboltWarnboltl1 W wyatt fraaslfraas1fraasa and michael R Frisinafnsina2frisina22

ABSTRACT the objective of this study was to determine if crude protein varies significantly during late summer and midwinter among stands of bitterbrushbitterbrush purshia tntridentata pursh in southwestern montana A secondary objective was to determine if leaves when present contribute significant additional protein in the region nine sites with different environmental conditions and within a radius of 14514.514 5 km were studied bitterbrushBitterbrush leaves and leaders collected in august 1990 and 1991 and february 1991 were used for crude protein and leaf to leader ratio determinations crude protein differed P 00010.0010 ooi001 among sites for both leaves and leaders on individual collection dates crude protein in leaves was nearly twice the level found in leaders because few leaves were present in february they increased crude protein in total foliage by only 030.30 3 over twigs alone february crude protein levels averaged 686.86 8 for total foliage which is below the estimated requirement for wintering deer

key words purshia tntritrldentata bitterbrushbitterbrush crude protein winter range big game nutrition montana

protein is one of the most important nutri- METHODS ents for wintering ungulaungulatesungulatedtes dietz 1972 study sites welch et al 1983 estimated that winter crude protein levels of purshia dentatatritridentatatridentate pursh bit nine study sites were chosen primarily to terterbrushterbruschbrush are not high enough to meet ungu- represent bitterbitterbrushbrush stands from a range of late requirements but postulated that protein environmental conditions table 1 this in- content might vary with populations of bitter cluded burned sites and bitterbrushbitterbrush sites protected from browsing study brush differences in bitterbrushbitterbrush protein con- all sites were located within a radius of 14514.5 km near tent between sites have been noted giunta et butte and anaconda in southwestern montana long- al 1978 although not between local habitat term climatic records were available for the types morton 1976 blausen slausen and ward 1986 general study area from the anaconda weather found no difference in crude protein among 3 station at 1700 m elevation annual precipita- colorado accessions in a common garden but tion at anaconda averages 340 mm with 47 welch et al 1983 found differences in a com- received between april and july NOAA 1991 mon garden test with plants from a wider geo- vegetation types at all but 3 sites burn graphical area no differences in nutrient con- unburn and high rye were seraiseral stages of the tent have been found at varying browse levels bitterbrushbitterbrush bluebunchbluebunch wheatgrasswheatgrass agropyron of bitterbrushbitterbrush plants dietz et al 1962 shep- spicatumspicatum pursh habitat type mueggler and herd 1971 crude protein levels were higher stewart 1980 the dominant shrub was bitter when winter leaves were present dietz et al brush but understory vegetation was regressed al 1962 but winter leaf presence varies between fraas et 1992 on the other 6 sites from the described potential climax youtie populations of bitterbitterbrushbrush welch et al 1983 composition et al 1988 our objective was to determine if crude pro- the butte site at maude S canyon near tein varies significantly during late summer and butte montana was selected because it re- midwinter among stands of bitterbitterbrushbrush south- in ceives no ungulate browsing the plant com- western montana secondarily we wished to munity consisted ofbitterbrushbitterbrush centaurea mac determine if bitterbitterbrushbrush leaves in our region alosaulosa lam spotted knapweed ribes cereuecereum contribute significant additional crude protein dougl squaw currant and rosa woodsiiwoodsie quantities when present lindl woods rose

I1 department of animal and range sciences montana state university bozeman MT 59717 montana fish wildlife and parks butte MT 59701

205 206 GREAT BASIN naturalist volume 56

tableTABLLTABLF 1 topographic characteristics of the 9 study sites east edge of the MHWMA big game winter data from the last 4 sites were obtained from guenther range plant community consisted ofbitter 1989 the brush and spotted knapweed the willow creek site elevation slope aspect site was near the top of a grassy ridge 150 m m degrees above willow creek this site supported a rel- butte 1730 26 234 atively large amount of elymus cinereuscicinereousnereus scribn cattle exclosure 1830 16 188 & merr basin wild rye along with other cattle deer 1820 10 190 perennial grasses and bitterbitterbrushbrush area burn 2010 21 220 this unburn 2010 24 180 was used as winter range by mule deer elk Powerline 1640 16 85 and moose the railroad gulch site was also willow creek 1780 31 110 on the deer and elk winter range this site gulch railroad 1650 32 115 occupied a midslopemidslope position 30 m above an high rye 1940 38 120 intermittent stream where the plant community consisted of bitterbrushbitterbrush and spotted knapweed the high rye site was 1500 m higher in at dry cottonwood creek in the Deerdeerlodgelodge elevation than the other MHWMA sites and district of the Deerdeerlodgelodge national forest a appeared to receive the greatest snowsnowpackpack livestock exclosure with deer only use was the plant community on the high rye site was studied and known as the cattle exclosure site typical of the bitterbrushbitterbrush rough fescue fes- near the exclosure a bitterbitterbrushbrush stand was tuca scabrella torrey ex hook habitat type studied and known as the cattle deer site mueggler and stewart 1980 with those species because it sustained both cattle and mule deer currently dominant guenther 1989 found the browsing these 2 sites have a scattered over- least amount of big game use at this location story of pseudotsuga menziemenziesiimenziesnmenziesiasiislisn birbmirb franco among the 4 MHWMA sites the MHWMA douglas fir A high number of native peren- study sites received insignificant levels of live- nial forbs occurred in the understory on these stock grazing sites sampling and analysis two sites were selected to gauge the impacts of burning bitterbitterbrushbrush in southwestern mon- leaves and leaders current year stem tana the 2 sites burn unburn were situated growth minus leaves were collected at each on either side of the burn line on the south study site from 10 randomly selected plants for flank of steep mountain 8 km northwest of crude protein analysis on each of the sampling butte in the butte district of the Deerdeerlodgelodge dates the same plants were sampled to deter- national forest the plant community on these mine leaf to leader ratios material was col- 2 sites was a bitterbitterbrushbrush mountain big sage- lected the ist week of august prior to or at brush artemisia ttltntrltridentata nutt sspasp vastyanavasevaseyanayana seed set in 1990 and 1991 this was estimated redbrydb beetle bluebunchbluebunch wheatwheatgrassgrass associa-assoriaassocia to be the period of minimum soluble carbohy- tion intermediate to the big sagebrush blue drate content for bitterbitterbrushbrush plants menke bunch wheatwheatgrassgrass and bitterbrushbitterbrush bluebunchbluebunch and trlica 1981 material was also collected on wheatgrasswheat grass habitat types of mueggler and 12 february 1991 when mule deer were con- stewart 1980 the prescribed burn was con- centcentratedrated on these sites plant material was ducted 3 november 1981 after a year s rest from oven dried at 60c60goc C for 48 h and weight of dry livestock grazing to increase fuel loads live- matter determined leaves were separated stock use resumed 15 september 1982 when from leaders and weighed separately to deter- sampled for protein content bitterbrushbitterbrush on the mine leaf to leader ratios on a percent dry burned site was significantly lower in canopy matter basis leaves and leaders were then cover FP ooi0010.010 01 flower production P 010.10oi 1 ground to approximately I1 mm diameter in a and seed production P 010.1oi0 1 than on the un- grinder janke & kunkel kg type aioalo kjel- burned site fraas et aal 1992 dahl nitrogen analyses were used to arrive at four sites were located on the mount hag- crude protein contents winter crude protein gin wildlife management area MHWMA values were calculated with a weighted aver- owned and managed by montana fish wildlife age of winter leaf and leader protein levels and parks the werlinePopowerlinepowerlme site was on a slope this allowed comparison with other studies 50 m above a perennial stream on the north dietz et al 1962 welch et al 1983 199611996 VARIATION IN BITTERbitterbrushbltterbrushBRUSH CRUDE PROTEIN 207

soil samples were obtained at a depth of 15 protein levels also differed P 00010.001 cm below the surface from a soil pit in each among the 3 collection dates fig 1 when all study plot because soils at most sites con- sites were pooled august leaf protein increased tained a large rock fraction it was necessary to 11 between years P 005oos0.05 and february sample at the relatively shallow depth of 15 cm leaf protein decreased 21 from august levels to standardize sampling the montana state P 00010.001 leader crude protein did not vary university soil test laboratory performed significantly between years but was higher in organic matter determinations and total kjel- august 1991 than during the previous febru- dahl nitrogen analyses on all non MHWMA ary P 0050.05oos samples texture was determined by both the the browsedununbrowsedunbrowned butte site rated highest in hydrometer and bouyucous mechanical analy- crude protein fig 1 for 3 of the 6 measure- sis methods soil ph was determined in I1 part ments although none was significantly higher soil to 2 parts water extractions topographic than the next lower site when the butte site information was also recorded at each site was compared to the aggregated crude protein aspect was determined by taking a compass levels of the other 8 sites it was significantly P bearing from the major slope slope was mea- 0050.05 higher for both leaves and leaders in sured with a clinometer elevation was deter- august 1990 but did not differ from browsed mined from USGS topographic maps the sites in february or august 1991 thus it does informationinf6rmation from MHWMA sites was derived not appear that browsing affects crude protein from guenther 1989 levels protein A one way ANOVA with site as the factor values for the 4 MHWMA sites were was conducted for each sampling date and pro- lower for august 1990 leaves P 0070.07 and tein source combination snedecor and coch- leaders P ooi0010.01 than for other sites and col- lectively ran 1989 this was done with the knowledge rated lowest for 4 of the 6 measure- that protein sources leaves or leaders con- ments these site differences were not expected from mortonsmortens 1976 work but tained very different levels of crude protein were supported by that of giunta et al 1978 and welch al within each sampling date site was also the et 1983 factor in an ANOVA for percent leafinesslealeakinessfiness for bitterbrushBitterbrush crude protein levels on the deer the february 1991 sample the least significant cattle site were 1 higher P 005oos0.05 than difference LSD method FP 0050.05oos protected on the adjacent cattle exclosure site for august by a prior F test FP 0050.05oos was used for com- 1990 leaves fig 1 other protein levels did paring treatment means snedecor and cochran not differ significantly between these 2 sites 1989 although a difference in use might thus seem to affect protein levels on these sites the RESULTS AND discussion browsedununbrowsedunbrowned butte site had higher protein levels than browsed sites in august 1990 P 0050.05oos levels P 1 crude protein differed 00010.0010 00 and no difference in february or august 1991 among sites within each protein source and related to these site and possible popula- collection 1 date combination fig thus we tion alderfer 1977 differences are soil differ- rejected the hypothesis that crude protein val- ences soil samples from shrub interinterspacesspaces ues are equal during august and february table 2 contained 49 more soil nitrogen at among local stands of bitterbrushbitterbrush crude pro- the butte site than at the burn and unburn tein in the leaves when averaged over all sites sites and 78 more than at the cattle exclosure varied with a 13 to 10 decline from august and deer cattle sites bayoumiqayoumi and smith 1990 to february 1991 and subsequent increase 1976 found a positive response of bitterbrushbitterbrush to 15 by august 1991 crude protein in the protein levels to fertilization with nitrogen leaders for these 3 dates was 717.1 656.5gs and although tiedemann 1983 found slightly neg- 727.2 respectively when averaged over all ative to no response to fertilization however sites these crude protein levels generally most desert shrubs accumulate nutrients under agreed with previous reports for bitterbitterbrushbrush their canopiescanopies and the surrounding interinterspacesspaces throughout its range dietz et al 1962 bay have low nutrient content garcia moya and oumi and smith 1976 morton 1976 tiede- mckell 1970 tiedemann and klemmedson mann 1983 welch et al 1983 1973 conditions that we did not sample 208 GREAT BASIN naturalist volume 56 leaves

A a b b aceC bdb d percent A c b crude A e ccdd e ekreko000 d protein b 18 bl 16 14 12 highi gkbyeRY e 10 ffiafirailroadffiroad gulch 8 willjkcreekavvcreek Powepowerildennierilde 6 L Un urn 4 bumRUr n 7 IFqatlleandtlQ d deer 2 BL catlleekclosureowneCOOR E losure 0v buiterbutteratteutte augaug9090 feb 91 aug91aug 91

leaders

T percent crude

protein b bd ID C a e 18 c C d C 16 J c 14 ae d 12 d c d higrpvee 10 V gulch A IPVVirailroadrailnailnall 8 power e 6 unburfunburur 4 B urq CQ U I1 d deer 2 cattiecattle exclosure aug 90 feb91feb 91 aug91aug 91

fig 1 average percent crude protein in bitterbitterbrushbrush leaves and leaders found in august 1990 february 1991 and august 1991 protein values within each protein source and collection date with similar lowercase letters are not significantly different LSD P 005oos0 05 insufficient leaf material was available for statistical analysis in february 1991 199611996 VARIATION IN BITTERbitterbrushbltterbrushBRUSH CRUDE PROTEIN 209

TABLE 2 soil characteristics for study areas including ph organic matter OM total kjeldahl nitrogen N percent sand silt and clay and textural class soil nitrogen was sampled at only 5 sites data from the last 4 sites were obtained from guenther 1989

site ph OM N sand silt clay textural class M N butte 56 26 011 63 24 13 sandy loam cattle exclosure 57 10 006 80 12 8 loamy sand cattle deer 57 10 006 80 12 8 loamy sand burn 63 11 007 67 23 10 sandyloamsandy loam unburn 63 11 007 67 23 10 sandy loam Powerline 58 14 65 15 20 sandyloamsandy loam willow creek 52 36 65 18 17 sandy loam railroad gulch 57 10 72 18 10 sandyloamsandy loam high rye 67 28 69 15 16 sandy loam

protein levels at our study sites were therefore guenther 1989 reported that deer pellets not necessarily related to soil nitrogen levels from the MHWMA sites contained large although most leaves had fallen by febru- amounts of rocky mountain juniper juniperus ary all sites contained plants that had retained scopulorum sarg and oregon grape berberis some leaves at that time bitterbrushBitterbrush phenol- rapensrepens lindl protein values for small winter ogy seems to vary more by season and climate samples of oregon grape and juniper from the than by ecotype shaw and monsen 1983 most willow creek site were 848.4 and 696.9gg re- leaves are deciduous dropping in response to spectspectivelyively these values are below those re- moisture stress in late summer or fall shaw ported by welch et al 1983 and like bitter and monsen 1983 but some small leaves over- brush are also below what they considered to winter on some populations alderfer 1977 be the necessary threshold of 898.9 crude pro- dietz et al 1962 alluded to the high protein tein for wintering deer hamlin and mackie level of leaves in winter but did not quantify 1989 suggested that mule deer have more those levels welch et al 1983 reported that need for high quality forage in the fall while winter leafinesslealeakinessfiness presumably weight of leaves building energy reserves than in the winter compared with weight of stems of plants from bitterbrushBitterbrush in southwestern montana may idaho colorado utah and california ranged supply this needed level of nutrients in the fall from 595.9sg to 155iss15.5 while combined leaf and as we observed delayed leaf fall on wind pro leader crude protein ranged from 595.9 to teatedtected bitterbitterbrushbrush plants in late november 1990 797.9 these ranges are similar to values found but we did not sample plants at that time for these montana sites lealeafinessleakinessfiness percent restoration efforts for ungulate winter weight of leaves per weight of stems of 15lsis1.5 to ranges capable of maintaining bitterbrushbitterbrush may 15815.8 and combined crude protein of 616.1glgi to benefit through consideration of our results 767.6tg table 3 because so few leaves were we have found that bitterbrushbitterbrush populations of present in february table 3 crude protein in even a localized ecotype such as we studied total foliage increased by only 030.3 over twigs should not be expected to attain the same lev- alone for all sites els of crude protein over different environmen- although we concluded that leaves contain tal conditions that will vary between sites significantly more crude protein than leaders revegetation of bitterbrushbitterbrush ranges will involve on our study sites leaf scarcity during winter consideration for obtaining the best possible in our region prevents total leaf and leader plant materials our evidence indicates that crude protein from meeting deer requirements plant characteristics other than protein con- welch et al 1983 the february crude pro- tent should likely be of primary concern as tein levels for total foliage averaged 686.8 across protein can be expected to vary by site condi- sites which were below the estimated neces- tions regardless of plant material however it sary threshold of 898.9sg for wintering deer welch appears that consideration should be made of et al 1983 however september through bitterbitterbrushbrush genotypes that maintain a high per- november protein levels might have been centage of leaves into the winter these geno- higher as many plants retained leaves through types may provide a higher level of crude pro- that period tein that is desirable for wintering ungulaungulatesungulatedtes 210 GREAT BASIN naturalist volume 56

TABLE 3 winter crude protein content percent of bit HAMLIN K L AND R J MACKIE 1989 mule deer in the terterbrushterbruschbrush leaves and leaders combined and percent lean- missouri river breaks montana montana depart- ness percent weight of leaves per weight of stems for ment of fish wildlife and parks study sites sampled february 1991 column entries with MENKE J W AND M J TRLICA 1981 carbohydrate similar letters are not significantly different LSD P reserve phenology and growth cycles of nine col- 005 orado range species journal of range management 3426934 269 277 finess site crude protein Lealeanness MORTON M A 1976 forage relationships of mule deer in N the bridger mountains montana nutritional values of mule deer forage plants in the butte 76 136ab13 6 important winter in unpublished master s cattle exclosure 664ed4 65d6 ad3d badgerbndgerbridger mountains montana cattle deer gge 99gge thesis montana state university bozeman MUEGGLER STEWART grassland and burn 6711e6 7 996ed6 W AND W L 1980 habitat types of western montana USDA unburn 76a7 6sas 105bc10 5 shrubland and range Powerline 61dgid6 id 85ed8 5 foiestforest service intermountain forest experiment general technical report INT willow creek 71all7 ply 8 led1 station 66 UT railroad gulch 721 158a15 8aaa ogden NOAA 1991 climatological data montana 84 941 13 high rye 61d6gidP iselse15e1 56 national climatic data center asheville NC SHAW N L AND S B MONSEN 1983 phenology and growth habits of nine antelope bitterbitterbrushbrush desert bitterbitterbrushbrush stansbury clifcliffrosefrose and apache plume literature CITED accessions pages 55 69 in A R tiedemann and K L johnson compilers proceedings of the research and management of Bitterbitterbrushbitterbrusbbrush and cliffroseCliffrose in ALDERFERALDERrER J M 1977 A taxonomic study of bitterbrushbitterbrush in derfer western USDA service gen- purshia tntritrldentata pursh DC in oregon unpub- north america forest lished master s thesis oregon state university cor- eral technical report INT 152 vallis SHEPHERD H R 1971 effects of clipping on key browse colorado colorado game BAYOUMI M A AND A D SMITH 1976 response of big species in southwestern parks 28 game winter range vegetation to fertilization journal fish and division GFP R T denver CO of range management 29 44 48 SLAUSEN W L AND R T WARD 1986 exogeneticecogeneticEcogenetic pat- arid DIETZ D R 1972 nutritive value of shrubs pages terns of four shrub species in semisemiarid communities of colorado southwestern naturalist 289 302 inm C M mckell J P blaisdell and J R northwestern coodingoodin editors wildlandwindlandWildland shrubs their biology and 3131931 319 329 utilization USDA forest service general technical SNEDECOR G W AND W G COCHRAN 1989 statistical methods iowa state press ames report INT 1 ogden UT university TIEDEMANN A R 1983 response of bitterbrushbitterbrush and asso- DIETZ D R R H UDALL AND L E YEAGER 1962 broadcast phospho- chemical composition and digestibility by mule deer ciated plant species to nitrogen pages of selected forage cache la poudre range rus and sulfur fertilization 240 253 in A R species proceed- colorado colorado game and fish department tiedemann and K L johnson compilers technical publication 14 ings of the research and management of Bitterbitterbrushbrush and cliffroseCliffrose western north america USDA FRAAS W W C L WAMBOLT AND M R FRISINA 1992 in for- general 152 prescribed fire effects on a bitterbrushbitterbrush mountain big est service technical report INT sagebrush bluebunchbluebunch wheatwheatgrassgrass community pages TIEDEMANN A R AND J 0 klemmedson 1973 effect of on physical and chemical properties of the 212 216 in W P clary E D mcarthur D bedunah mesquite range 29 and C L wambolt compilers proceedings of the soil journal of management 26 27 B S B MONSEN AND N L SHAW 1983 symposium on ecology and management of riparian WELCH L shrub communities USDA forest service general nutritive value of antelope and desert bitterbitterbrushbrush pages technical report INT 289 ogden UT stansbury cliffrosecliffrose and apache plume 173 185 A R and K L johnson GARCIA MOYA E AND C M MCKELL 1970 contribution in tiedemann compilers and manage- of shrubs to the nitrogen economy of a desert wash proceedings of the research plant community ecology 51 81 88 ment of cliffroseCliffrose in westeinwestern north america USDA general technical report INT 152 GIUNTA B C R STEVENS K R JORCENSENJORGENSEN AND A P forest service YOUTIE B A B GRIFFITH AND M PEEK 1988 succes- PLUMMER 1978 antelope bitterbrushbitterbrush an important J wildlandwindlandwildland shrub utah division of wildlife research sional patterns in bitterbitterbrushbrush habitat types in north central washington journal of range management publication 781278 12 122 GUENTHER G E 1989 ecological relationships of bitter 4112241 126 brush communities on the mount haggin wildlife management area montana department of fish received I1 november 1995 1 may wildlife and parks and montana state university accepted 1996 department of animal and range sciences boze man great basin naturalist 563 0 1996 appp 211 224

DAM FORMING CACTI AND NITROGEN enrichment IN A pironPINON JUNIPER WOODLAND IN northwestern ARIZONA

molly thomas hyselllhysell1 and charles C gnerigner2grier2

ABSTRACT in a pifionpigionpmon juniper woodland in northwestern arizona connected basal cladodescladoduscladodes of a prickly pear cactus opuntia littoralishttoralislittoralis var martmartinianaimana form check dams that cause deposition of N nchrich detritus in interinterspacesspaces otherwise lacking litter seventy eight percent of connected basal cladodescladoduscladodes measured in transects grew at an angle with respect to the slope contour 45 an orientation facilitating deposition of flood borne debris soil total N was significantly greater P ooi00100.0101 and organic C was greater but not significantly above cactus dams compared to below cactus dams soil total N and organic C both above and below cactus dams were significantly greater P 0 0001000010.0001 compared to adjacent interspacesinterspaces soil total N and organic C above cactus dams were equal to areas beneath canopiescano pies tree and shrub combined net n03 0 5 cm depth above cactus dams was significantly greater P 0 0001000010.0001 than below cactus dams at interinterspacesspaces and beneath canopiescanopies net nhanh4NH 0 5 cm soil depth above cactus dams was significantly greater P ooi00100.0101 than below cactus dams and interinterspacesspaces and was greater but not significantly than beneath canopiescanopies at 5 10 cm soil depth differences in net nhanh4 and net n03 between sampling locations were not significant except for the difference in net n03 above and below cactus dams P 005oos0 05 the litter layer above cactus dams had twice as much total N P ooi0010.010 01 as the litter layer beneath canopiescanopies tree and shrub combined differences in net mineralized N were not significant between litter layers over the course of a single rainy season detritus depth behind cactus dams increased up to 23 cm with a mean increase of 434.34 3 cm sys 06250.6250 625 P 0 000100001

key words prickly pear cactus nitrogen enrichment growth habit soil characteristics check dams detritus runoff bulk density total nitrogen organic carbon mineral nitrogen pifionpigionpmon juniper woodlands islands of fertilityoffertility

the growth habit of opuntia littlittoralisoralis var 1981 litter occurs in patches due to the non- martiniana L benson L benson consists of contiguous canopy cover and soil N distribu- connected basal cladodescladoduscladodes growing across wood- tion corresponds to litter and canopy distribu- land slopes roughly along the contour clad tion debano and klopatek 1987 tiedemann odes in contact with the ground sprout adven- 1987 in mixed species stands patches may be titious roots and become anchored sequentially mosaics of different litter components anchored cladodescladoduscladodes function as check dams dur- interspace and canopy area soils usually dif- ing runoff events causing deposition of flood fer in characteristics such as concentrations of borne detritus including surface soil animal nutrients ph bulk density soil water and in feces and litter of pinon pines juniper and numbers and species of resident microorgan- oak isms and micromicroarthropodsarthropods everett and shar- pinon juniper woodlands occupy at least 17 row 1985 klopatek 1987 klopatek and klo- X 106 ha in the western US with widespread patek 1987 although there are exceptions to distribution in colorado new mexico arizona this generalization debano et al 1987 soil eastern california nevada and utah west organic matter and nutrients are concentrated 1988 these woodlands fall between mesic near the soil surface west and klemmedson conditions that support closed forest canopiescanopies 1978 lyons and gifford 1980 debano and and andaridarld conditions in which plants are widely klopatek 1987 and runoff from storms can spaced compared with forests of wetter envi- carry considerable amounts of detritus rich in ronmentsronments pinon juniper woodlands have low organic matter and N fletcher et al 1978 biomass leaf area and primary productivity objectives of this study were 1 to charac- grier et al 1992 woodland structure varies terize the angle of growth relative to slope con- but can generally be described as single trees tour of connected basal cladodesclacladodusdodes of opuntia and shrubs and clumps of trees and shrubs sur- littoralislittoralis var martiniana 2 to compare litter rounded by a network of interinterspacesspaces lanner and soil properties above and below cactus

1 departmentldepartment of forest resources college of natural resources utah state university logan UT 84321 2departmentepaitment of forest science colorado state university fort collins CO 80523

211 212 GREAT BASIN naturalist volume 56

TABLE 1 sites of measurement of angle relative to the slope contour of connected basal cladodescladoduscladodes samples for soil comparisons were taken only in the Huhualapaikualapaialapai mountains see table 2 transect soil pinon pine length parent elevation aspect slope cover location Wm material m 0 0 cerbatcarbat mountainsamountainshMountmountains1ainsa 401 granite 1930 SSSES SSE 10 25 10 30

Huhualapaikualapaialapai mountainshM aunauntamstams 110 granite 1524 N 15 45 30 40

music mountainsmountainshmountains00 302 vesicular 1712 E 30 40 70 basalt

112727 kinkmkiuklu nwofkingNW of kmgminan AZ kitkutlat3527litlat 3527 long11409longiong 11409 t24nr18ws23nw 111212 kmkinkruklu SE ofkingminofkingmln AZ bitbrtlat3508litlat 3508 long11355long 11355 t20nr16wslswt20nr16ws1sw 53 kmkinkiuklu NE ofofkingminkinginanKin ginan AZ lat3541lat 3541 long11349longiong 11349 t27nr36ws36ne

dams and 3 to compare litter and soil proper- summer rain sometimes occurs as intense ties above and below cactus dams with inter thunderthundershowersshowers sellers and hill 1974 spaces and areas beneath canocanopiespies we first observed dam forming cacti in the hualapaikualapaiHualapai mountains rising to over 2438 m METHODS 12 km southeast of kingman arizona in the course of data collection for studies of pinon two distinct physiographic provinces come juniper woodland productivity we subsequently together in northwestern arizona southeast visited 2 nearby ranges the cerbatcarbat mountains west and north of kingman arizona is the over 2133 m at highest point 29 km northwest basin and range province characterized by of kingman and the music mountains over north trending fault block mountain ranges 2011 mlm 53 km northeast of kingman and separated by broad desert valleys the col- found dam forming cacti in these locations to orado plateau lies to the east this area is the characterize the angle of growth of connected interface of 3 deserts as well as a physiographic basal cladodescladoduscladodes of prickly pear cacti our ist ob- interface north of kingman is the great basin jecjectivetive we took angle measurements in july desert west is the mojave desert and south- 1991 on all cacti intercepting straight line tran- west is the sonoran desert the climate of sects in the 3 mountain ranges table 1 start- northwestern arizona is semiarid sellers and ing points of line transects were randomly lo- hill 1974 precipitation is bimodal occurring cated and direction of transects was along ran- mostly in winter and summer months with dom azimuths A total of 233 angle measure- more rainfall during winter than summer ments were recorded sequentially connected

77 450 300 68 16 450

451 4040.40

fig 1 measurement of angle of growth of connected basal cladodescladoduscladodes with respect to the slope contour point of origin indicated by solid cladode 199619961 DAM FORMING CACTI AND NITROGEN enrichment 213 basal cladodescladoduscladodes with series ranging from 040.4 in TABLE 2 characteristics of litter and soil sampling site in a pinon juniper woodland in the Huhualapaikualapaialapai mountains of to 252.5 in in length were measured with an I1 northwestern arizona records 1967 1991 of licensed engineer s adjustable triangle as shown in fifig-9 livestock grazing show year round grazing of cattle and ure 1 a direction of growth parallel to slope horses with year to year variation in season of heaviest use contour was 0 while a direction of growth per- and in number of animalsofanimals USDA BLM 1991 pendipendicularcular to slope contour either upslope or elevation 1524 m 5000 ft downslope was 90 slope 204020 40 aspect north soil and litter sampling soil parent material granite soil texture sandy loam SITE description we restricted litter and soil classificationclassificationsclassification33a barkerville seriesserles soil sampling to I1 of the 3 transect locations loamy mixed mesic shallow the Huhualapaikualapaialapai mountains 12 kinkm southeast of udorthentic haplustolls 21 kingman table 2 fig 2 to minimize con- 11 other soil characteristicscharacteristiesbcharacteristics11 such as different soil types founding factors al horizon 10 cm deep 39 coarse frag site histories and land management practices ph surface soil interspace 65 about 40 of the study site is open interspacesinterspaces ph surface soil under canopy 808.0so combined data unpublished from eighteen 2 nonnoncalcareouscalcareous throughout X 2 m plots using daubenmire s 1968 cover- species and covert 57 age classes and from 12 permanently marked finnsfinusrinusfenuspinus monomonophyllaphylla subspfallax 360 s F 575.7 lgig 25 m long line transects using methods de- juniperus osteospennaosteosperma 40 T 161.6 scribed in meeuwig and budy 198119811 inter quercus turbinella 30030.0 sys 62 yucca bacattabagatta 404.0 8 11 mostly soil and rock surface spaces are bare opuntia littlittoralisoralis var mattmartinartinianamartmianamartimianamiana 191.9ig s 11iill1.1 with 3 grass cover mostly bouteloua gracigracilisfishisbis rhus trilobata 070.7 s 040.4 HBK lag ex steudel and B curtipendula ceanothus greggiigreggio 040.4 sys 02 michamichx torr and traces of litter herbs and canitiacanotia holocanthaholocantha 030.3 sys 030.3 bouteloua gracilis 292.9 sys 131.3 gams mostly scrub oak quer- cryptocryptogamscryptograms shrubs Gutiercutierriziagutierriziarizia sarothraesarothrae 1 cus turbinella greene cover about 30 of the richmondarichmond and richardson 1974 study area pifionpigion pines pinus monomonophyllaphylla varvan bUnpublishedunpublishedbunpublished data this study fallaxfallad little silba cover about 36 of the twoc1wo methods were used to estimate covetcover forror all species estimates weiewere made on eighteen 222 m plots accordingacco iding to coveragecover age class ratings daubenmireDaubennine area and juniperus osteosperma torr little 1968 tree and shrub cover were estimated on 12 permanent 25 m line tran- 4 of com- sects as cover 2531425314transect253 Transect length sum of crown diametersdiame teis about the added cover vegetation meeuwigMeen wig and budy 1981 values leportedreported heiehelehere for trees and shrubs aiealeare ave-aver- ponents is greater than total vegetation cover rageages of both methods and standard enorserrors aream fromhiomblom pooled variances due to the presence of different vertical layers of shrub and tree canopiescanopies and aggregation of paired samples 10210.2 cm above litter present vegetation in clumps trees ranged in age from little no cactus seedlings to about 260 yr estimated from annual and below to litter present above and ring counts of cores unpublished data age axes to compare soil properties we estimates are approximate due to occurrence below cactus dams took additional sam- of false rings in wood of pinonpinnon pines and ples from bare interinterspacesspaces and from areas junijunipersjuniperuspers beneath tree and shrub canopiescanopies to compare size range of soil surface patches covered these areas with the areas above and below by cacti and associated litter accumulations cactus dam Interinterspacesspaces were considered to lie was estimated by measuring every cactus dam beyond the influence of canopiescanopies and associ- on a 25 X 25 m plot we recorded length width ated litter and beyond the influence of cactus and circumference for each cactus dam and dams and associated litter vegetation and litter associated litter accumulation 32 total the were scant to absent in interinterspacesspaces beneath area of soil surface covered by cactus dams and tree and shrub canopy sampling included litter was calculated as the area of a circle plus pinon pines scrub oaks junipersjunijuniperuspers and occa- 12 the difference between the area of a rectan- siosionallynally mixed species canopiescanopies roughly in pro- gle and a circle portion to the presence of these components SOIL AND LITTER SAMPLING APPROACH as estimated by percent canopy cover on the sampling was stratified by woodland micro site table 2 the sampling location beneath habitats above cactus dams below cactus dams canocanopiespies was at 23 canopy radius out from the interinterspacesspaces and beneath canopiescanopies we took stem or clump center litter of yucca baccalabaccata 214 GREAT BASIN naturalist volume 56

17600

7600

udd

35 08n 5200

11 T 56w arizona

legend north

636.3 cm 040.4 km 252.5 in 025025.025 milemlle

122 rn fig 2 soil and litter sampling area in the Huhualapaikualapaialapai mountains of northwestern arizona the contour interval is 12212.2 40 ft enlarged from US geological survey rattlesnake hill arizona quadrangle

torr and a few other species was occasionally sized individual samples compositingCompositing followed though rarely present in litter samples along guidelines in peterson and calvin 1986 and with litter of the dominant species with the was suitable for the present study since we exception of bulk density samples soil and lit- were not examining variation within microhab ter samples were comcompositescompositedposited within microhab itatsitaas As pointed out by crepin and johnson itaticat strata by combining equal numbers ofofequalequal 1993 composite sampling can be used in 199619961 DAM FORMING CACTI AND NITROGEN enrichment 215 conjunction with stratification ie the land- for percent organic C and methods described scape can be divided into meaningful units and by gee and bauder 1986 for particle size good averages of soil properties obtained by analysis compositingcompositing samples within each unit all soil NET mineralized N the total amount of and litter sampling was conducted in july 1991 N liberated from organic matter is gross min- BULK DENSITY bulk density was deter- eralization the quantity remaining after micro- mined by the excavation method blake and bial immobilization is net mineralization carl- hartge 1986 twenty two paired samples were yle 1986 net mineral N the N available for taken 10210.2 cm above anani 1 below cactus axes 10 plant uptake is an index of soil fertility to com- samples were taken from i terspaceserspaces and 10 pare soil N fertility among woodland sites net were taken from beneath tree and shrub mineral N was assessed by laboratory aerobic canopiescanopies soil was excavated with a bulb planter incubations binkley and vitousek 1989 diameter 555.5ss cm at cutting edge creating a seven permanent plots were created on the hole 7 cm deep A thin tough plastic bag was study site the ist plot serving as a central placed in the hole filled with water and then point from which 6 satellite plots were created emptied into a graduated cylinder to deter- each 32 in from the central point at 60 intervals mine hole volume extracted soil was dried at beginning with a random azimuth because of 105 C and weighed resulting in a weight to topography I1 plot was relocated 32 in from the volume measurement center of a satellite plot from each plot center TOTAL N TOTAL ORGANIC C AND SOIL TEX- 8 cacti 05os050.5 to 5 in frowfrom center were selected TURE thirty pairs of soil cores mineral soil at 45 intervals beginning with a random surface to 7 cm deep were extracted with a azimuth for a total sample of 56 cacti ss55 bulb planter diameter 555.5 cm at the cutting paired soil samples were taken 10210.2 cm log above edge adjacent to cactus axes 10210210.2 cm from cactus axes on all 7 permanent plots be- from cano and below cactus axes 30 beneath ginning from the easternmost cactus and mov- from samples were pies and 30 interspacesinterspaces ing clockwise samples were comcompositedcompositesposited com- of 6 satellite plots estab- taken near each the bining 4 individual samples into 1 composite for mineralization study see lished the net sample compositingCompositing and field processing see all from surface to mineral below litter litter below were performed immediately upon the soil was retained for determination of total N extraction of 4 cores for example on the ist samples were air dried and stored in paper plot 4 cores 10210.2 cm above cactus axes in the wrappers soil samples originally taken for 90 270 hemisphere of the plot were taken determining bulk density see above were cacomeomcompositescompositedposited and field processed before the added to these soil samples for a total of 51 c0 4 were drawn ensured pro- samples from each side of cactus dams 40 next cores this fresh soil composite sample samples from beneath canopiescanopies and 40 from cessing fourteen were prepared interinterspacesspaces one of the 22 paired bulk density pairs approximately midpoints of the six 32 m samples was lost and could not be included at satellite plots 2 samples samples were combined to create compos- lines creating were cano pines sampled ites above cactus dams 51 samples of soil were taken beneath canopiespies pinon oak mixed comcompositedcompositesposited to make 3 samples of soil and 30 most heavily followed by scrub samples of litter were combined to make 3 lit- species canopiescanopies and juniper and 2 from inter ter samples below cactus dams no litter pre- spaces composites of 4 individual samples sent 51 samples of soil were comcompositescompositedposited to were prepared and field processing completed make 3 soil samples beneath canocanopiespies 40 soil immediately as each set of 4 cores was drawn samples were comcompositescompositedposited to make 3 samples of three composite samples were prepared soil and 30 litter samples were comcompositescompositedposited to samples were taken with a 2 cm diameter make 3 litter samples from interspace areas soil corer to a depth of 10 cm preparation of no litter present 40 samples were comcompositescompositedposited samples for analysis followed methods outlined to make 3 samples of soil analysis was by utah in vitousek et al 1982 in the field cores were state university soils testing lab following divided into 3 components litter layer top 5 the kjeldahl method bremner and mulvaney cm of mineral soil and mineral soil between 5 1982 to determine percent total N the walk and 10 cm soil depth and comcompositescompositedposited com- ley black method nelson and sommers 1982 posite soil samples were sievedsievek through a 2 216 GREAT BASIN naturalist volume 56

6 TABLE 3 size distribution of cactus dams and associated 2 litter accumulations on a 25 X 25 m plot the area mea- 50 sured was the soil surface covered by cactus dams and

40 associated litter class of 0 size number mam2 cactus dams

005 1 10 1 I JLW 010 1 101.0loio1 0 16 0 9 1019 202920 29 303930 39 L404940 49 505950 59 606960 69 7079 8090 09 11 20 6 degrees 21 30 6 cla with fig 3 angle of growth of connected basal cladodescladodusdodes 31 40 2 respect to slope contour zero degrees is a direction of growth parallel to the slope contour 90 degrees is a direc- 1030 1 tion of growth perpendicular to the slope

mmMITI screen litter was not sievedsievek subsamples oratory at utah state university for determina- were sealed in bags for determination of mois- tion ofnh4 and n03 US EPA 1983 ture content while a and2nd subsample of approx- CHANGE IN DEPOSIT DEPTH depth of de- imately 10 g was placed in 100 ml I1 N KCI posits above cactus dams ie above con- adjusted with hc1 to ph 252.5 with phenylmer nected basal cladodescladoduscladodes was measured before curiecuriceurieeuriceunie acetate PMA added as a preservative july and after september the rainy season of solutions were refrigerated transported to the 1991 on 6 of the 7 plots designated for net min- laboratory mixed frequently for 4 d then eralization sampling see above two sampling allowed to settle for 48 h after settling the points could not be relocated at the end of the solution was removed with a pipette and rainy season making a total sample size of 46 nhanh4NH and n03 were determined at bilby cactus dams ie 6 plots 8 cacti per plot minus research facility at northern arizona univer- 2 depth was measured from base to top of sity using methods described by keeney and deposits in the area of greatest accumulation nelson 1982 statistical analysis the remainder of comcompositescompositedposited field samples after removal of the above 2 subsamples was A heterogeneity chi square analysis followed transported to the laboratory and incubated by a chi square analysis zar 1984 was per- aerobically following procedures in vitousek et formed with the 3 data sets of angle of cactus al 1982 soils were wetted to approximately growth from the 3 mountain ranges field moisture capacity assessed visually SOIL AND LITTER ANALYSES tests of nor- for data placed in plastic covered cups each of which mality were performed each set above had a small air hole and kept in a dark moist cactus dams below cactus dams interinterspacesspaces soil chamber at a constant temperature of 22 C and beneath canopiescanopies of each and litter oc during an 8 wk incubation period samples characteristic sampled A paired t test received distilled water applied as a fine mist 0050.05 was used to compare means of soil char- to the surface with no mixing as needed to acteristicsacteristics above and below cactus dams and maintain an approximately constant moisture to compare the depth of deposits at cactus content so as not to disturb incubating sam- dams before and after the rainy season an ples moisture content was assessed by visible analysis ofofvariancevariance F test oc 0050.05 for unbal- soil color easily observable through the clear anced sample sizes the GLM procedure in plastic incubation cups SAS software SAS 1985 was used to compare at the end of 8 wk subsamples approxi- sample means of soil above and below cactus mately 10 g of incubated samples were taken dams with beneath canopy and interspace sam- for determination of moisture content and ple means plots of residuals were generated to subsamples of approximately 10 g were placed assess equality of variance significant differ- in the KCI solution described above these ences between means were separated and solutions were shipped to the soils testing lab ranked using a multiple comparison method 199619961 DAM FORMING CACTI AND NITROGEN enrichment 217

TABLE 4 results of paired t tests comparing sample means of soil characteristics above and below cactus dams and comparing depth of detritus above cactus dams between early july and mid september A minus B refers to the value above cactus dams minus the value below paired t test statistics attribute N mean difference sy t P A minus B bulk density galgml 22 0222 0117 1901 00711 natural log 0 7 cm depth total N 3 0043 0003 13000 00059 0 7 cm depth organic C 3 0933 0231 4035 00563 0 7 cm depth net mineralized N ugg 0 5 cm depth NH 14 9621 2353 4088 00013 0 5 cm depth n03 14 41979 5398 7776 00001 5 10 cm depth nhanh4 12 2050 2063 0993 03418 5 10 cm depth n03 12 7550 2672 2826 00165 change in depth cm 46 124011 0124 10038 00001 detritusofdetntusof above cactus dams natural log 0 7 cm depth meanamean difference of september detritus depth minus july detritus depth

REGWF cited as being compatible with the were nonnonformalnonnormalnormal and were not normally dis- overall analysis of variance F test SAS 1985 tritributed when transformed with standard A t test oc 0050.05 was used to compare sam- transformations therefore results of total N ple means of total N and net mineralized N analyses should be interpreted with caution from the litter above cactus dams with the lit- residual plots indicated equality of variance reasonable ter layer beneath woodland canopiescanopies assumptions were bulk density above and below cactus dams significantly different FP oos0.05 RESULTS was not at 005 table 4 bulk density was significantly lower P pattern of angle of connected basal 000010.0001 in soil deposits above cactus dams the dams and canocanopiespies cladodesclacladodusdodes with respect to slope contour was below cactus beneath tree compared to soil from interinterspacesspaces table 5 similar in the 3 mountain ranges sampled data fig 4 soil above and below cactus dams was pooled based on results of a heterogene- were also lower in bulk density than soil beneath ity chi square analysis analysis of pooled data 2 tree canopiescanopies although this difference was not 854 P 0.001 indicated that X 85485.4 0001 orienta- significant at P 0050.05oos there was little differ- tion of connected basal cladoduscladodescladodes of opuntia lit ence in soil texture among the 4 micromicrohabitatshabitats toraliscoralis var martiniana was nonrandom growth table 5 was most frequently parallel to the woodland soil total N above cactus dams was greater slope contour fig 3 the size range of cactus P 001ooiool0.01 than below cactus dams table 4 dams and associated litter on a 25 X 25 m plot organic C was not significantly different P at the Huhualapaikualapaialapai mountains study area is given 0050.05oos above cactus dams compared to below in table 3 cactus dams soil total N and organic C were SOIL AND LITTER ANALYSES the null hy- 2 3 times greater P 000010.0001 in both cases in popothesisthesis for normality was not rejected for soil above and below cactus dams than in most of the data sets however total N data interspace soil table 5 fig 4 soil total N and 218 GREAT BASIN naturalist volume 56

A bulk density 3

F 9299.29 p 000010.0001 2 1 I1E m111111

11 mm 0 above below interinterspacesspaces beneath cactus damsdarnsdanns cactus darns canopiescano pies

B total N 0220.22 0.20020 020 F 35357735.7777 018-018 016olg p 000010.0001 ninnlnKIM z 0140.14 ii3riier p lliali JS 0120.12 Q 0100.10olo sp 0080.08 006-006 004 002- 000ooo

above below interspacesinterspaces beneath cactus damsdarns cactus damsdarns canocanopiespies

C organic C

6 5 F 380038.00 0 p 000010.0001 g 4- 1 3- 0 2 inmbinmM

above below interinterspacesspaces beneath cactus dams cactus dams canocanopiespies

fig 4 comparisons of soil characteristics above cactus dams below cactus dams beneath canopiescanopies tree and shrub combined and in bare interinterspacesspaces

organic C above cactus dams were equal to net mineral nhanh4NH and n03 at 0 5 cm areas beneath canopiescanopies below cactus dams soil depth were significantly greater P 00010.001 total N was significantly lower than beneath and FP 000010.0001 above cactus dams compared canopiescanopies and organic C was not significantly to below table 4 at 5 10 cm depth net min- different compared to beneath canocanopiespies while eral n03 was significantly greater P soil organic C and soil total N differed among 001650.0165 above cactus dams compared to below woodland locations the CN ratio was similar net mineral N in soil 0 5 cm deep above cac- between locations table 5 tus dams was over 3 times that in interspace 199619961 DAM FORMING CACTI AND NITROGEN enrichment 219

D nhanh4 050 5 cm depth 16 14 F 4934.93 p 000670.0067 roCD 12 T 10 0CD Z5- 8 it 6 z 414 1 2 0 j above below interinterspacesspaces beneathdeneath cactus dams cactus dams canocanopiespies

E n03 050 5 cm depth 90 80 Ff1121112111.21 0 70 p 000010.0001 60 T 0D 50 agogeagogg 40 g 0 30 Z 20- romememsmoms11 1411 N 1001 0 above below interinterspacesspaces beneath cactus damdams cactus daadamdams canocanopiespies

1 F nhanh4 515 1 acmocm10cm depth 6 5 F 0740.74 p 053740.5374 1 0CD I 4- 0 3 2 zX

0 above below interspacesinterspaces beneath cactus dams cactus dams canopiescano pies

G n03 515 10cmdepth1 acmocm depth 16 14 F 0970.97 p 042190.4219 01 12 10 0 1 8 IZ KS 0 6 0zz 4- 2 0 above below interspacesinterspaces beneath cactus dams cactus dams canopiescanopies

fig 4 continued 220 GREAT BASIN naturalist volume 56

TABLETABLL 5 comparison of sample means of soil characteristics at 4 woodland micromicrohabitatshabitats above and below cactus dams interinterspacesspaces and beneath canopiescanopies at the Huhualapaikualapaialapai mountains site superscript letters separate means significantly differentdiffeidiffee ent at 0050.050oos05 for texture s sand sl silt and cl clay samples are composites except for bulk density N sample size and is followed in parentheses by the number of individualofindividual samples that were comcompositescompositedposited location attribute above cactus dams below cactus dams Interinterspacesspaces beneath canopiescanopies soilsolisoiltextuietexture sandy loam sandy loam sandy loam sandy loam separates s 650 s 630 s 637 s 627 sl 250 sl 260 sl 276 sl 283 cl 90 cl 110 cl 87 cl 90 n 3 51 n 3 51 n 3 40 n 3 40

bulk density Tx 0099b9913 Tx 113b1 13b x 185a1 8521 af3fx 140b1 4011 gal9mlgml 011 092 010 016 nn2222 n 22 n 10 n 10

total N x 016aolgaoiga0 16s1 xT 001211215 xC 006c0 06 xjf 01tolt0 171 0 7 cm depth 0006 0003 0007 016 n 3 51 n 3 51 n 3 40 n 3 40

organicoigamccC xT 39 xT 30a30s1 YX isb15b15k Tx 363 0 7 cm depth 024 007 004 026 n 3 51 n 3 51 n 3 40 n 3 40

CNC N ratio 244 250 250 212 net mmelmineralizedahzedahmed N ugggg 0 5 cm depth nhanh4 xT 148a14 81 xaf5f 53b Tx 20b x5 ilsalisa115a11 26 16 02 13 NOn03 Yx 887a88 71 xF 467146.7146 7 Tx 271127 x 542154 2 69 59 26 64 n 14 56 n 14 56 n 3 12 n 3 12 5 10 cm depth nhanh4NL Tx 53 Tx 32 xY 09 Tx 45 11 18 097 15 NOn03 xT 316131.6131 6611 xT 24240bob Tx 166 xT 309 40 44 30 82 n 14 56 n 14 56 n 3 12 n 3 12

soil and almost twice that in soil beneath tree discussion canopiescanopies table 5 fig 4 net mineral N below cactus dams was greater than in interinterspacesspaces the similarity of soil texture above cactus but the difference was not statistically signifi- dams below cactus dams beneath tree and cant shrub canopiescanopies and in interinterspacesspaces agrees with litter accumulated at cactus dams had total findings of schlesinger et al 1989 that desert N 0740740.74 over twice as high as litter beneath soils receiving overland flow and adjacent soils tree and shrub canopiescanopies 0320320.32 t 848.4 P deprived of overland flow were similar in fine ooi0010.01 NHL and n03 in the litter layer were material or clay content the effects of cactus greater beneath canopiescanopies than above cactus dams and associated litter and detritus deposits dams but not significantly table 6 fig 5 on bulk density total N organic C and net from early july to mid september depth of mineralized N of nearby soil were expected detritus behind cactus dams increased signifi- based on a number of studies in shrub lands cantly P 000010.0001 from 2 cm to 23 cm with and woodlands documenting islands of fertility an average of 43434.3 cm safs5fs 06250.625 figg ie localized areas of nutrient enrichment 199611996 DAM FORMING CACTI AND NITROGEN enrichment 221

TABLE 6 comparison of total N and net mineralized N in the litter layer beneath canopiescanopies with litter accumulations above cactus dams N sample size and is followed in parentheses by the number of individual samples that were comcompositedcompositesposited above beneath cactus dams canopiescanopies P attribute mean sV mean STs t total N 07370.7370 737 00470 047 0320 0015 84275 00095 n 3 30 n 3 30

net mineralized N ugg NH 6023360 233 1501815.01815 018 84550 11250 1 29292999129291.2929 0 2426024260.2426 n 21122211 12 n 12b 48

n03n0na 2108 6410 4005040.05040 050 3495034 950 11865 04398 n 2 12 n 12 48

three compositecomposites weiewere preparedprepiredared however initial beforebebbie incubation net mineral N values were not obtained for I1 sample valuesbvalues before incubation were not obtained for 2 of the original 14 comcompositedcompositesposited samples

garcia moya and mckell 1970 tiedemann basal cladodescladoduscladodes and to 0120.12 below table 4 and klemmedson 1973 barth and klemmed- nitrogen enrichment and soil amelioration son 1978 barth 1980 doescher et al 1984 associated with deposits at cactus dams may everett et al 1986 garner and steinberger increase cactus productivity nobel et al 1987 1989 schlesinger et al 1990 observed that while annual aboveground pro- deposits at cactus dams of opuntia littlittoralisoralis ductivity of prickly pear cacti can be high under var martiniana raised soil total N from 0060.06 optimal conditions cacti productivity is often interspace soil to 0160.16olg above connected limited by low levels of soil N nobel et al

litter total N and net mineral N

120 p 024260.2426 100 p 000950.0095 p 04398 Z 80- 7 a Z 9.9 60 E R

C 40

20 total N net NH net n03

IM cactus dams liliiiii1111 canocanopiespies

fig 5 nitrogen in the litter accumulated above cactus dams compared with the litter layer beneath canopiescanopies trees and shrubs combined 222 GREAT BASIN naturalist volume 56

july depth 30 M september depth

25 E 0 C 5 20

15 september mean depth 114 cm 10 july mean depth 7177.11 cm 5

0 deposition at individual cactus dams

fig 6 deposition at cactus dams during I1 season of summer thunderthundershowersshowers depths of detritus accumulations at 46 cactus dams in july and in september

1987 nobel 1989 increased productivity in acknowledgments desert prickly pear cacti is positively correlated with both number of new cladodesclacladodusdodes produced we thank ron lanner and helga van and cladode size nobel et al 1987 we do not miegroetmingroetMiegroet for advice encouragement and assis- know if similar patterns occur in woodland tance the senior author thanks chuck grier species of prickly pear additionally nobel for accepting an unconventional student and his scientific 1988 describes a tendency for daughter clad for sharing abundant cajolery and 11 acumen odes to replicate the orientation of mother cladodescladoduscladodes and points out that if a particular CITED direction of growth is favorable it may be per- literature petuapetuatedted this happens because favorably ori- BARTH R C 1980 influence of pinyon pine trees on soil ented cladodescladoduscladodes are expected to be more pro- chemical and physical properties soil science soci- ductive than other cladodescladoduscladodes and produce more ety of america journal 44 112 114 and larger similarly oriented clacladodescladodusdodes this may BARTH R C AND J 0 klemmedson 1978 shrub induced spatial patterns of dry matter nitrogen and be occurring in dam forming cacti but it was organic carbon soil science society of americaofamerica jour- not investigated in this study nal42804nal 4280442 804 809 cactus dams lower soil bulk density and BINKLEY D AND P VITOUSEK 1989 soil nutrient avail- enrich patches of woodland interspace with ability pages 75 96 in R W pearcy J R Ehlenehleringerngerngen H A mooney and P W rundel editors plant physi- organic matter total N and net mineral N sug- ological ecology field methods and instrumentation gesting that they may play roles in nutrient chapman and hall londonnewLondon New york cycling and other ecosystem processes some BLAKE G R AND K H HARTGE 1986 bulk density pages 363 375 A klute editor methods of soil possible functions of cactus dams are to 1 inin analysis part 1 physical and mineralogical methods crease woodland detritus storage 2 increase add2nd2dd edition soil science society of america inc the rate of N turnover 3 mitigate nutrient loss madison WI in interspace areas 4 reduce soil erosion and BREMNER J M AND C S MULVANEY 1982 nitrogen page al dampen effects of disturbances 5 provide total pages 595 694 in A L et editors methods of soil analysis part 2 and2nd edition agron- seedbeds and 6 provide habitat for other omy monographs 9 american society of agronomy organisms and soil science society ofamericaof america madison WI 199619961 DAM FORMING CACTI AND NITROGEN enrichment 223

CARLYLE J C 1986 nitrogen cycling in forested ecosys- 391 396 in R L everett compiler proceedings tems forestry abstractsabshabah acts 47 306 336 pinyon juniper conference general technical report crapincr9pinCREPIN J AND R L JOHNSON 1993 soil sampling for en- INT 215 intermountainIntelmountain research station ogden UT vironvironmentalmental assessment pages 5 18 in M R carter KLOPATEK C C AND J M KLOPATEK 1987 Mycorrhizamycorrhizalmycorrhizaemycorrhizaelel editor soil sampling and methods of analysis lewis micmicrobesrobes and nutrient cycling processes in pinyon publishers boca ratonantiratonannRatonAnnnAnti arborlondonArbor London tokyo juniper systems pages 360 364 in R L everett daubenmire R 1968 plant communities a textbook of compiler proceedings pinyon juniper conference plant synecology harper & row new york general technical report INT 215 intermountain DEBANO L F AND J M KLOPATEK 1987 effect of man- research station ogden UT agement on nutrient dynamics in southwestern pinyon LANNER R M 1981 the pinonpinnon pine a natural and cul- juniperjumper woodlands pages 157 160 min C A troendle tural history university of nevada press reno M R kaufmann R H hamre and R P winokur LYONS S M AND G F GIFFORD 1980 impact of incre- coordinators management of subalpine forests build- mental surface soil depths on plant production tran- ing on 50 years of research general technical report spiration ratios and nitrogen mineralization rates RM 149 rocky mountain research station fort journal of range management 33 189 196 collins CO MEEUWIG R 0 AND J D BUDY 1981 point and line DEBANO L FE H M PERRY AND S T OVERBY 1987 effects intersect sampling in pinyon juniper woodlands of fuelwood harvesting and slash burning on biomass general technical report INT 104 intermountain and nutrient relationships in a pinyon juniperjumper stand research station ogden UT pagesages 382 386 in R L everett compiler proceed- NELSON D W AND L E SOMMERS 1982 total carbon ings pinyon juniperjumper conference general technical organic carbon and organic matter pages 539 579 in report INT 215 intermountain research station A L page et al editors methods of soil analysis ogden UT part 2 and2nd edition agronomy monographs 9 ameri- DOESCHER P S R F MILLER AND A H WINWARD 1984 can society of agronomy and soil science society of soil chemical patterns under eastern oregon plant america madison WI communities dominated by big sagebrush soil sci- NOBEL P S 1988 environmental biology of agacesagaves and ence society of america journal 48 659 663 cacti cambridge university press cambridge EVERETT R L AND S H SHARROW 1985 soil water and 1989 A nutrient index quantifying productivity of temperature in harvested and harvestednonnonharvested pinyon agavesagaces and cacti journal of applied ecology 26 lumperjuniperjumper stands forest service research paper INT 635 645 342 intermountain research station ogden UT NOBEL P s3saS C E RUSSELL P FELKERFELKEB G C MEDINA AND EVERETT R L S H SHARROW AND D THAN 1986 soil E ACUNA 1987 nutrient relations and productivity nutrient distribution under and adjacent to singlesingleleafleaf of prickly pear cacti agronomy journal 79 550 555 pinyon crowns soil science society of america jour- PETERSEN R G AND L D CALVIN 1986 sampling pages nal50788nal 5078850 788 792 33 51 in A klute editor methods of soil analysis FLETCHER J IE1 D L SORENSEN AND D B PORCELLA part 1 physical and mineralogical methods and2nd edi- 1978 erosional transfer of nitrogen in desert ecosys- tion soil science society ofamericaof america inc madison tems pages 171 181 in N E west and J skujins WI editors nitrogen in desert ecosystems dowden RICHMOND D L AND M L richardson 1974 general hutchinson & ross inc stroudsburgStrouds burg PA soil map and interpretations mohave county ari- GARCIA MOYA E AND C M mckell 1970 contribution zona US department of agriculture soil conserva- of shrubs to the nitrogen economy of a desert wash tion service plant community ecology 51 81 88 SAS 1985 SAS user s guide statistics version 5 edition GARNER W AND Y steinberger 1989 A proposed mech- SAS institute carscarygary NC anismaniam for the formation of fertile islands in the schlesinger W H P J FONTEYN AND W A REINERS desert ecosystem journal of andaridarld environment 16 1989 effects of overland flow on plant water rela- 257 262 tions erosion and soil water percolation on a mojave GEE G W ANDandlJ W BAUDER 1986 particle size analysis desert landscape soil science society of america pages 383 411 inm A klute editor methods of soil journal 53 1567 1572 analysis part 1 physical and mineralogical methods schlesinger W H J FE REYNOLDS G L cunningham and2nd edition soil science society of america inc L F HUENNEDE W M JARRELL R A VIRGINIA AND madison WI W CG WHITFORD 1990 biological feedbacks in global GRIER C C K J ELLIOTT AND D G mccullough desertificationdecertification science 247 1043 1048 1992 biomass distribution and productivity of pinus SELLERS W D AND R H HILL EDITORS 1974 arizona adulisedulis juniperusJumjamperusperos monospennamonospermamonospermymonomonos pennapeonasperma woodlands of north climate 1931 1972 university of arizona press central arizona forest ecology and management 50 tucson 331 350 TIEDEMANN A R 1987 nutrient accumulations in pinyon KEENEY D R AND D W NELSON 1982 nitrogen inor-inorlnor juniper ecosystems managing for future site pro- ganic forms pages 643 698 in A L page et al edi- ductivity pages 352 359 in R L everett compiler tors methods of soil analysis part 2 2ndand edition proceedings pinyon juniper conference general agronomy monographs 9 american society of agron- technical report INT 215 intermountain research omy and soil science society of america madison station ogden UT WI TIEDEMANN A RANDJR AND J 0 klemmedson 1973 effect of KLOPATEK J M 1987 nutrient patterns and succession in mesquite on physical and chemical properties of the pinyon juniper ecosystems of northern arizona pages soil journal of range management 26 27 29 224 GREAT BASIN naturalist volume 56

USDA BUREAU orOF LAND management 1991 record of W D billings editors north american terrestrial licensed livestock Huhualapaikualapaialapai ph allotment 1967 1991 vegetation cambridge university press cambridge kingmankmgmdn resource area kingman AZ WEST N E AND J 0 klemmedson 1978 structural dis- USU S EPA 1983 methods for chemical analysis of water tributiontribution of nitrogen in desert ecosystems pages and wastes EPA 6004 79 020 revised march 1983 1 16 in N E west and J skujins editors nitrogenNiti ogen method 350 1 method 3532353.2353 2 inm desert ecosystems dowden hutchasonhutchmsonhutchinson & ross VITOUSEKVMOUSEK P M J R gosz C C GRIER J M MELILLOMEEIEEO inc stroudsburgStrouds burg PA pren AND W A REINERS 1982 A comparative analysis of ZAR J H 1984 statisticalbiostatisticalBio analysis and2nd edition potential nitrification and nitrate mobility in forest tice hall inc englewood cliffs NJ ecosystems ecological monographs 52 155 177 westWLSF N E 1988 intermountain deserts shrub steppes received 20 april 1995 and woodlands pages 209 230 in M G barbour and accepted 14 march 1996 great basin naturalist 563 C 1996 appp 225 236 distribution AND ecological characteristics OF LEWISIA longipetalaLONGIPETALA PIPER CLAY A HIGH ALTITUDE ENDEMIC PLANT

anne S haifordHalfordhalfordl2halford1212 and robert S nowakl3nowak13

ABSTRACT lewisia longilongipetalapetala piper clay is a high altitude endemic found in the northern sierra nevada the characteristics of 12 sites with L longilongipetalapetala which represent all known populations were studied to define habitat requirements of the species meso and micromicroscalescale characteristics of the habitat were examined including characteristics of the associated plant community average plant size and plant density of L longilongipetalapetala were also determined for each population similar measurements were made on 6 populations of lewisia pygmaeapygmaean A gray robinson a more common lewisia populations of L longilongipetalapetala that had larger plants and higher plant density were associated with gently sloped north facing sites that were near large persistent snowbanks and had low vegetative cover plant species associated with populations of L longipetalalongipetala were similar among the 12 sites and were indicative of mesicmesiemesle rocky alpine sites these types of plant communities found near persistent snowbanks are often termed snow bed vegetation in contrast L pygmaeapygmaean was found to be less site specific lewisia pygmaeapygmaean was found adjacent to or interspersed with L longipetalalongipetala at 5 sites but was found in areas associated with a higher percentage of herbaceous cover and a wider vari-varivarl ety of species this integration of ecological and community information for L longilongipetalapetala populations contributes to the interim management and longtermlong term monitoring of this species by providing needed information concerning its habitat and environmental specificity

key words lewisia longipetalalongi petala lewisia pygmaeapygmaean site characteristics snow bed vegetation alpine endemic plant size plant density

the recent implementation of programs to species within the genus lewisia portula- preserve rare plant taxa indicates the elevated caceae are well known in horticulture elliot concern for effective and longtermlong term steward- 1966 mathew 1989 however little informa- ship of sensitive species sutter 1986 one of tion exists regarding these species in their the initial steps toward the protection of rare native environments only 4 species within the plants is to document their occurrences utter genus lewisia have relatively wide distribu- and hurst 1990 mountain ranges are typi- tions lewisia pygmaeapygmaean A gray robinson L cally rich in endemicsendemics major 1989 and within nevadensis A gray robinson L triphylla S the sierra nevada they comprise a high per- watson robinson and L rediredivivaredivivoviva pursh the centage of the flora stebbins and major 1965 remaining 15 species have considerably smaller factors that characterize the species habitat distributions and 9 that occur in california are are inferred from the species geographic dis- listed by the US fish and wildlife service as environmentally tributiontribution and often suggest candidates for threatened or endangered status the distribution of sen- imposed limitations on lewisia longilongipetalapetala piper clay is a federal plant taxa baskin and baskin 1988 sitive candidate 2 species which implies that data on hutchings 1991 nelson and harper 1991 for identifiable threats are insufficient to support example some limitations that influence en- listing as threatened or endangered demic plants within alpine environments are federal and pavlik 1994 longipetalalongipetala snowbank depth and duration markovakomarkovaKokomdrkovd 1975 skinner lewisia webber et al 1976 and levels of disturbance Isis an endemic species with limited distribution plant classi- to root systems from needle ice fitzgerald et Aal that the california native society 1990 to help ensure the survival of rare plant fies as a category I1 RB species which is a cate- species habitat and biological information gory for rare threatened or endangered plants should be integrated with longtermlong term monitor- within california lewisia longilongipetalapetala popula- ing programs sutter 1986 baskin and baskin tions are fairly remote and most exist in US 1988 hutchings 1991 forest service wilderness areas although L

department of environmental and resource sciences mail stop 199 university of nevada at reno reno NV 89557 resent2present address bureaubuieauoiof land management 785 N main st suite fP bishop CA 93514 author to whom reprint requests should be submitted

225 226 GREAT BASIN naturalist volume 56 longilongipetalapetala populations are not an immediate and L oppositioppositifoliafolia S watson robinson later management concern one population basin descriptions munz 1959 placed L longilongipetalapetala peak is on private land and mining claims as a subspecies of L pygmaeapygmaean more recently within close proximity of the site pose a poten- L longilongipetalapetala was again recognized as a dis- tial threat furthermore the potential also exists tinct species dempster 1993 a distinction for activation of mining claims within wilder- supported by morphological as well as chromo- ness areas as well as increased ski area devel- somal differences between L longipetalalongipetala and opment within the vicinity of the other L longi L pygmaeapygmaean stebbins 1968 halford 1992 petala populations lewisia longilongipetalapetala fig 1 is an herbaceous the first specimen of L longilongipetalapetala was col- perennial with a basal tuft of green linear lected by J G lemmon in 1875 in the moun- leaves an individual plant produces numerous tains west of truckee california in 1913 piper scapes 30 60 mm long each bearing 1 3 pale described L longilongipetalapetala as Oreooreobromabroma longi pink flowers with petals 11 20 mm long the petpetalumalum an intermediate between L pygmaeapygmaean two sepalssapals are distinctly fuchsia in color 4 10

7 J

meinITIKhemn

leitulleittldeiderLei tuL toktpeI1 p e tcdsxfa ia i 92

figFig 1 line drawing of lewisia longipetalalongipetala piper clay showing growth habit 199611996 distribution AND ECOLOGY OF L longipetaialongipetalaLONGI PETAIAPETALA 227 mm long and conspicuously glandular dentate elliot 1966 mathew 1989 in contrast infloinflow rescences of L pygmaeapygmaean do not have pronounced scapes and the flowers are smaller mor with petals 6 10 mm long flower color forlaorlforror L 0 pygmaeapygmaean ranges from pink to white and sepal 09 akeoke krnarn from green to fuchsia both of 6 color varies for okoosoCIO ahoechoe species seeds are similar in size isls151.5 mm long 0 0 200 numerous 50 60 per capsule and passively ejected as the capsule dehisces however L longilongipetalapetala seeds are black whereas those of L pygmaeapygmaean are reddish brown the primary goal of this study was to integrate baseline ecological and community information for lewisia longilongipetalapetala this information could then be used to 1 describe suitable habitat requirements 2 provide necessary environmental criteria to search for additional populations of L longilongipetalapetala and 3 establish guidelines for the interim management and potential longtermlong term monitoring of this species fig 2 distribution of lewisia longilongipetalapetala known popu- factors that influence the size of individual plants lations of L longilongipetalapetala are indicated by open circlesen clescies and the occurrence and size of populations include environmental and community attributes such as elevation slope aspect proximity detailed measurements of L longipetalalongipetala three to snowbanks parent material and percent 15 X 15ism m plots were established at basin cover of vegetation surface rock surface water peak 1 in plot 1 plants of L pygmaeapygmaean were litter and bare ground to help delineate the present but no plants of L longilongipetalapetala were habitat requirements of L longipetalalongipetala 6 popu- present plot 2 was adjacent to plot 1 and had lations of L pygmaeapygmaean which also occurs in plants of both L longilongipetalapetala and L pygmaeapygmaean alpine areas but is a more widespread lewisia plot 3 was approximately 10 m north of plot 2 were also studied the 2 objectives of our study and also had both lewisia species were 1 to determine environmental and com- to help delineate the habitat requirements munity characteristics for L longipetalalongipetala and 2 of L longipetalalongipetala a total of 6 L pygmaeapygmaean popu- to determine correlations between the size of lations were also sampled five of these popu- L longipetalalongipetala plants and site characteristics lations basin peak 1 basin peak 2 granite such as elevation slope slope aspect percent chief keith s dome and dick s lake were cover and proximal location to snowbanks as selected because L pygmaeapygmaean plants were near well as between plant density of L longipetalalongipetala or interspersed with L longilongipetalapetala the 2 populations and these site characteristics species were interspersed at basin peak 1 and basin peak 2 and measurements of L bygpyg METHODS maea were made in areas that had predomi- nantly L pygmaeapygmaean at granite chief keith s A total of 12 L longipetalalongipetala populations fig dome and dick s lake the distribution of the 2 which represent the total number of known 2 species did not overlap for these sites the populations of L longilongipetalapetala were examined population of L pygmaeapygmaean that was nearest the nine populations were determined from cali- L longilongipetalapetala population was sampled the ath6th fornia department of fish and game database population of L pygmaeapygmaean was at plute pass maps during this study 3 additional populations which is an area previously described as hav- were located basin peak populations I1 and 2 ing L longipetalalongipetala but where no L longipetalalongipetala are the most northern populations and the re- plants were documented in this study maining 10 populations extend south through for each population we collected a set of 9 the granite chief and desolation wilderness site characteristics that included environmental areas of the sierra nevada fig 2 for more characteristics elevation slope aspect slope 228 GREAT BASIN naturalist volume 56

and distance from nearest uphill snowbank as ticsbics the 3 models with the highest r2ra with 3 well as more detailed community data percent site characteristics and the 3 models with the cover of rock bare ground litter surface water highest r2ra with 4 site characteristics next a and vegetation by species in addition we also series of forward stepwise regressions were per- noted geologic parent material to obtain the formed that started with each of these 12 mod- cover data three 15 m transects were placed els finally a backward stepwise regression was within each of the populations except basin performed that started with all 9 site character- peak 1 the starting point for all 3 transects istics statisticstatistixStatistix version 414.1 analytical software was the geographic center of the population tallahassee FL was used for all analyses and transects radiated out from this starting results were considered significant ifaififpP 0050.05 point along 3 randomly selected compass directions at basin peak 1 the three 15 m transects vegetation association analyses were uniformly spaced along contours within floristic data were analyzed using multi- each of the three 15 X 15ism m plots cover values variate methods that grouped populations were obtained using a cover point projector based on the fidelity of species to a particular model 2 ESCO and associates inc boulder population or stand three different programs CO were used 1 a 2 way indicator species analy- estimates of plant density were obtained by sis TWINSPAN hill 1979a 2 Detrdetrendedended direct count of individuals within a 05osha ha area correspondence analysis DCA hill 1979b until the number of individuals exceeded 500 and 3 nonmetric multidimensional scaling the 05 ha area was orientaorientatedted such that it en- program NMDS TWINSPAN and NMDS compassed the topographical extent of each pop- are classification programs that divide the plots ulation and was centered on the geographic cen- into a series of groups based on percent simi- ter of the population in addition plant size was larity NMDS differs from TWINSPAN in that estimated from the clump diameter of an indi- NMDS recovers gradients of high beta diver- vidual plant up to 20 L longipetalalongipetala individuals sity that may alter the ordering of samples were selected for clump diameter measure- minchen 1987 DCA is an ordination pro- ments if the population size was less than 20 gram that orders the plots along a series of axes all individuals were measured if population size such that the distance between plots in the was greater than 20 then 20 individuals were multidimensional space is proportional to the chosen for measurement by randomly selecting differences between them to graphically rep- 20 numbers between 000.0oo and 15015.0 in 010.1oi incre- resent the results of the classification and ordi- ments these numbers represented sampling nation analyses populations and species were points along the transect tape if I1 or more plotted in the 2 dimensional space formed by plants were within I1 m of the designated sam- DCA axes 1 and 2 hierarchical classifications pling point then the plant nearest the desig- that were constructed from TWINSPAN eigen nated sampling point was measured if no plants values were then used to draw the groupings were within I1 m of the sampling point then of populations and species on the DCA plots additional sampling points were randomly se- where NMDS groupings differed from TWIN- lected for the and2nd and ard3rd transects as necessary SPAN NMDS grouping were also drawn regression analyses were used to determine to examine relationships between species correlations between plant density and the set composition and environmental site character- of 9 site characteristics elevation slope slope istics for these populations we used a rotational aspect distance from nearest uphill snowbank correlation analysis to generate correlation co- bare ground cover litter cover surface rock efficients between the DCA axis I1 scores and cover surface water cover and total vegetative the site characteristic dargie 1984 tueller et cover for L longipetalalongipetala as well as between al 1991 correlation matrices were developed clump diameter and the 9 site characteristics for the set of 9 site characteristics elevation the ist regression analysis used multiple re- slope aspect snowbank distance total vegeta- gressionsgressions to determine the 3 models that had tive cover surface rock cover bare ground the highest r2ra values for regressions with 1 cover surface water cover and litter cover for site characteristic the 3 models with the high- all correlation analyses P 0050.05oos was the level est r2ra for regressions with 2 site characterscharacteris of significance used 199619961 distribution AND ECOLOGY OF L longipetalaLONGI PETALA 229

RESULTS populations with the largest plants generally were also those with the highest plant density lewisia longilongipetalapetala table 1 regression analyses between clump the majority of L longipetalalongipetala populations diameter and 9 site characteristics yielded a were in shallow north facing basins table 1 single model that all forward and backward fewer populations were on steep slopes or had stepwise regressions converged upon table 3 a southern exposure most populations were mean plant diameter from each population between 2700 and 2900 in elevation but L was inversely correlated with distance from longilongipetalapetala populations were found up to 3200 the nearest uphill snowbank the value of the in elevation populations were not restricted to regression coefficient for surface litter cover only I1 rock type but were found on substrates was significantly different from zero at the 6 derived from basaltic and granitic rocks carex probability level rather than the 5 level and scopulorum var bracteosabracteose and AntenantennarianatlanaTia media plant size was inversely correlated with the were the 2 species that co occurred with most amount of surface litter cover L longipetalalongipetala populations and had the highest classification and ordination of the floristic mean cover table 2 data corroborated these results fig 3 four the largest populations oflkoflof L longilongipetalapetala were site characteristics were found to be significantly on low gradient north to northeast facing sites correlated with DCA axis I1 scores table 4 table 1 populations with lower densities were and these site characteristics are shown in fig on steep slopes 30 with west southwest 3aaa as vectors that indicate the directional or southeast facing slopes regression analyses increase of slope surface rock cover bare between plant density and 9 site characteris- ground cover and total vegetative cover tics did not yield 1 best model but rather 2 TWINSPAN classified the 12 populations into models that had similar adjusted r2ra values 3 groups fig 3aaa and the species groupings table 3 for both models slope was a signifi- associated with the TWINSPAN population cant dependent variable and plant density was groupings are shown in fig ab3b the basin inversely correlated with slope ie as slope peak populations had higher vegetative cover increased plant density decreased surface whereas the other populations had higher rock water and surface rock cover were significant cover fig 3aaa these populations that are dependent variables in I1 model and plant associated with increasing rock cover contain density was positively correlated with both of species such as antennaria media cassiope these dependent variables in the and2nd model mertensianamertensiana and kalmia polipolifoliafolia var micropymicropsy total vegetative cover was a significant depen- llaliaiia that are indicative of such environments dent variable and L longipetalalongipetala density was fig ab3b L longipetalalongipetala populations at granite inversely correlated with vegetative cover chief top lake mt price 2 mt price 3 and

TABLE 1 descriptive site attributes for 12 lewisia longipetalalongipetala populations ordered from north to south mean standard error of plant diameter from 20 randomly selected plants as well as plant density is given for each population parent elevation slope plant diameter plant density population material anm1n aspect cm peiper 050 5 ha basin peak I1 basalt 2800 NNE 2 8 970397 03 185 basin peak 2 basalt 2840 NNE 2 8 3901393.9 01 10 pole creek I1 basalt 2733 NNE 2 6 13013.01300909 500 pole creek 2 basalt 2733 NNE 2 6 8204828.2 04 500 granite chief granite 2800 N 30 6501656.5 01 135 dick s lake granite 3033 NNE 2 10 868604040.4 500 top lake granite 2866 W 30 420242 020.2ob 12 mt pricepnce33 granite 3133 WSW 30 636.36304040.4 35 mt price I1 granite 3200 SSE 2 8 340234 020.2 40 keith s dome granite 2800 NNE 2 8 10810.810 8 040 4 500 mt price 2 granite 2966 SSW 30 8303838.3 03 30 pyramid peak granite 2787 WNW 30 4102414.1 02 25 230 GREAT BASIN naturalist volume 56

TABLE 2 mean percent cover for species found within lewisia longipetalalongipetala populations and the number of L longipetalalongipetala populations that contained that species species are listed from highest to lowest cover hickman 1993 was used as the authority for all species letter codes used in figures 3 and 4 are given in brackets for each species species mean of species code cover pop carex scopulorum holm var bracteosebracteosa L bailey FE herm casebcascbcasab 62 9 antennariaantennataantennanaAntennana media E greene acmeanme 51 9 juncus mertensianusmertensianus bong jume 42 4 erigeronengeron peregrinus pursh E greene eipelerpelelpel 29 2 lupinus brewerbrewen A gray lubraubr 26 1 lewisia pygmaeapygmaean A gray robinson lepylepyldepyl 19 5 lewisia longipetalalongipetala piper clay lelo 17 12 arnica mollis hook armolanno 14 1 mimulus guttatusgutgustatustatus DC migu 12 3 salix artica pallus saar 12 3 aster alpialpigenusgenus torrey & A gray A gray sspasp andersonandersonniiandersonmiminiinil A gray M peck asala 10 5 calyptridiumcalyptndium umbellatumumbellatum torrey E greene caum 09 4 phleum alpinumalptnum L phal 09 2 juncus drummondii E meyer audrjudr 08 3 sibbaldiaSibbaldia procumbensprocumbent L siersiprisipr 08 4 dodecatheon alpinum A gray E greene doaldoaidoall 07 2 cassiope mertensianamerten siana bong don came 07 2 kalmia polifoliapolifolia wangenhwangene sspasp micromicrophyllaphylla hook calder & roy taylor kacomkapom 05 2 lycopodium sp lycsp 05 5 mimulus pnmuloidesprimulprimuloidesoidesoldes benth mierlmiprmiprl 05 3 poa towheeheelenletiietileri vasey jowhpowh 03 1 polygonum bistortoidesbistort oides pursh pobi 03 2 eriogonum incanum torrey & A craygray erin 02 1 Penpenstemonpentstemonstemon rydberg ii nelson sspasp oreooreochansoreocharischanscharischarlschang E greene N Hohnholmgrengren peryolgeryolperyo 02 1 phyllodoce bremenbrewen A gray maxim chbrphbr 02 2 anemone drummonddrummondiiii S watson andr 01 1 poa secunda J S freslfresipreslpres sspasp secunda poses 01 1 sedum roseum L scop sspasp integnfoliumintegrifolium raf hulten serol 01 1

pyramid peak were situated in cracks on steep basin peak I1 plots to the group of 7 plots to the granitic slabs and one of the most common right of basin peak 2 in the 2 dimensional species found associated with these sites was space defined by DCA axes I1 and 2 basin rock sedum sedum roseum sspasp integrifolium peak 2 appears to be transitional in its floristic NMDS results differed slightly from the composition between the basin peak I1 plots TWINSPAN classification by the separation of and this group of 7 populations species that the top lake population from all other popula- contributed to these different classifications of tions and by a change in association of the the basin peak populations were phleum alpin basin peak 2 population from the group of 3 um and lupinus brewebremebreweiibreweribrebrewerewerimerlmefiii P alpinum was within

TABLE 3 multiple regression results between each of 2 dependent variables plant density and plant diameter and the set of 9 site characteristics for 12 lewisia longipetalalongipetala populations

Rregressionegression statisticsstatisties variable statisticsst itistics dependent variable N adaadj r2ra p model variables coefficient p plant density 12 081 001 slope 185 001 surface water cover 112 001 surface rock cover 69 001

12 081 001 slope 166 001 total vegetative cover 65 001 plant diameter 12 053 001 snowbank distance 007 001 surface litter cover 024 006 199611996 distribution AND ECOLOGY OF L LONGIlongipetalalong1petalapetaiaPETALA 231

400 A vegetation cover slopesiope 300 regroundbaregroundBa cover

granite chief 0 top lakeloke 200

BP 131 3 BP 111 1 1 prce loo100 0 ad5d mt price I 0mtamt 2 BP 121 2 dicks lk keiths dome oleck I1 00 oboleopole ck 2 N 0 basinbosin pk 2 C 1 0 0 0 pyramid pk price V2 mt 3 X rock cover

1 loo100 1 B peryoandrseroi u powhasalaPowhAsala erin Q 300 0 chbrphbr0 ojudrojudd siproocaum 200 dumejume0 lepyjepy 0anme Ca m e 100 olubr 0 armo lycsp apomoagomo erpearpe 0 0 lelo 0 sooro 0 casabcascb liguomiguo oodoal opobi mipro poses0 loo100 phalphol 0

1 1 1 200 1 1 1 loo100 0 loo100 200 300 400 500 600goo DCA axis I1

fig 3 A population ordinations generated by DCA for L longipetalalongipetala B species groupings associated with the populations for both graphs circled groups were determined from TWINSPAN dendrograms broken lines indicate NMDS groupings letter codes for each species are given in table 2 232 GREAT BASIN naturalist volume 56

TABUTABLE 4 correlation coefficients generated from a rota- large difference in vegetative cover persisted tional correlation program foiforhorbor all lewisia longilongipetalapetala and even when all populations were considered lewisia pygmaeapygmaean DCA axis 1 scores variables with an are significant at the 0050.050 05 level for all the known L longipetalalongipetala populations mean vegetative cover was 31831.8 s 585.8 for species 6 L pygmaean variables correlation coefficients the pygmaea populations used in this study mean vegetative cover was 53753.7 sifsafs5fs 838.3 lewisialewista longipetalalongipetala although this difference was significant 2 elevation 0240.240 24 sample test 16 P that slope 0580 58 t df 00460.046 note our aspect 0490.490 49 original selection of L pygmaeapygmaean populations snowbank distance 0410 41 was not designed to be a random sample of all bare ground cover 066 L pygmaeapygmaean populations and thus extrapolation litter cover 0 36 covel 036 to all L pygmaean sursuisurfaceface rock cover 071 pygmaea populations is not statistically surface water cover oil0110.11 justified totalmotal vegetative cover 0670.670 67 TWINSPAN results for L pygmaeapygmaean populations grouped the basin peak populations sep- lewisia pygmaeapygmaean arately from populations elevation 0530 53 other fig 4aaa but slope 017 the environmental site attributes that were sig- aspect 0140.140 14 nificnificantlyantly correlated with DCA axis I1 scores snowbank distance 0380 38 differed between L pygmaeapygmaean and L longi bare ground cover 0920 92 petala table 4 cover was a significant litter cover 09200.9292 litter surface rock cover 0340 34 site attribute for L pygmaeapygmaean but slope and surface water covelcover 0260.260 26 surface rock cover were not the vegetative sl total vegetative cover 0780.780 78 cover vector increased toward the basin peak population which suggested that these populations contained a greater herbaceous compo- the sampled transects ofbothof both basin peak I1 and nent the species indicative of such areas 2 populations but not within any of the other include erigeron peregrinus salix artica and populations on the other hand L breweribrewere wasas arnicaamica mollis fig 413 the bare ground vector only within the basin peak 1 populations in also increased toward the basin peak stands general it should be noted that classification although the concomitant increases in bare inferences based solely on location within the ground and vegetative cover may seem contra- 2 dimensional space defined by any 2 DCA dicdictorytory surface rock cover tended to decrease axes can be misleading for example mt price toward basin peak thus surface rock was 1 and mt price 2 are close together in the 2 replaced by vegetation and bare ground ie dimensional space defined by DCA axes I1 and inorganic soil along these vectors high litter 2 but they do not classify into the same group cover was commonly associated with the gran- in either TWINSPAN or NMDS because they ite chief and keithkeiths s dome populations are on different planes in the 3 dimensional plute pass is an area historically thought to space defined by the addition of a ard3rd axis contain L longipetalalongipetala however only L png individuals verified lewisia pygmaeapygmaean maea were at this site the area is south of yosemite national park cali- lewisia pygmaeapygmaean grew in areas where total fornia which makes it the southernmost site vegetative cover was greater than that where surveyed for L longipetalalongipetala environmental L longipetalalongipetala was found the strongest evi- attributes of plute pass are similar to those of dence for this difference in site characteristics other L longipetalalongipetala and L pygmaeapygmaean popula- was from the 5 sites where L longilongipetalapetala and tions except for some differences in species L pygmaeapygmaean coexisted in proximity to each other composition namely the relative preponder- basin peak 1 basin peak 2 granite chief ance of dodecatheon jeffrejeffreyyii dick s lake and keith s dome total vegetative cover for areas with L pygmaeapygmaean averaged discussion 60460.4 s 595.9sg which was 55 greater than the mean cover of 39039.0 safsyfs5fs 12312.3 for areas with L site characteristics that were most highly longipetalalongipetala this difference was significant at P associated with the occurrence of L longi 0100.10olo paired t test 4 df P 00670.067 this petala and also correlated with plant size and 199611996 distribution AND ECOLOGY OF L LONGIloncipetalalongipetalaPETALA 233

400 A litter cover vegetation cover 300 regroundbaregroundBa cover 200 z BP 121 2 0 piufepiume passposs 0 dicks lk 100 0 BP 111 1 0 0 0 granite chief keiths dome

CM 0 BP 1 3 0 0 basinbosin pk 2 V X 100 1 B lelo 0 300 0 migu nQ lubraubr 0 judr 200 0 audr anceoanmeo

andr lycspcaum miarmipr camecome 100 jume 0 000ooo 000goolepyjepy 0 0 kacomkapom dole chbrphbr 0 doal senoisenol siarsipr 0 0 erpearpe 0 0 A rmoamo casch 0 phaiphalphoi 100 isaarosaar 0 200 feryoperyo

I1 I1 I1 1 300 I I1 100 0 100 200 300 400 500 DCA axis I1

fig 4 A population ordinations generated by DCA for L pygmaeapygmaean B species groupings associated with the populations for both graphs circled groups were determined from TWINSPAN dendrograms letter codes for each species are given in table 2 except doie dodecatheonjeffreyidodecatheon jeffrejeffreyjjeffreyzyz 234 GREAT BASIN naturalist volume 56 density were proximity of snowbanks steep- the restriction of some species to sites with ness of slope slope aspect and cover of vege- low vegetative cover may be related to reduced tation surface rock and surface water these interspecific competition ostler et al 1982 inferences are supported by inspections of the for example competition partially accounts for site characteristics and by statistical analyses the reduced growth of talinum calcaricumcalcaricum a the L longipetalalongipetala populations with higher highly restricted rock outcrop species of the density were found on gently sloping sites with portulacaceae family in herbaceous sites domi- a northern exposure that were close to snow- nated by boapoa pratensispratensis ware 1991 viable banks and had low vegetative cover of all populations of the endangered furbish s louse- pole species for example creek I1 and 2 keiffkeith s wort pedicularis furbishiae occur on mesic dome and dick s lake have populations that rocky sites that experience intermediate distur- exceeded 500 individuals whereas basin peak bances from hydrological processes which which overall had the most herbaceous cover remove potential competitors menges 1990 of any of the other populations had much potentilla robbinsianarobhinsianarobbinsianaslana an endemic from new lower plant density plant density of L longi longiong hampshire s white mountains also requires petala populations increased with increased rocky mesic sites that are moderately dis- cover of surface water and rock but decreased turbed in this case by frost heaving that limits with total vegetative cover and slope steep- other species fitzgerald et al 1990 the lower ness plant size as measured by clump diame- densities and smaller L longipetalalongipetala plants in ter increased with decreased distance from areas with high vegetative cover and high soil snowbanks and decreased litter cover further- organic matter halford 1992 suggest that more at basin peak I1 as well as other sites of interspecific competition may also restrict this L longipetalalongipetala populations plants that were species but specific studies need to be con- more distant from snowbanks or that were on ducted to explicitly test this mechanism south facing slopes were more water stressed environmental halford 1992 the site characteristics of L pygmaeapygmaean are broader than those of L longi site characteristics that are associated with petala populations of L pygmaeapygmaean have been more vigorous L longipetalalongipetala populations are documented in dense herbaceous meadows indicative of areas that high snowsnowpackpack receive cracks in steep rocks and open gravely depres- accumulations in alpine plant environments sions elliot 1966 major and taylor 1977 in communities whose occurrences are influ- our study plants of L pygmaeapygmaean were found enced by geomorphological characteristics that adjacent to 3 and interspersed with 2 of the 12 favor high snowpackpack snow accumulations are often L longipetalalongipetala populations which suggests that termed snow bed vegetation billings and bliss L pygmaeapygmaean and L longipetalalongipetala can grow in sim- 1959 kuramoto and bliss 1970 canaday and ilar environments however an important dif- fonda 1974 tomaselli 1991 some species in ference between the 2 species is that L bygpyg the sierra nevada that major and taylor 1977 maea was found in areas with more herbaceous commonly found associated with areas of high cover the less pronounced site specificity ex- snowsnowpackpack accumulations and that often occur hibited by L pygmaeapygmaean parallels other widely in mesic depressions with low vegetative cover distributed mesic alpine species whereas the are phyllodoce brebreweribrewereweri cassiope mertensianamertensiana relative restriction of L longipetalalongipetala to more kalmia polifoliapolifolia var micromicrophyllaphylla phleum open sites is similar to other restricted plant alpinumalpinum mimulus primulprimuloidesoides and M gutta taxa fitzgerald et al 1990 menges 1990 tus additional species that occur in mesic to the potential threats to L longipetalalongipetala are even hydric habitats include antennaria media not imminent at this time but include both sto- sibbaldiaSibbaldia procumbensprocumbent dodecatheon alpialpinumnum chastic and anthropogenic processes climatic and sedum roseum major and taylor 1977 events such as periodic droughts that reduce these species were also associated with L longi snowsnowpackpack accumulations as well as potential in- petala populations conversely species that are creases in interspecific competition may signif- more frequently associated with xeric sites icantly reduce the viability of L longipetalalongipetala pop- such as lupinus breweribrewere and juncus drum ulatulationsions especially those that already have low mondiimondai chabot and billings 1971 nachlinger densities of individuals human activities may 1985 were less frequently associated with L also have significant impacts for example if longipetalalongipetala populations slopes above populations are altered by mining 199611996 distribution AND ECOLOGY OF L longipetalaLONGI PETALA 235

activity or ski area development the displace- HICKMAN J C EDITOR 1993 the jepson manual higher ment of substrate could alter the topography plants of california university of california press berkeley 1400 appp and hence hydrology of the site through changes HILL M 0 1979a TWINSPAN a FORTRAN program in snow accumulation and melt water runoff for arranging multivariate data in an ordered two to enhance the longtermlong term viability of this way table by classification of the individuals and endemic species primary management goals attributes cornell university ithaca NY 1979b DECORANA a FORTRAN program for 1 should include monitoring of L longipetalalongipetala detrendedDetr ended correspondence analysis ordination populations to gauge how changes in site cornell university ithaca NY hydrology may influence fluctuations in plant HUTCHINGS M J 1991 monitoring plant populations density and 2 acquisition by land conservation census as an aid to conservation pages 62 86 inm FE B goldsmith editor monitoring for conservation and groups of sites that may be impacted due to ecology chapman and hall new york london mining or ski area development KOMRKOVKOMARKOVA V 1975 alpine vegetation of the indian peaks area front range colorado rocky mountains acknowledgments unpublished doctoral dissertation university of colorado boulder KURAMOTO R T AND L C bussBLISS 1970 ecology ofsubof sub this study was partially funded by a grant alpine meadows in the olympic mountains wash- from the hardman foundation as well as cost ington ecological monographs 40 317 347 share funds from the california native plant MAJOR J 1989 endemism a botanical perspective pages 117 A myers S giller society and the tahoe tahoe basin and 146 in A and P editors anal- lake ytical biogeography chapman and hall london eldorado national forests additional support MAJOR J AND D W TAYLOR 1977 alpine in M G bar- was provided by the nevada agricultural bour and J major editors terrestrial vegetation of experiment station the excellent field assis- california john wiley and sons new york MATHEW press tance of kirk halford is gratefully acknowl- B 1989 the genus lewisia timber inc portland OR as is edged the drawing of lewisia longipetalalongipetala MENGES E S 1990 population viability analysis for an by annaliese miller endangered plant conservation biology 4 52 62 MINCHEN P R 1987 an evaluation of the relative robust- CITED ness of techniques for ecological ordination vegetatiovegetation literature 69 89 107 MUNZ P A 1959 A california flora and supplement uni- BASKIN J M AND C C BASKIN 1988 some considera- versity of california press tions in evaluating and monitoring populations of rare nachlinger J 1985 the ecology of subalpine meadows plants in successional environments natural areas in the lake tahoe region california and nevada journaljournals62626 30 unpublished master s thesis university of nevada BILLINGS W D AND L C BLISS 1959 an alpine snow- reno bank environment and its effects on vegetation planpianplantpiantt NELSON D AND K HARPER 1991 site characteristics and development and productivity ecology 397 40 388 habitat requirements of the endangered dwarf bear CANADAY B B AND FONDA 1974 R W the influence of claw poppy arctomecon humilis coville papaver- subalpine snowbanks on produc- vegetation pattern aceae great basin naturalist 51 167 175 tivity and phenology bulletin of the torrey botanical OSTLER K W K T HARPER K B mcknightMCKNIGIIT AND D C club 101 340 350 ANDERSON 1982 the effects of increasing snowsnowpackpack CHABOT B FE AND W D BILLINGS 1971 origins and on a subalpinesub alpine meadow in the uinta mountains utah ecology of the sierran alpine flora and vegetation USA arctic and alpine research 14 203 214 ecological monographs 42 163 197 SKINNER M W AND B M PAVLIK EDITORS 1994 inven- DARGIE T C D 1984 on the integrated interpretation of tory of rare and endangered vascular plants of cali indirect ordinations site a case study using semisemiaridarid fornia california native plant society no 1 ath5th edi- vegetation in southeastern spainspam vegetatiovegetation 55 37 55 tion 336 appp DEMPSTER L M 1993 in J C hickman editor the jep- STEBBINS G L 1968 A lost species rediscovered news- son manual higher plants of california university of letter of the california native plant society 4 1 5 press california berkeley and los angeles STEBBINS G L AND J MAJOR 1965 endemism and spe- ELLIOT R C 1966 the genus lewisia bulletin of the ciation in the california flora ecological monographs alpine garden society 34 1 76 35135 1 35 fitzgerald B T K D KIMBALL AND C V COGBILL SUTTER R D 1986 monitoring rare plant species and nat- 1990 the biology and management of potentilla rob ural areas ensuring the protection of our invest- binsianabinsiana an endemic from new hampshire s white ment natural areas journal 6 3 5 pages mountains 163 166 in ecosystem manage- TOMASELLI M 1991 the snow bed vegetation in the ment rare species and significant habitats new york northern apennines vegetatiovegetation 94 177 189 museum bulletin 471 TUELLER P T R J TAUSCH AND V BOSTICK 1991 species HALFORD A S 1992 the ecology and distribution ofa cali- and plant community distribution in a mojave great fornia high altitude endemic plant lewisalewisialewisw longilongipetalapetala basin desert transition vegetatiovegetation 92 133 150 unpublished master s thesis university of nevada UTTER J M AND A W HURST 1990 the significance and reno management of relict populations of chamaehnumchamaelirium 236 GREAT BASIN naturalist volume 56

luttumluteum L gray pages 180 184 in ecosystem man- ives editors ecological impacts of snowsnowpackpack aug- agement lareiarerare species and significant habitats new mentation in the san juan mountains colorado final york museum bulletin 471 report of the san juan ecology project to the USU S wablwabeWAREWARL S 1991 influence of interspecific competition light bureau of reclamation division of atmospheric water and moisture levels on growth of rock outcrop tal- resources management colorado state university inum portulacaceae bulletin of the torrey botanical publication CSU FNR 7052 1 club 1181118 1 5 WEBBER P J J C EMERICK D C MAY AND V KOMARKOVA received 29 august 1995 1976 the impact of increased snowfall on alpine accepted 8 april 1996 vegetation pages 201 264 in H W stemhoffandjsteinhoff and J D great basin naturalist 563 0 1996 appp 237 246

LARGER ectoparasites OF THE IDAHO GROUND SQUIRREL spermophilus BRUNNEUS

eric yensenlyensen1 craig R bairdibaird2 and paul W Shermanshermanasherman33

ABSTRACT we sampled both subspecies of the idaho ground squirrel spermophilus brunbrunneusbrunneousneus to document the larger ectoparasites of this rare endemic S b brunbrunneusbrunneousneus was host new host record new idaho record to 4 flea species neopsyllaNeopsylla inopina oropsyllaOropsylla idahoensis 0 tuberculatatuberculatetuberculata and tarassisthrassisThrassis panpandoraedorae I1 tick ixodes sculptussculptus and an eyeworteyeworm nematoda rhabditis orborbitaorbitalsorbitalisorbitahsitalishs also ist records from sciuridae S b endemicusendemicus was host to a louse species neohaematopmusneohaematopinus laeviusculus 5 flea taxa rhadinopsyllarhadmopsylla sp 0 t tuberculatetuberculatatuberculata tarassisthrassisThrassis f francisi T f barnesibargnesi and T f rockwoodirockwoodtrockwotockrockwoodiodt and a mite androlaelapsAndrolaelaps fahrenholzi spermophilus brunbrunneusbrunneousneus had fewer known ectoparasite species than other convenerscongenerscong eners although all of their parasites had many other hosts S b endemicusendemicus and S b brunbrunneusbrunneousneus shared only a single parasite species in common whereas all but one of their ectoparasites also occurred on the closely related townsend s ground squirrel S townsendii the proportion of parasitized individuals and the parasite loads per individual were significantly lower in S b brunneusbrunbrunneousneus which lives in small isolated populations than in S b endemicusendemicus which has larger less fragmented populations suggesting a relationship between host population structure parasite loads and parasite species diversity all but one of the flea species have been linked to plague transmission

key words ground squirrels ectoparasites spermophilus brunneusbrunbrunneousneus idaho

the idaho ground squirrel spermophilus and yensen 1990 eliminating habitat the brunbrunneusbrunneousneus is one of the rarest and until recently remaining populations are presently isolated least known north american mammals sher- from each other by the encroachment of coni- man 1989 yensen 1991 yensen and sherman fers into meadows and by competition with in press this endemic species inhabits a 125 columbian ground squirrels yensen and sher- X 90 km area in west central idaho but it man in press today there is apparently little actually occupies only a small fraction of this or no gene flow among populations allozymeAllozyme limited range yensen 1991 despite the analyses of 55 protein loci in 12 populations species restricted geographic distribution gavin et al submitted indicated that the pro- there are 2 allopatric subspecies that are mor- portion of polymorphic loci was 11511.5 19219.2 phophologicallylogically and genetically differentiated and and heterozygosity values were 00410.041 00800.080 possibly have reached species level separation ftF was 03170.317 implying that there is genetic yensen 1991 gill and yensen 1992 gavin et differentiation among populations despite their al submitted geographic proximity and the apparent recency Spennspermophilusophilus b brunbrunneusbrunneousneus occurs in montane of their separation in 1993 the total number of meadows surrounded by coniferous forests at individual S b brunneusbrunbrunneousneus was 1000 1200 but elevations of 1035 to 1550 in in adams and val- the number fell to 600 800 in 1994 and 1995 ley counties yensen 1991 As of 1995 only 18 T A gavin PE W sherman E yensen per- of the 28 known populations remained and sonal observation only one of these contained 100 animals the spermophilusSpennophilus b endemicusendemicus occurs in rolling majority of the sites were within an area of 22 foothills at elevations of 670 to 975 m in gem X 9 km and totaled 300 ha of occupied habi- payette and washington counties yensen 1991 tat T A gavin PE W sherman and E yensen it is patchily distributed throughout its range unpublished data of 75 X 30 km although censuses of S b fire supression began in the area about 100 endemicusendemicus populations have not been made its yr ago subsequent succession and expansion total population is apparently much larger than of forests has filled in many of the natural that of S b brunbrunneusbrunneousneus the area occupied esti- meadows in the range of S b brunbrunneusbrunneousneus truksa mates of population densities and the amount

imuseurnimuseumIMus eurn of natural history albertsonaibeAlbe rhon college caldwell ID 83605 university2university of idaho parma research and extension center parma ID 83660 section of neurobiology and behavior cornell university ithaca NY 14853

237 238 GREAT BASIN naturalist volume 56

of remaining habitat are more than 2 orders mortem squirrels were placed individually in of magnitude greater than for S b brunbrunneusbrunneousneus plastic bags fleas ticks lice and larger mites yensen E personal observation were collected with forceps or a camel s hair parasites of S brunbrunneusbrunneousneus have not been pre- brush moistened with 70 ethanol as they left viously surveyed the only prior records baird the host squirrels were not examined under a and saunders 1992 were 2 flea species oro dissecting microscope so smaller mites were psylla t tuberculatatuberculatetuberculata and tarassisthrassisThthrassisfrancisirassis francisi rock not collected eyes were not examined for eye woodi collected from specimens now referred worms to S b endemicusendemicus yensen 1991 from 1987 to 1994 S b brunbrunneusbrunneousneus were we were interested in how ectoparasite live trapped for demographic and behavioral diversity and density are affected by reduction studies sherman 1989 and ongoing they in size and isolation of host populations were hand held and parasites were picked off according to epidemiological models ander- with forceps because the animals were not son and may 1979 may and anderson 1979 anesthetized all of the smaller and some of the the number of contacts between hosts and inin- larger ectoparasites may not have been seen fective stages of parasites determines the rate eyes were checked for eyewormseyeworms by pulling at which adult parasites are acquired mean back the upper lid specimens were removed parasite load should equal growth rate of the from the cornea of the eye with a cotton swab population divided by mortality from the dis- moistened with sterile water all parasites were ease thus as population growth slows para- placed in 70 ethanol in addition 21 S b site load per individual should drop at very endemicusendemicus were live trapped at sand hollow low host population densities there may be too payette county idaho in 1994 and examined few contacts even to maintain ectoparasite popu- for eyewormseyeworms lations thus we predicted that S b brunbrunneusbrunneousneus collected specimens of S brunbrunneusbrunneousneus were prepared as should have fewer ectoparasite species and standard museum study skins and skulls fewer ectoparasites per individual than con- and deposited in the albertson college museum generic more widely distributed western of natural history ACMNH caldwell ground squirrels spermophilus sppapp we also idaho and the national museum of natural predicted that due to its fragmented popula- history USNM they are identified below by museum number specimens of ectopara tion structure and smaller population sizes S b sites were sent to appropriate specialists for brunbrunneusbrunneousneus should have fewer ectoparasite species identification and deposited the entomologi- than S b endemicusendemicus in cal collections at the of because of questions about the taxonomic university idaho moscow and ACMNH differences in similarity of S b brunbrunneusbrunneousneus and S b eidemiendemi parasite loads between individuals and subspecies cus we also wished to learn if they had similar were analyzed with hand calculated mann ectoparasites and how similar their ectopara whitney 17 tests and chi square tests as sites were to those of other western ground appropriate squirrels further because of the limited geo- RESULTS graphic range and low number of small populations both subspecies of S brunbrunneusbrunneousneus would we examined 29 freshly collected individu- be vulnerable to extirpation by an epizootic als of S b brunbrunneusbrunneousneus and 53 of S b endemicusendemicus such as plague thus it was important to learn for ectoparasites these represent 43 of the if their ectoparasites were species involved in 192 museum specimens of this species known plague transmission to us yensen 1991 plus 4 additional specimens additionally we opportunistically collected ecto- METHODS parasitic arthropods from 12 live trapped individuals of S b brunbrunneusbrunneousneus and eyewormseyeworms from from 1980 to 1990 specimens of S brunbrunneusbrunneousneus another 36 we examined 21 S b endemicusendemicus were collected for a taxonomic study yensen for eyewormseyeworms 1991 to minimize negative impacts on small we collected 6 ectoparasite species from populations a mean of 050.50 5 mdividualssiteyrindividualssiteyr of spermophilus b brunbrunneusbrunneousneus 4 fleas 1 tick and 1 S brunbrunneusbrunneousneus was collected squirrels were killed nematode table 1 we collected 7 taxa of by shooting or by live trapping and injecting ectoparasites from S b endemicusendemicus 5 fleas 1 nembutalnembutol into the heart immediately post louse and 1 mite 199619961 spermophilus BRUNNEUS ectoparasites 239

TABLE 1 parasites of S brunbrunneusbrunneousneus that also occur on some other species ofwesternof western ground squirrels subgenus sper- momophilusphilus symbols known primary hosts records possibly accidental on host no records in references below host this study literature recordsrecords1recordsb1 parasite sbbabb sbeabe sto seesecsce sblabl sar sellseilselc sri swa spy LICE neohaematopinus laeviusculus FLEAS neopsyllaNeopsylla inopinainopine oropsyllaOropsylla idahoensis 0 t tuberculatatuberculatetuberculata rhadinopsylla s secsectilissectiussextilistilistills thrassisf barnesibargnesi TftffrancisiT f francisi tfT f rockrockwoodiwoodi T p panpandoraedorae 4 TICKS ixodes sculptussculptus MITES androlaelapsfahrenholziandrolaelapsAndrolaelaps fahrenholz i NEMATODA rhabditis orbitalisorbitalis

from records inin hubbard 1947 burgess 1955 stark 1970 hilton and mahrt 1971 whitaker and wilson 1974 holekamp 1983 lewis et al 1988 baird and saunders 1992 baird unpublished and this study blostbhosthost acronyms sbbabb spermophilus b brunbrunneusbrunneousneus sbeabe S b endeendemicusmicus stesto S townsendtownsendiitownsendtttownsendiaiitt gensusensu latu secseesce S columbianuscolumbianus sblabl S belbeibeldingibeldtngzdingi sar S armatusarmatos sel S elelegansdelegansegans sri S richardsoncharclsonnrichardsoniirichardsonianiinil swa S washingtoniwashzngtomwashingtondwashing toni spy S parryii confused inin the literature with S richardsorichardsoniinchardsonurichardsonianiinil the records here are those that unambiguously refer to this species and the total for S hchardsoniinchardsonn may include a few parasites of this species

the proportion of parasitized individuals in anopleuraAnopleura haematopinidae the 2 subspecies was strikingly different we neohaematopinusneohaematopintis laeviusculus grube found ectoparasites on 37 of 53 70 S b we found this louse on S b endemicusendemicus in endeendemicusmicus but on only 8 of 29 28 S b brun the following locations 11 mi 18 km N neus collected xax2 13413.4 d 1 FP 00010.001 di emmett gem co tan r2wraw sec 13 parasitized individuals of S b brunbrunneusbrunneousneus had t8n 4402n 1163rw11631w1163rW 830 in elev 21 february 1 3 175 species of ectoparasites each X 1751.75 n 1982 ACMNH 222 28 february 1982 8 and parasitized individuals of S b ACMNH 226 227 236 237 238 010.1oi mi E endemicusendemicus had 1 4 species of ectoparasites X payette co line 12612.6 mi 20 km N emmett 1591.59 n 37 this difference was not signif- gem co t8ntan r2wraw sec 12 4403n icant ug 154 P 0.505os us 05 however there was 11632w 810 in elev 28 february 1982 a significant difference between subspecies in ACMNH 224 weiser cove washington co the parasite load of parasitized individuals 4413n 11644w 715 in elev 7 march fleas were the only common group of ectopar 1982 ACMNH 228 229 230 lower mann asitesarites ofbothof both ground squirrel taxa there were creek 252.5 mi 4 km N jet weiser river road 414.1 fleas per parasitized individual in S b washington co 4416n 11651w 720 in brunbrunneusbrunneousneus and 787.8 in S b endemicusendemicus Usug elev 14 march 1982 ACMNH 231 240 242 95595.5 P 0050.05oos 243244243 244 this louse occurs from eurasia east to alaska ANNOTATED LIST OF ectoparasites and the northwest territories and south through western united states to mexico it is in the ectoparasite species accounts below apparently a species complex K C emerson letters and numbers in brackets refer to the personal communication lice of this complex number of male and female fleas eg 1 I1 m 2 flf have been collected from many ground squir- or to conversions of original collecting data to rels eurasian spermophilus major S citellus latitude longitude and metric units S pygmaeus S undulatusundulates and north american 240 GREAT BASIN naturalist volume 56

S beecheyibeecheyi S matusareatusararmatus S belbeldingidingi S colum siphonaptera ceratophyllidae S S townsendiitownsendia S washing blanus parryiiparryii townsendii oropsyllaOropsylla idahoensis baker toni and Ammoammospennophilusammospermophilusspermophilus leucurusleucurus as well this flea species was collected on S b marmota chipmunks as marmottmarmots mannotaflaviventrisfiauiflaviflaviventrisventris brunbrunneusbrunneousneus at the following locations price val- pocket perognathus tamias minimus mice ley 11626011626w 1270 m elev 3 june and deer peromyscus manicula 4501n parousporous mice canicula 1981 ACMNH 209 3 flf and OX ranch 1 2 tus rayburn et al 1975 shaw and hood 1975 km S 1 2 km E bear adams co 4500n records from national museum of natural 45wn his- ilg1161163939w 1340 m elev live trapping collec- tory although N laeviusculus is the most com- tions mon louse species taken from ground squirrels oropsyllaOropsylla idahoensis occurs from alaska to in idaho C R baird personal communication new mexico and is one of the most common K C emerson personal communication S b fleas of ground squirrels in the rocky moun- endemicusendemicus is a new host record tains and westward hosts include other west- siphonaptera hystrichopsyllidae ern ground squirrels of subgenus Spennspennophilusophilus table 1 golden mantled ground squirrels S neopsyllaNeopsylla inopinainopine rothschild lateralis and marmottmarmots Manmarmotanota sp lewis et we collected 8 individuals of this flea from al 1988 baird and saunders 1992 S b brun S b brunbrunneusbrunneousneus in the following locations lick neus is a new host record creek adams co t19n r3wraw sec 14 4459n 11640w 1290 ra elev 17 april Orooropsyllapsylla tuberculatetuberculatatuberculata tuberculatetuberculatatuberculata baker 1983 ACMNH 305 1 m 2 f ACMNH 306 this was the most common flea on both 1 I1 flf I1 mi iglg161.6 km NE bear guard station S b brunbrunneusbrunneousneus and S b endemicusendemicus occurring adams co 4505n 11637w 1480 m at nearly all locations from which we collected 2 june 1988 ACMNH 518 1 Qf and price ectoparasites we found 0 t tuberculatetuberculatatuberculata on valley 45orn4501n45 olnOrN 11626w 1270 m elev S b brunbrunneusbrunneousneus at the following localities price 3 june 1981 ACMNH 209 1 m I1 ffl ACMNH valley 45orn4501n45 olnOrN 11626w 1270 m elev 210 if1 fl 3 june 1981 ACMNH 209 1 m ACMNH this flea occurs from british columbia 210 1 m mill creek summit 5 km N hornet south to oregon and nevada and eastcast to guard station adams co t18n r3wraw sec saskatchewan north dakota and utah lewslewis 25 4500 elev 4453n 11639w 1370 m 2 et al 1988 it has been collected from other june 1985 ACMNH 510 2 m 3 f ACMNH western ground squirrels of subgenus sper- 512 2 m 3 flf lick creek adams co t19n momophilusphilus table 1 and from badger taxidea r3wraw sec 14 4454n 11640w 1290 m taxus dens lewis et al 1988 baird and saun- elev 17 april 1983 ACMNH 305 4 m 3 f ders 1992 S b brunbrunneusbrunneousneus is a new host record ACMNH 306 1 flf round valley valley co 4421n 11600w 1460 m elev 18 may rhadinopsylla sp 1985 ACMNH 315 1 flf we collected 1 female specimen of this flea records from S b endemicusendemicus are as follows genus from S b endemicusendemicus unfortunately it sucker cr 11 mi 18 km N emmett gem co could not be identified to species the locality t8ntan r2wraw sec 13 4402n 11631w 830 was dry creek road payette co 141.4 mi 22222.2 m elev 21 february 1982 ACMNH 221 222 km E little willow creek t9ntan r2wraw sec 18 223 28 february 1982 ACMNH 225 226 440744 07n 11637w 815 m elev 26 february 2273227 3 may 1987 ACMNH 544 1 mml 01OI0.1oi mi 1983 ACMNH 318 1 flf reported in baird and E payette co line 12612.6 mi 20 km N emmett saunders 1992 gem co t8ntan r2wraw sec 12 4403n the flea is most likely R s secsectilissextilistilis which 11632w 810 m 28 february 1982 ACMNH occurs in many western states on deer mice 224 236 237 238 reported in baird and peromyscus sp and ground squirrels includ- saunders 1992 dry creek road 141.4 mi 22222.2 ing S townsendtownsendiitownsendiaii and S washingtoniwashingwashingtondtoni lewis et km E little willow creek payette co t4ntan al 1988 baird and saunders 1992 rhadino r2wraw sec 18 4407n 11637w 815 rn psylla are uncommon fleas and have popula- elev 20 february 1983 ACMNH 318 10 m tion peaks in the colder months lewis et al 13 f 26 february 1983 ACMNH 317 8 m 1988 this is the ist record of any rhadino 3 Qf weiser cove washington co 4413n4413 psylla species from S brunbrunneusbrunneousneus 11644w 715 m elev 7 march 1982 199619961 spermophilus BRUNNEUS ectoparasites 241

ACMNH 228 229 230 lower mann creek common hosts are S armalusannatusarmatus and S eleganseldelegansegans 252.52 5 mi 4 km N jet weiser river road wash- rather than S townsendtownsendiitownsendiaii mollis the usual host ington co 4413n 11651w 720 m elev of T f francisi stark 1970 felt that host asso- 14 march 1982 ACMNH 231 232 233 240 ciationsciations may separate the 2 subspecies of T 242 243 244 washington co lower mann francisi although the 2 fleas appeared to inter- creek 33 mi 53 km N jet weiser river grade in eastern nevada S b endemicusendemicus is a road 4417n 11651w 730 m elevclev 14 new host record march 1982 ACMNH 239 this is a very common flea in most of the thrassistarassisThthrassisfrancisifrancisirassis francisi francisi fox western united states and western canadian we collected 14 individuals of this flea from provinces baird and saunders 1992 hosts S b endeendemicusmicus at I1 locality dry creek road include other western ground squirrels of sub- 141.4 mi 22222.2 km E little willow creek payette genus Spennspermophilusophilus table 1 antelope ground co t4ntan r2wraw sec 18 4407n 11637w squirrels ammospennophilus leucurusleucurus wood 815 m elev 26 february 1983 ACMNHlswelsw318 rats neotoma sp and badgers lewis et al 1 m 5 f sm2sma 2 m 3 ffl 24 february 1986 1988 baird and saunders 1992 it was previ- ACMNH 920 2 m I1 flfj ously recorded from S brunbrunneusbrunneousneus by baird and this flea is known from the great basin saunders 1992 desert of eastern oregon idaho south of the snake river eastern nevada utah and parts tarassisthrassisThrassis panpandoraedorae panpandoraedorae jellison of wyoming it occurs primarily on S town we found I1 specimen of this flea on S b sendil but the white tailed prairie dog onocyno- brunbrunneusbrunneousneus at lick creek adams co t19n mys leucurusleucurus is the usual host in wyoming r3wraw sec 14 4454n 11640w 1290 m stark 1970 there are incidental records from elev 17 april 1983 ACMNH 305 1 m several species of ground squirrels table 1 this flea is distributed from washington to marmarmotsmarmottmots and deer mice stark 1970 our california and east to colorado stark 1970 it records are the ist from any host north of the is found most frequently on spermophilusSpennophilus ahnaannaarma snake river in idaho stark 1970 lewis et al tus S belbeldingidingi and S elegansdelegans nchardsonuiichardsonii in 1988 baird and saunders 1992 S b eidemiendemi stark 1970 but also occurs on S columbianuscolumbianus cus is a new host record S elegansdelegans table 1 and a variety of other rodents lagomorphslagomorpha and carnivorescarnivores stark thrassistarassisThthrassisfrancisirassis francisi rockrockwoodiwoodi hubbard 1970 S b brunneusbrunbrunneousneus is a new host record two males of this flea were collected from S b endeendemicusmicus at a single locality sucker tarassisthrassisThthrassisfrancisirassis francisi barnesibargnesi stark creek 11 mi 18 km N emmett gem co we found this flea on S b endeendemicusmicus at t8ntan r2wraw sec 13 4402n 11631w 830 sucker cr 11 mi 18 km N emmett gem co m elev 21 february 1982 ACMNH 223 28 t8ntan r2wraw sec 13 4402n 11631w 830 february 1982 ACMNH 227 2 m reported m elev 31 may 1981 ACMNH 220 3 m in baird and saunders 1992 4 ffl 3 may 1987 ACMNH 540 4 m af3f this subspecies has been recorded from east- ACMNH 541 2 m I1 ffl ACMNH 542 1 m 6 f ern oregon northwestern nevada and north- ACMNH 543 4 m 7 f ACMNH 544 1 m ern california where it occurs most commonly I1 f ACMNH 545 2 m I1 f ACMNH 547 on S belbeldingidingi although collections have been 4 m 9 ffl ACMNH 548 1 ffl ACMNH 549 made from S townsendtownsendiitownsendiaii stark 1970 lewis 3 m 7 flf 7 mi 11 km N emmett gem co et al 1988 t7ntan RIW sec 5 4358n 11629w 920 m elev 23 may 1987 ACMNH 546 4 m 2 Qf acarina ixodidae sand hollow 565.65sg 6 km N 505.05so 0 km E payette hodesbodesixodes sculptussculptus neumann payette co t9ntan r4wraw sec 7 44wn4408n we collected specimens of this tick from S b 1165rw11651w1165rW 750 m elev 30 march 1989 USNM brunbrunneusbrunneousneus at I1 locality OX ranch 1 2 km S 565927 3 m 2 Qf 1 2 km E bear adams co 4500n this flea occurs north of the snake river in 1340 m elev live trapping collectionslw western idaho and on both sides of the river this widespread tick occurs from western in eastern idaho and south into central utah canada south to california and texas and east and eastern nevada stark 1970 its most across the great plains it occurs on several 242 GREAT BASIN naturalist volume 56 western ground squirrels of the subgenus sppapp mice apodemusApodemus sppapp and mus muscu- spermophilus table 1 prairie dogs cynomys lus and rats rattus norvegicus poinar 1965 sp marmottmarmotsmarmots voles microtus sp pikas kinsella 1967 cliff et al 1978 hominick and ochotona sp gophers thomomys sp jump- aston 1981 schulte 1989 S b brunbrunneusbrunneousneus is a ing mice zapus sp domestic animals and new host record the ist record of any rhabdi- various carnivoresearncarnivores doss et al 1974 S b tis from sciuridae and also the ist record of R brunbrunneusbrunneousneus is a new host record orbitalisorbitalis from idaho acarina Laelapidae epizootics androlaelapsfahrenholziandrolaelapsAndrolaelaps fahrenholzfahrenholzii berlese in 11 field seasons april june of work we collected 8 specimens of this mite from with S b brunbrunneusbrunneousneus we found only 2 dead indi- S b endemicusendemicus at the following localities sucker vividuals and none were observed sick or dying cr 11 mi 18 km N emmett gem co t8ntan while a number of populations have declined r2wraw sec 13 4402n 11631w 830 in T A gavin PE W sherman and E yensen per- elev 21 february 1982 ACMNH 227 4 f 2 sonal observation mortality occurred while the deutodeutonymphsnymphs lower mann creek 2522.55 mi 4 animals were in hibernation rather than during km N jet weiselweiserwelser river road washington co the active season the most serious population 4416n 11651w 720 in elev 14 march declines were estimated to be around 50 in 1982 ACMNH 233 2 ffl I1 yr rather than the 95 100 active season this mite is widespread inm eurasia north mortality typically associated with plague america whitaker 1979 and central america lechleitner et al 1968 rayor 1985 although strandtmann 1949 it occurs on a wide vari-varivarl numbers of fleas on individual squirrels were ety of mammals including marsupials didel- relatively low especially in S b brunbrunneusbrunneousneus all phis sp insectmsectivoresinsectivoresivores bats several families of flea species we collected are important in rodents lagomorphslagomorpha carnivoresearncarnivores and birds plague epidemiology in other hosts pratt and strandtmann 1949 whitaker and wilson 1974 stark 1973 and could potentially play a role in rayburn et al 1975 Oposopossumssums insectivoresmsectivoresinsectivores an idaho epizootic and rodents are the primary hosts but A fahrenholzi has the least host specificity and discussion widest geographic range of any north american ectoparasitic mite whitaker 1979 these collections of ectoparasites from S brun are the ist records from S brunbrunneusbrunneousneus neus have resulted in new state records for the flea tarassisthrassisThrassis rockrockwoodiwoodi and the eye nematoda francisi rhabditidae worm rhabditis orbitalisorbitalis plus 9 new host rhabditis Pelodera orborbitaorbitalsorbitahsorbitalisitalishs sudhaus and records because there have been no previous schulte studies of S brunbrunneusbrunneousneus the new records are we observed this parasitic eyeworteyeworm only in hardly surprising however the records of live trapped S b brunbrunneusbrunneousneus from OX ranch 1 2 thrassistarassisThrassis ff francisi and T ff rockrockwoodiwoodi on S b kmkni S 1 2 km E bear adams co 4500n endemicusendemicus were unexpected tarassisthrassisThrassis f bar IIGWW 1340m1340 in elev nesi occurs north of the snake river in the all specimens were collected inm april and snake river plain stark 1970 and is the sub- may 1990 to 1994 we found them in I1 eye 01or species of tarassisthrassisThrassis francisi that would be ex- both eyes of yearling and adult S b brunbrunneusbrunneousneus pected to occur in the range of S b endemicusendemicus the number per eye vanedvariedvalled from 0 to 1272 instead we found tarassisthrassisThthrassisfrassis f francisi which is the museum specimens were not checked for common in S townsendtownsendiitownsendiaii mollis south of the eyewormscyewormseyecyeworms in 1994 T A gavin and P W sher- snake river and T f rockrockwoodiwoodi for which the man examined 21 live trapped S b endeendemicusmicus nearest locality is from oregon across the snake from sand hollow payette co and found no river stark 1970 a major biogeographic bar- eyewormseyeeyewoiwormsms rier in southern idaho davis 1939 this inter- this eyeworteyeworm has been reported previously esting situation merits further study from eurasian and north american voles and with the exception of eyewormseyeworms ectopara lemmings microtus sppapp lemmus trimucronatnmucrona sites of S brunbrunneusbrunneousneus are all known from multi- tus dicrostonyx groenlandgroenlandicusicus pitimys subterhubter ple other species of ground squirrels table 1 raneus arvicola terrestristerrestnstetreterrestris and clethrionomys thus it is curious that S b brunbrunneusbrunneousneus and S b 199619961 spermophilus BRUNNEUS ectoparasites 243 endemicusendemicus shared only a single ectoparasite ancesences in hair density and diameter although oropsyllaOropsylla t tubertuberculatatuberculateculata a widespread flea found these were not quantified by yensen 1991 on at least 4 other species of ground squirrels possibly S townsendtownsendiitownsendiaii is inhabitable by the by contrast the geographically and taxonomi- entire set of ectoparasites and each subspecies cally close nadler et al 1984 S townsendtownsendiitownsendiaii has of S brunbrunneusbrunneousneus is a suitable host for about half all but one of the ectoparasite species found on the set thus pelage differences hypothesis 3 both S brunneusbrunbrunneousneus subspecies however sper- are a possible explanation for the lack of over- momophilusphilus townsendtownsendiitownsendiaii is now recognized hoff- lap in ectoparasite species between 2 very mann et al 1993 as a complex of 3 closely close relatives but it would not explain the dif- related sibling species with different kary ferencesferences in parasite loads or the low percent- stypesotypes and it was not always clear to us from ages of nonparasitized individuals the literature table 1 which parasites were anderson and may 1979 argued that para- associated with which host consequently we site infestations should be sensitive to host have treated S townsendii as a single entity population structure hypothesis 4 As popula- herein tion size declines and populations become more there are several possible explanations for isolated the probability of parasite species loss the lack of shared ectoparasites between S b should increase our data were consistent with brunbrunneusbrunneousneus and S b endemicusendemicus 1 they are geo- this pattern the proportion of parasitized S b graphically separated and their ranges are brunbrunneusbrunneousneus was significantly lower than that of inhabited by different ectoparasites 2 they S b endeendemicusmicus the former has smaller more occur in different habitats and therefore have isolated populations different ectoparasites 3 pelage differences the isolated S b brunbrunneusbrunneousneus populations between them may be different micromicrohabitatshabitats would also retard exchange of ectoparasites for ectoparasites 4 possibly the formerly among populations thus there might be sto- shared ectoparasites on one or the other sub- chastic losses of parasite populations with low species have been lost via a founder event due probability of recolonization anderson and may to population structure or because of popula- 1979 the differences in incidence of parasites tion bottlenecks and 5 we did not adequately between S b brunbrunneusbrunneousneus and S b endemicusendemicus sample all ectoparasites on either subspecies are consistent with this interpretation among these hypotheses 5 is the least inter- the low density and wide dispersion of in- esting evolutionarilyrevolutionarily and 4 is the most inter- dividualsdividuals within S b brunneusbrunbrunneousneus populations at a esting site E yensen and PE W sherman personal most western ground squirrel species are observation may also retard direct transfer of allopatric or parapatricparapatric thus there is little pos- ectoparasites and consequently S b brunbrunneusbrunneousneus sibisibilitylity of direct transmission of ectoparasites populations may not be able to support large among them historically the 2 subspecies of ectoparasite populations the low incidence of S brunbrunneusbrunneousneus were separated by 19 km 250 m in parasitism in idaho ground squirrels thus elevation and a habitat change from andaridarld shrub appears to be related to population structure steppe vegetation to montane meadows yensen because we did not examine ground squir- 1991 at present the nearest extantextant popula- rels under a microscope we do not suppose tions are separated by 48 km because S town that all ectoparasites were collected hypothe- sendiisendia is allopatric to S brunbrunneusbrunneousneus occurs in non sis 5 however there was no systematic bias montane habitats and has all ectoparasites found in the sampling that would account for the dif- on both subspecies of S brunbrunneusbrunneousneus differences ferencesferen ces in the proportion of parasitized ani- in geography hypothesis 1 and habitats 2 are mals and parasite load differences between S b unlikely to be the sole explanations for the dif- brunbrunneusbrunneousneus and S b endemicusendemicus the low propor- ferencesferences in ectoparasites between S b brun tion of parasitized S b brunbrunneusbrunneousneus 28 and S b neus and S b endemicusendemicus endemicusendemicus 70 in this study may have been there are significant differences in pelage partially because our collecting techniques length between S b brunbrunneusbrunneousneus and S b andemendem missed smaller ectoparasites however the same icus yensen 1991 interestingly the pelage of techniques were used for both subspecies S townsendtownsendiitownsendiaii is intermediate in length between therefore the sampling differences between the 2 S brunbrunneusbrunneousneus subspecies E yensen un- them should reflect real differences in parasite published data there also appear to be differ load thus with the number of animals and 244 GREAT BASIN naturalist volume 56

localities sampled the low overlap in lists of been identified to date but we can now add parasites is striking the following records further the low proportion of S brunneusbrunbrunneousneus with ectoparasites 55 especially in S b neopsyllaNeopsylla inopinainopine brunneusbrunbrunneousneus is atypical of Spennspermophilusophilus for ex- adams co 151.5lsis km N 151.5lsis km E bear guard ample hilton and mahrt 1971 found that in station 28 april 1995 6 m 7 flf adams co alberta 100 of S columbianuscolumbianus and S frank steve s creek 2 km S 2 km E bear 15 april liliilinii and 92 of S richardsorichardsoniirichardsonianii had ectopara 1995 8 m 7 flf adams co mouth of cold sites we were collecting S townsendtownsendiitownsendiaii and S springs creek 14 may 1995 1 m 1 f columbcolumbianusianus at the same time as S brunbrunneusbrunneousneus and were impressed by the much higher para- oropsyllaOropsylla idahoensis site loads on those species adams co 151.5lsis km N 151.5lsis km E bear guard although we did not observe plague in S station 28 april 1995 1 m 2 f adams co brunbrunneusbrunneousneus during this study it does occur in steve s creek 2 km S 2 km E bear 15 april southwestern idaho serum samples positive 1995 4 m 2 Qf adams co 3 km S bear 16 for tersina bestispestis the plague bacterium were april 1995 1 f reported from S townsendtownsendiitownsendiaii during a major ground squirrel die off in 1941 42 in ada oropsyllaOropsylla tuberculatetuberculatatuberculata tuberculatetuberculatatuberculata canyon and payette counties immediately adams co 151.5 km N 151.5lsis km E bear guard south of the range of S b brunbrunneusbrunneousneus hubbard station 28 april 1995 18 m 16 ffl adams co 1947 link 1955 in 1975 1977 positive anti- steve s creek 2 km S 2 km E bear 15 april body titers to plague were found in 72 91 1995 20 m 21 ffl adams co mouth of cold of badgers in the snake river birds of prey springs creek 14 may 1995 3 f area 50 km south of the range of S b eidemiendemi cus messick et al 1983 badgers are impor- thrassistarassisThrassis panpandoraedorae panpandoraedorae tant predators of ground squirrels eight of 9 adams co 151.5isls km N 151.5 km E bear guard dead townsend s ground squirrels examined station 28 april 1995 28 m 31 flf adams co by messick et al 1983 were positive for YV steve s creek 2 km S 2 km E bear 15 april bestispestis the plague bacterium has been detected 1995 8 m 15 f adams co 3 km S bear 16 in other species of Spennspermophilusophilus in all 5 idaho april 1995 1 m counties where S brunbrunneusbrunneousneus populations exist but until 1995 no S brunbrunneusbrunneousneus hadbad been exam- catallagiaCatallagia sp prob descipiens ined idaho department of health and welfare adams co 151.5lsis km N 151.5 km E bear guard personal observation in april 1995 T A gavin station 28 april 1995 1 f found a dead S b brunbrunneusbrunneousneus at the OX ranch and sent it to the wyoming state veterinary foxelladoxella ignota laboratory laramie where it was assigned case adams co steve s creek 2 km S 2 km E 95w3914 the carcass was found to be nega- bear 15 april 1995 4 m 3 ffl tive for Y bestispestis E williams personal comment nonetheless in the event of a plague epizootic spermophilus b brunbrunneusbrunneousneus is a new host local populations of S brunbrunneusbrunneousneus could easily be record for catallagiaCatallagia sp and foxelladoxella ignota decimated with only a small number of popu- Catallcatallagiaagia decipiensdecipiens is widely distributed in lations remaining plague could jeopardize the the western united states and is usually found survival of both subspecies ofofsjofsS brunbrunneusbrunneousneus on deer mice baird and saunders 1992 foxelladoxella ignota is commonly found on pocket NOTE ADDED IN PRESS gophers in the northern rocky mountains hubbard 1947 six hibernaculahibernacular of S b brunbrunneusbrunneousneus were exca- these new records also indicate that differ- vated in spring 1995 yensen and sherman ent sets of ectoparasites occur on S b brun unpublished data nests recovered from the neus and S b endeendemicusmicus thus corroborating hibernaculahibernacular were placed in plastic bags in the the earlier results the same 4 flea species field taken to the laboratory and then placed were again found associated with S b brun in berlese funnels small invertebrates were neus and neither catallagiaCatallagia nor foxelladoxella is collected in 70 ethanol only the fleas have known from S b endemicusendemicus 199611996 spermophilus BRUNNEUS ectoparasites 245

acknowledgments brunbrunneusbrunneousneus with reference to otherothel species of sper- momophilusphilus submitted GILL A YENSEN we thank elizabeth J dyni elizabeth E AND E 1992 biochemical differentiation in the idaho ground squirrel spermophilus domingue thomas A gavin D brad ham- brunbrunneusbrunneousneus rodentia sciuridae great basin natural- mond david oneill daniel A stevens and ist 52 155 159 william FE laurance for assistance in the field HILTON D FE J AND J L MAHRT 1971 ectoparasites robert E lewis iowa state university from three species of Spennspermophilusophilus rodentia sciuridae in alberta journal of zoology B in canadian 49 richard eads and eduardo campos centers 1501 1504 for disease control and elizabeth domingue HOFFMANN R S C G ANDERSON R W thorington cornell university kindly identified the fleas JR AND L R HEANEY 1993 family sciuridae pages nixon wilson university of northern iowa 419 465 in D E wilson and D M reeder editors mammal J E keirans rocky mountain laboratory and species of the world and2nd edition smith sonianaoniansoman institution press washington and london joann tenorio bishop museum identified the 1206 appp ticks and mites K C emerson national mu- holekampHOLEKAMF K E 1983 proximal mechanisms of natal dis- seum of natural history identified the lice persal in belding s ground squirrel spermophilus susan E wade cornell university identified belbeldingidingi belbeldingidingi unpublished doctoral dissertation university of californiaofcalifornia berkeley the nematode eyewormseye and amy worms doerger HOMINICK W M AND A J AsASTONTONmon 1981 association be- fields university of wyoming tested the spec- tween Pelodera stronglyoidesstrongly oides nematoda rhabditirhabditis imen for plague financial support was pro- dae and wood mice apodemusAp odemus sylvaticussylvaticus parasitolparasital vided by the national science foundation agyogyogy8383 67 75 HUBBARD 9225081 C A 1947 fleas of western north america DEB national geographic society their relation to public health iowa state college grant 3485863485 86 to PE W sherman and E press ames 533 appp yensen george C timtiatim hixon and univer- KINSELLA J M 1967 helminthshelminthes of microtinae in western sity of idaho agricultural experiment station montana canadian journal of zoology 45 269 274 R R L KARTMAN M I1 goldenberg AND Jjohnohn and jeanne dyer and pro- lechleitner tim hixon B W HUDSON 1968 an epizootic of plague min gun- vided encouragement housing I1 and access to nison s prairie dogs cynomys gunngunnisonigunmsomisoni in south research sites we thank sherilyn robison and central colorado ecology 49 734 743 rita colwell for helpful discussions and william LEWIS R E J H LEWIS AND C MASER 1988 the fleas of H clark eric eldredge james munger john the pacific northwest oregon state universityuniveiuniver sity press corvallis 296 appp whitaker and an anonymous referee for 0 jr LINK V B 1955 A history of plague in the united states constructive comments on an earlier version of ofamerica public health monograph 26 120 appp the manuscript MAY R M AND R M ANDERSON 1979 population biology of infectious diseases part II11 nature 280 455 461 CITED literature MESSICKJMESSICK J PPGG W SMITH AND A M BARNES 1983 seriserlsen ologicblogic testing of badgers to monitor plague in south- ANDERSON R M AND R M MAY 1979 population biol- western idaho journal of wildlife diseases 19 1 6 ogy of infectious 1 diseases partpartipantipanni nature 280 361 367 NADLER C F E A LYAPUNOVA R S HOITMANNHOFFMANN N N BAIRD C R AND R C SAUNDERS 1992 an annotated VORONTSOV L L shaitarova AND Y M BORISOV checklist of the fleas of idaho of university idaho 1984 chromosomal evolution in holarctic ground college of ofagriculture agriculture research bulletin 148 34 appp squirrels spermophilus II11 geimsa band homolo BURGESS G D 1955 arthropod ectoparasites of richard- gies of chromosomeschromo somes and the tempo of evolution son s ground squirrel journal of parasitology 35 zeltZeitzeitschriftzeitschnftschrift fur saugetierkundesdugetierkunde 49 78 90 325 352 POINAR G 0 1965 life history of Pelodera stronglystronglyoidesoides CLIFECLIFF ANDERSON G M R C AND FE F MALLORY 1978 schneider in the orbits of mundmuridmurld rodents of great dauelDauerdauerlarvaelarvae of Pelodera stronglyoidesstrongly oidesaides schneider britain proceedings oftheodtheof the helmmthologicalhelminthological society 1860 nematoda rhabditidae in the conjunctival of washington DC 32 148 lsiisi151511 sacs of lemmings canadian journal of zoology 56 PRATTPRATTHH DANDHD AND H E STARK 1973 fleas of public health 2117 2121 importance and their control US department of DAVIS W B 1939 the recent mammals of idaho caxton health education and welfare centers for disease printers ltd caldwell ID 400 appp control atlanta GA DHEW publication 74826774 8267 doss M A M M FARR K F ROACH AND G ANASTOS 40 appp 1974 index catalogue of medical and veterinary zool- RAYBURN J D M W HOOD M D KIRBY AND J H SHAW ogy special publication no 3 ticks and tick borne 1975 index catalogue of medical and veterinary diseases I1 genera and species of ticks part 2 gen- zoology supplement 19 part 5 parasite subject cata- era HNH N USU S department of agriculture agricul- logue parasites arthropodaartbropoda and miscellaneous phyla tural research service washington DC 593 appp USU S department of agricultureofagriculture agricultural resetichreseaichresearchReseaichsealch GAVIN T A P W SHERMAN E YENSEN AND B MAY pop- service washington DC 403 appp ulation structure and gene flow among disjunct pop- RAYOR L S 1985 dynamics of a plague outbreak in gunni ulationsulations of idaho ground squirrels spermophilus soisolsonss prairie dog journal of mammalogy 66 194 196 246 GREAT BASIN naturalist volume 56

SCHULTE F 1989 history life of rhabditis PeFelodera TRUKSA A S AND E YENSEN 1990 photographic evi-evievl orbitaorbitoorbitalsorbitalisorbitahsorbitalishs a larval parasite in the eye orbits of avri dence of vegetation changes in adams county idaho calidcolid and mundmuridmurld rodents proceedings of the helmint- journal of the idaho academy of science 26 181018 40 hologicalth 1 isioisto society of washington DC 56 7 WHITAKER J 0 JR 1979 origin and evolution of the ex- SHAW H AND M HOOD J W 1975 index catalogue of ternal parasite fauna of western jumping mice genus medical and veterinary zoology supplement 20 part zapus american midland naturalist 101 49 60 5 parasite parasites subject catalogue arthropoda WHITAKER J 0 JR AND N WILSON 1974 host and dis- and S miscellaneous phyla USU department of agri- tributiontribution lists of mites acarlacariacan parasitic and phoreticphophoneticretic culture agricultural research service washington in the hair of wild mammals of north america north DC 223 appp of mexico american midland naturalist 91 1 67 SHERMAN P W 1989 mate guarding as paternity insurance YENSEN E 1991 taxonomy and distribution of the idaho in idaho ground squirrels nature 338 418 420 ground squirrel spermophilusSpennophilus brunbrunneusbrunneousneus journal of STARK H E 1970 A revision of the flea genus thrassmthrassusthoasThrassus mammalogy 7258372 583 600 jordan 1933 siphonaptera ceratophyllidae with YENSEN E AND P W SHERMAN in press spehnSpennspermophilusophilus observations on ecology and relationship to plague brunneusbrunbrunneousneus mammalian species university of california publications in entomology 53153 1 184 received 29 march 1995 strandtmann R W 1949 the bloodsuckingblood- sucking mites of the accepted 2 may 1996 genus haemolaelaps acarina laelaptidae in the united states journal of parasitology 35 325 352 great basin naturalist 563 V 1996 appp 247 253

ROOST SITES OF THE SILVER HAIRED BAT lasionycterislasioivycteris noctivacansnoctivagantnoctivagans IN THE BLACK HILLS SOUTH DAKOTA

todd A Mattsonmattsonl2mattson1212 steven W BuskirBuskirkkl1 and nancy L stantonlStanton1

ABSTRACT we investigatinvestigate the roosting ecology of silver haired bats lasionycterislasionyctens noctivagantnoctivagans in the black hills of western south dakota using i oteleotelemetrymetry we located 39 boostsroosts 10 of which were maternity aggregations containing 6 to 55 bats the boostsroosts were mostly in ponderosa pine pinus ponderosa snags that averaged 39 cm diameter at breast height solitary bats preferred roosting under loose bark or in crevices in trees regularly moving among trees all maternity aggregations were found in tree cavities primarily those created by woodpeckers roost trees were located in patches of forest with relatively high snag densities about 21 snagshasnagsha this study suggests that snags play an important role in maintaining silver haired bat populations in ponderosa pine ecosystems

key words lasionyctenslasionycteris noctivagansnoctivagant silver haired bat boostsroosts snags

the silver haired bat lasionycteris noctivanoctwa hills of south dakota although forests in this gans occurs widely across north america at region have been intensively managed for tim- highly variable densities barbour and davis ber boldt and van deuendauendeuoen 1974 silver haired 1969 kunz 1982a studies conducted in the bats are relatively abundant compared to the 9 northwestern united states suggest that silver other bat species present in the region matt- haired bats occur more frequently in late suc son 1994 although mattson 1994 captured cesssessionalcessionalional forests dominated by trees over 200 twice as many males as females pregnant or yr old than in early seres perkins and cross lactating females were not uncommon our goal 1988 thomas 1988 this association is attrib- was to characterize roost selection by silver uted to the presence of high concentrations of haired bats in terms of attributes potentially standing dead trees some of which have exfoli affected by current forestry practices atingabing bark cracks in the wood and cavities excavated by birds sites that may be pre- STUDY AREA ferred by bats for roosting perkins and cross 1988 thomas 1988 campbell et al in press our study area is located in the southern little information on summer roost sites for black hills of south dakota near the town of silver haired bats is available kunz 1982a custer 4346n 10335w most of the study particularly in areas that lack abundant stands area is in the black hills national forest and of late successional forests barclay et al 1988 occurs at elevations from 1360 to 1985 m asl searched trees in manitoba and found silver the topography of the area varies from rolling haired bats roosting under folds of loose bark highlands with parklike valleys to narrow steep during the migration period parsons et al canyons with rocky ridge tops the climate of 1986 reported observations of 2 small silver the black hills differs from the surrounding haired bat maternity colonies in hollow trees in semiarid plains in that it is moister and less canada likewise maternity colonies have subject to temperature extremes average maxi- been found in cavities of both live and dead mum temperature at custer in july is about 23 trees in california rainey et al in press C while mean annual precipitation is 457 mm despite these records a clear understanding the forests of the area are dominated by pure of silversliver haired bat boostsroosts and roost habitat is stands of ponderosa pine pinus ponderosa still lacking small stands of quaking aspen populus tremuteemu to better understand the roost requirements loides precede ponderosa pine on disturbed of silver haired bats we investigated roost sites paper birch betula papyrpapyriferaifera grows in selection by the silver haired bat in the black small clusters in more mesic sites whereas

1 department ofzoologyandof zoology and physiology university of wyoming laramie WY 82071316682071 3166 present address PIC technologies inc 309 south ath4th street suite 201 laramie WY 82070

247 248 GREAT BASIN naturalist volume 56

rocky mountain juniper Jumjunipentsjumperusperus scopulorum with others we returned to the tree before grows on dry ridges dusk to watch and count bats leaving the site the forests of the black hills have been we attempted to approach the tree quietly to managed for timber production since logging reduce disturbance we used a bat detector first began in the 1870s during the past 100 bat box liililIII111 stag electronics st agnes eng- yr most areas have been cut once and many land to listen for echolocation calls these have experienced multiple partial cuts alexan- along with body size and flight pattern were der 1987 in all nearly 12 X 106 m3ma of timber used to confirm that bats in a given roost were has been removed only a few small scattered only silversliver haired bats stands of unharvested forest remain boldt and remain roost measurements van deusen 1974 although clearcuttingclearcuttmgclearelearcutting was once the primary means of harvest shelter- we located 18 roost trees in the jewel cave wood cutting a method using a seriesserles of cuts study site and 21 in the hazelrodtnazelrodtHazelrodt study site is now standard when possible the type of roost ie wood- we delineated two 10110loi10.11 X 10110loi 1 km study pecker cavity crevice loose bark etc was sites in areas in which we located silver haired recorded each roost tree was classified as bat boostsroosts the jewel cave study site encom- being used by either a maternity aggregation passes jewel cave national monument and or solitary bats maternity roosis located by adjacent areas of the black hills national for- tracking pregnant and lactating females always est the hazelrodtnazelrodtHazelrodt study site is located south- contained 6 or more bats solitary boostsroosts con- east of custer on national forest land and tained only a single bat and were located by custer state park much of the hazelrodtnazelrodtHazelrodt tracking males or females that did not appear study site burned during a fire in 1990 that pregnant or lactating or were post lactating covered over 5670 ha we categorized the aspect of the roost exit as northeast 0 89 southeast 90 179 south- maremxreMALERIALSRIALS AND METHODS west 180 269 or northwest 270 359 each roost tree was identified to capture and tracking techniques species and its height and diameter at breast height silver haired bats were captured using mist abbdbhdbb measured we placed each roost tree into nets set above small ponds and streams be- 1 of 7 decay stages decay stage I1 included live tween 25 june and 4 august 1994 we deter- trees with intact bark and branches whereas mined the sex and reproductive condition for decay stage 7 included dead trees beginning to all captured bats using external features racey decompose with broken tops and no loose bark 1988 bats were classified as adult or juvenile thomas et al 1979 based on fusion of the epiphyseal diaphysialdiaphyseal plot measurements suture of the finger bones anthony 1988 we attached 070790 7 g radio transmitters model within a 5 m radius 78 m2ma circular plot BD ab2b holohil systems ltd woodlawn centered at each roost tree we measured aver- ontario to 4 adult males and 12 adult females age tree size total basal area and snag density after fur had been trimmed from the bats trees were defined as standing woody stems transmitters were attached to the area between isls151.5 in in height and 10 cm abbdbhdbb we also the shoulder blades using a cyanoacrylate based recorded whether disturbance by fire or log- glue fing rs camarilloCamanlloaniloaniio CA bats to which ging had taken place in each plot disturbance transmitters were affixed weighed 11 14 g so by fire was considered to have occurred if there that transmitters represented 5 646.46 4 of body was any charred woody material in the plot mass slightly over the 5 maximum recom- and disturbance by logging was noted if we mended by aldridge and bnghambrighambangham 1988 we observed any saw cuts on woody material in did not use any other marking technique to the plot identify individuals to compare characteristics of roost site plots hand held 3 element yagi antennas and with the surrounding areas we located four 5 portable receivers model TR 2 telonicstelomcsTelomcsmes mesa m radius neighborhood plots for each roost AZ were used to track bats to roost trees if plot and recorded the same information as for we were unable to determine where in the tree roost plots we located the center of the neigh- the bat was roosting or whether it was alone or borhood plots by pacing 100 in from the roost 199611996 SILVER HAIRED BAT ROOSTS 249

tree in each of the cardinal directions north which occurred in trees nine adult females south east west and then pacing an additional were tracked to 10 trees that were used by 30 m in a randomly selected direction maternity aggregations averaging 22222.222 2 49 we measured elevation and distance to the sifsicsacs3cs individuals range 6 55 three other nearest source of water for each roost tree using females and 4 adult males were tracked to 25 topographic maps 75757.5 minute series USGS roost trees none of which were used by mater- denver CO for comparison we randomly nity aggregations three of the females that located a point in the jewel cave study site or originally used maternity aggregations were hazelrodtnazelrodtHazelrodt study site for each roost tree found later followed to 4 trees where they roosted in that site to examine roost site selection on a alone maternity boostsroosts were found exclusively larger scale we calculated the number of snags in tree cavities primarily those created by wood- all in neighborhood plots to estimate snag den- peckers picidae cavity openings were 757.57 5 10 sity for the study site generally this estimation cm in diameter solitary bats roosted under was made by dividing the total number of snags loose bark n 15 in a tree crack or crevice in the 156 neighborhood plots by their total n 5 or in a woodpecker cavity n 1 we area the fire in the hazelrodtnazelrodtHazelrodt study site in- could not determine the specific roost location flated snag densities in this area to remove the for 8 trees these trees were placed in the soli- influence of fire we calculated snag densities tary category because bats tracked to these 8 within the study sites by removing the 77 neigh- trees were always observed ostingroostingboostingro alone at borhood plots that had been disturbed by fire other trees maternity boostsroosts were 10210.210 2 151.5lsis1 5 m range 3.1313 1 13.813138 8 analysis 31 138 aboveground the height of measured solitary boostsroosts averaged 343.43 4 050.50os 5 m chi square tests for goodness of fit jelinski range 090.9og0 9 898.98 9 cavity openings of maternity 1991 were used to compare observed with ex- boostsroosts and solitary bat boostsroosts were found more pected roost aspects and tree decay stages by frequently on the south side of tree boles over roost type maternity vs solitary for the latter other aspects xax2 15815.815 8 Hd f 3 P 000100.001ooi001 test because of small sample size we pooled of 39 roost trees 38 97 were ponderosa the roost trees into 3 decay stage categories pine and 1 3 was aspen of 508 trees on stage 1 3 stage 4 and stage 575 7 neighborhood plots 483 95 were ponderosa to compare continuous attributes between pine and 25 5 were other species aspen roost plots and neighborhood plots we sub- juniper and paper birch the 10 trees used by tracted attribute means for the 4 neighborhood maternity aggregations of silver haired bats plots from corresponding means for the roost ranged from decay stage 2 to 7 median 5 plots so each roost plot was compared only to the 29 trees used by solitary bats varied from its 4 neighborhood plots we tested the null tree decay stage 3 to 7 median 4 trees in hypotheses that the mean differences did not neighborhood plots ranged from decay stage I1 differ from 0 using paired t tests chi square to 7 median 1 bats in maternity aggrega- tests for homogeneity jelinski 1991 were used tions selected roost trees in significantly differ- to compare observed with expected disturbances ent decay stages than solitary roosting bats xax22 at roost plots expected disturbances were based 10210.210 2 dfd f 2 P 000620.00620 0062 fig 1 roost trees on the proportion of neighborhood plots that averaged 14214.214 2 090.90og 9 m range 373.73 7 24 1 in total had burned or been logged we used 2 sample height and 39 2 cm dahdbh range 13 63 they t tests to compare the means for elevation and averaged 17 2 cm larger in dahdbh than neigh- distance to nearest water for roost sites and borhood trees the 10 maternity roost trees random sites to avoid type I1 errors that may averaged 44 4 cm dahdbh range 29 62 24 4 result from using a number of inferential statis- cm larger than neighborhood trees the 29 soli- tical tests with the same predictor variable we tary roost trees averaged 37 2 cm dahdbh range arbitrarily set a 00250.025 12 55 15 3 cm larger than neighborhood trees maternity and solitary roost trees did not RESULTS differ in diameter t 1641.641 64 P 00120.1212 roost attributes the 9 bats found in maternity aggregations returned to the same roost tree for a mean of we radio tracked 16 bats for a mean of 8 d 8 d range 1 21 we tracked 1 bat from a tree range 1 20 and located 39 boostsroosts all of containing a maternity aggregation of 55 bats 250 GREAT BASIN naturalist volume 56

so-

d 0 40 0001 s 30 i 20- 10 I1 1 a 0 r 131 3 4 575 7 tree decay stagstage e categories

available n132 YM aggregation nton10 W solitary n29

fig 1 percentages of trees in each tree decay stage category used by maternity aggregations and solitary roosting silver hairedhailhall ed bats and available trees in the black hills south dakota june august 1994 to a and2nd tree with a maternity aggregation of live and dead trees on roost plots were 656.5gs 44 bats about 440 m away the following eve- 171.7 cm larger in abbdbhdbb on average than those on ning no bats were observed exiting from the neighborhood plots t 3773.77 P 000060.0006 ist roost tree but it is not clear how many bats roost plots also had basal areas of both live from the ist roost tree moved to the and2nd tree and dead standing trees that were 140714.07 3463.46 with the bat we were tracking cm2cnm2m2 greater t 4064.06 P 000020.0002 than we tracked 10 bats that used solitary boostsroosts neighborhood plots neither fire disturbance 2 to a mean of 3 solitary roost trees range 1 6 xaXx2 00050.005 df 1 P 0940.94 nor logging for the most part these bats switched trees disturbance x2xaqc2 2722.72 df 1 P 00990.099 dif- daily however on 5 occasions solitary bats fered between roost and neighborhood plots used the same tree on consecutive days three maternity and solitary plots did not differ in the of the 7 solitary roosting bats that we followed attributes studied table 1 roost trees tended to multiple trees returned at least once to trees to be located higher in elevation than random they had used several days before solitary points t 1671.67 P 0100.10 roost sites were roosting bats traveledtiaveled a mean of 405 93793.793 7 m significantly farther from water than random 2.78 P 0.007 n 13 between successive roost trees radio points t 278 0007 all 156 neighborhood plots we calcu- tracked bats traveled a mean of 2060 440 in using lated snag density for the area to be 117 snags n 12 from the capture point to their first ha after removing 77 neighborhood plots that roost tree significantly farther t 3673.673 67 P were disturbed by fire we recalculated snag 00040.0040 004 than the distance between successive successive densities to be 21 snagshasnag roost trees sha plot attributes discussion roost plots had 171.71 7 060.60og 6 more live trees t boostsroosts used by maternity aggregations dif- 3093.093 09 P 00040.0040 004 than neighborhood plots fered from those used by solitary silver haired 199619961 SILVER HAIRED BAT ROOSTS 251

TABLE 1 comparison between solitary and maternity roost plot attributes in the black hills south dakota june august 1994 solitary maternity attribute n 29 n 10 T FP value live trees 4506454.5 06 5207525.2 070.7 072 047 noblotnoplotno plotpiot snags 212.1gi91210505 222.222052905050.5 015 088 noblotnoplotno plotpiot mean tree dbhdah 26726.726 7 323 2 27527.527 5 191.9iglg1 9 022 083 cm total basal area 17817.817 8 131 3 25325.325 3 474.74 7 154 013 cm22cmcm2m222

bats maternity aggregations always used a hol- of cavities with south facing entrances reflects low cavity within a tree bole usually these the selections ofbats or woodpeckers cavities were created by woodpeckers likely silver haired bats roosted exclusively in trees hairy woodpeckers ficoidesPicpicoidesoides villosisvillosusvillosus or black during the summer although all but one of the backed woodpeckers P arcticusarcticus based on the boostsroosts were located in ponderosa pine trees size of the openings terres 1980 although the dominance of ponderosa pine in our study rare in the black hills black hills national area prevented us from testing for tree species forest 1989 lewis woodpeckers melanerpes preference the wide geographic distribution lewis northern flickers colantescolaptes auauratusauranusratus or of silver haired bats relative to that of pon- three toed woodpeckers ficoidesPicpicoidesoides tridactylustridactylustridactylous derosa pine and the use by silver haired bats of may have excavated some of the cavities soli- both coniferous and deciduous roost trees in tary boostsroosts were located under loose bark or in other parts of their range novakowski 1956 a natural crack or crevice in the tree bole only parsons et al 1986 barclay et al 1988 camp- once did a solitary bat use a woodpecker cavity bell et al in press rainey et al in press sug- although silver haired bats are cryptically col- gest that these bats select for the structure of ored they were never observed roosting openly the roost itself rather than for a particular tree on a tree trunk or limb or in foliage this be- species As for other tree roostingboosting bats tide- havior differs from other cryptically colored mann and havelnaveiflavelmavel 1987 it is unlikely that tree tree roostingboosting bats eg lasiurusLasiurus sppapp which species is important to silver haired bats extend to roost among tree foliage shump and cept that at the local level 1 species may tend shump 1982a 1982b boostsroosts required by ma- to have preferred attributes ternity aggregations may limit silver haired bat roost trees were standing dead and larger abundance clearly trees with cavities are less than average in diameter the single living tree available than are those without reproductive selected as a roost was dying stage 2 and females seem to require boostsroosts that provide a missing its top it also had many dead limbs relatively enclosed and unexposed space for and several woodpecker holes high in the bole protecting young from predators or maintain- there was an observed difference in tree decay ing the necessary thermal environment stage between roost trees used by maternity cavity openings of maternity boostsroosts and aggregations and solitary bats solitary roosting solitary bat boostsroosts occurred more frequently bats frequently used trees in decay stage 4 than expected on the south side of tree boles which are characterized by the presence of we hypothesize that these boostsroosts are warmer loose bark alternatively maternity boostsroosts tended than sites facing north because of insolation to be found in older more decomposed trees and that these differences result in energetic decay stages 5 7 trees that are more com- savings providing more energy for growth and monly used by excavating woodpeckers thomas development mcnab 1982 reller 1972 has et al 1979 although the importance of snags shown that several species of woodpeckers ori- as roost sites in other forest types remains in ent their nest cavity openings southwesterly for question large snags appear to be important warming by the sun andor ventilation by the resources for silver haired bats in ponderosa wind however it is unclear whether bat use pine forests 252 GREAT BASIN naturalist volume 56

clearly solitary roosting silver haired bats study site an area with a large number of fire switch boostsroosts regularly this lack of fidelity may killed trees how fire suppression and logging be related to the abundant nature of potential practices have affected the number of snags in boostsroosts brigham 1991 or a predator avoidance the black hills remains unclear however early strategy kunz 1982b because they will return photographs suggest that many forested areas to roost trees used several days previously and were more open with many standing dead these boostsroosts are often close together solitary trees knight 1994 because snags are used for bats may use a series of trees in the same area nests or boostsroosts by a large number of vertebrate and thus maintain a level of site familiarity species thomas et al 1979 reduced snag conversely maternity aggregations tend to re- densities may increase interspecific competi- main in the same boostsroosts for longer periods tion we hypothesize that forest management this may be related to the less abundant nature practices that reduce snag densities will lead to of tree cavities and the importance of retaining declines in local silver haired bat populations boostsroosts that are suitable for raising offspring at least some of the maternity aggregations appear acknowledgments to switch boostsroosts during the reproductive period the reason for this is not clear although it may funding was provided by the university of involve predator or ectoparasite avoidance wyoming national park service research cen- lewis 1995 ter the national park service rocky moun- we expected bats to select boostsroosts relatively tain region and the national biological sur- close to water bodies minimizing energetic vey midcontinentMidcontinent ecological research center costs of moving between roosting areas and we especially appreciate support from kate areas potentially used for drinking and forag- cannon and the staff of jewel cave national ing although trees were abundant in the study monument from mike bogan of the national sites bats traveled an average of 2 km from biological survey and from joel tigner oscar point of capture to their ist roost tree and sig- martinezMartmez and alice lippacher of the black hills nificantlynificantly farther from water than expected national forest mike bjelland and jay grant randomly this seems to support other avail- provided valuable field assistance we thank able evidence for insectivorous bats in that thomas H kunz and R mark brigham for com- roost site location is not strongly influenced by menting on an earlier draft of the manuscript commuting costs fenton et al 1985 brigham 1991 roost sites located farther flomfrom water literature CITED than random points appear puzzling but may represent the large number of roost trees located ALDRIDGE H D J N AND R M BRIGHAM 1988 load and along hillbill ridge carrying maneuverability in an insectivorous bat or tops sites with potentially a test of the 5 rule of radio telemetry journal of higher snag densities mammalogy 6937969 379 382 silveisilversliver haired bat roost trees were found at ALEXANDER R R 1987 silvicultural systems cutting sites that differed from nearby areas in a num- methods and cultural practices for black hills pon- ber of attributes roost plots differed having derosa pine USDA forest service general technical in report RM 139 rocky mountain forest and range more large trees and hence a higher total basal experiment station fort collins CO 32 appp area than surrounding plots roost trees located ANTHONY E L P 1988 age determination in bats pages in areas that are ideal for tree growth or are 47 58 in T H kunz editor ecological and behav- logged infrequently might explain why the roost ioral methods for the study of bats smithsonianSmithsoman institution press washington plots have more larger trees DC BARBOUR R W AND W H DAVIS 1969 bats of america undoubtedly snags are important in pro- university of kentucky lexington 286 appp viding roost sites for silver haired bats in the BARCLAY R M R P A FAURE AND D R FARR 1988 black hills As suitable boostsroosts are critical roostingboostingRoosting behavior and roost selection by migrating silversliver haired bats resources for bat survival 1982b lasionycterislasionyctens noctivagantnoctivagans journal kunz snag of mammalogy 69 821 825 availability likely influences the distribution BLACK HILLS NATIONAL FOREST 1989 black hills national and abundance of this species forest stands foiestforest checklist ofbirdsofbirds USDA forest service containing silver haired bat boostsroosts had snag BOLDT C E AND J L VAN DEUSEN 1974 silviculture of densities of 21 snagshasnagsha a value much higher ponderosa pine in the black hills the status of our knowledge USDA paper than objectives forest service research current management these RM 124 rocky mountain forest and range experi- densities were even higher in the hazelrodtnazelrodtHaz elrodt ment station fort collins CO 199619961 SILVER HAIRED BAT ROOSTS 253

BRIGHAM R M 1991 flexibility in foraging and roosting teristefistens noctivagantnoctivagans in ontario and saskatchewan joinjourjoun behaviour by the big brown bat eptesicus fuscus nal of mammalogy 67 598 600 canadian journal of zoology 69 117 121211 PERKINSPEBKINS J M AND S P CROSS 1988 differential use of CAMPBELL L A J G HALLET AND M A OCONNELL in some coniferous forest habitats by hoary and silversilveisliver press conservation ofbatsof bats in managed forests use of haired bats in oregon murrelet 69 21 24 boostsroosts by lasionycteris noctivagantnoctivagans journal of mam- RACEY P A 1988 reproductive assessment in bats pages malogy 31 45 in T H kunz editor ecological and behav- FENTON M B R M BRIGHAM A M MILLS AND I1 L ioral methods for the study of bats smithsonian rautenbach 1985 the roosting and foraging areas institution press washington DC of epomophorus wahlwahlbergibergi pteropodidae and scoto RAINEY W E J SIPEREK AND R M MILLER in piesspress philus viridis vespertilionidae in kruger national colonial maternity boostsroosts of the silver haired bat park south africa journal of mammalogy 66 lasionycterislasionyctens noctivagantnoctivagansnoctwagans in northern california 461 468 forests american midland naturalist JELINSKI D E 1991 on the use of chi square in studies RELLER A W 1972 aspects of behavioral ecology of red- of resource utilization canadian journal of forest headed and red bellied Woodpeckwoodpeckerseisels american research 21 58 65 midland naturalist 88 270 290 KNIGHT D H 1994 mountains and plains the ecology of SHUMP K A AND A U SHUMP 1982a lasiurusLasiurus borealis wyoming landscapes yale university press new mammalian species 183 american society of mam haven and london 338 appp malogists new york 6 appp KUNZ T H 1982a lasionycteris noctivagantnoctivagans mammalian 1982b lasiurusLasiurus cictnereus mammalian species 185 species 172 american society of mammalogists american society of mammalogists new york 5 appp new york 5 appp TERRES J K 1980 the audubon society encyclopedia of 1982b roostingboostingRo osting ecology of bats pages 1 55 in north american birds wings books avenel NJ T H kunz editor ecology of bats plenum press 1109 appp new york THOMAS D W 1988 the distribution of bats in different LEWIS S E 1995 roost fidelity of bats a review journal ages of douglas fir forests journal of wildlife man- of mammalogy 76 481 496 agement 52 619 626 MATTSON T A 1994 the distribution ofbatsofbats and the roost THOMAS J W R G ANDERSONANDEBSON C MASER AND E L BULL ing ecology of the silver haired bat lasionycteris 1979 snags pages 60 77 in J W thomas editor noctivagantnoctivagans in the black hills of south dakota wildlife habitats in managed forests the blue moun- unpublished master s thesis university of wyomingofwyoming tains of oregon and washington USDAUS DA forest ser- laramie vice agricultural handbook 553 mcnab B K 1982 evolutionary alternatives in the physi- TIDEMANN C R AND S C FLAVEL 1987 factors affect- ological ecology of bats pages 151 200 in T H kunz ing the choice of diurnal roost site by tree hole bats editor ecology ofbatsofbats plenum press new york microchiropteraMiciochiroptera in southeasternsouth eastern australia aus- novakowski N S 1956 additional records of bats in sas- tralian wildlife research 1445914 459 473 katekatchewankatchhewanbewanewan canadian field naturalist 70 142 PARSONS H J D A SMITH AND R E WHITTAM 1986 received 13 december 1995 maternity colonies of silversliver haired bats lasionyc accepted 17 may 1996 cleatgleatgreat basin naturalist 563 0 1996 appp 254 260

perceptions OF UTAH ALFALFA GROWERS ABOUTAROUT WILDLIFE DAMAGE TO THEIR HAY CROPS implications FOR MANAGING WILDLIFE ON PRIVATE LAND

terry A MessmermessmerlMessmessmer1merlmeri1 and sue Schroeder 2

AnsTHAABSTRACTCr weWC conducted a survey of utah alfalfa medwagomedicagoMedwago satica growers in 1993 to identify wildlife damage pioblemsproblemsproblerns to hay crops suehsuchstich surveys can provide wildlife managers with important insights regarding landowners wildlife damage management concerns and needs pocket gophers thomomys sppapp and mule deer odocoileusodecoileus hemionus were perceived by growers as causing the most damage respondents reported a total annual loss of 350000 oior 2479ha247924 79haha 282 8 of the total crop value because of wildlife damage in alfalfa crops decreased hay quantity was the most frequently cited problem caused by wildlife compensation and incentive programs were preferred over assistance and informationmfoimafoi mationmatlon programs foiforboibolborror managing wildlife damage in alfalfa crops

key words wildlife damage perceptions alfalfa growers wildlife damage management wildlife management

alfalfa is an important livestock forage in improving wildlife habitat on private property 1994 over 58 million tons of alfalfa hay were will succeed conover 1994 harvested in the US on 9802400 ha of pri- there is consensus among professionals vately owned land this represents over 40 of working for federal and state wildlife and agri- the haybay harvested as livestock forage national cultural agencies that wildlife damage reduces agricultural statistics service 1995 the profitability of US agriculture conover alfalfa hay is the most important cash crop and decker 1991 professionals agree that wild- grown in utah in 1994 utah farmers harvested life depredation has increased over time but 2205000 tons of alfalfa on 210000 ha of pri- disagree over the seriousness of the impact vately owned land this crop was worth 158 although the actual costs associated with wild- million gneiting 1994 life depredation are difficult to estimate and rodents lagomorphslagomorpha ungulaungulatesungulatedtes and water- can differ on each farm or ranch and crop type fowl can impact alfalfa production piper 1909 tebaldi and anderson 1982 austin and umessurness sauer 1978 luce et al 1981 dunn et al 1982 1987a 1987b 1989 1993 lewis and obrien packalpackampackarn and connolly 1992 austin and urness 1990 landowners have demonstrated an abil- 1993 conover 1994 big game grazing of alfalfa ity to accurately assess crop losses caused by during the growing season creates conflicts be- wildlife decker et al 1984 conover 1994 tween growers and wildlife managers austin melvormcivor and conover 1994a crop losses and and urness 1993 potential future losses caused by or related to conflicts also may arise between landown- the presence of wildlife must be assessed to ers and wildlife managers because of differing determine if control is warranted rennison perceptions about the extent of wildlife damage and buckle 1988 in cultivated crops farmers may feel that wild- several great basin states including utah life managers are unaware of the extent of crop wyoming colorado new mexico nevada losses caused by wildlife and hence are insen- idaho and arizona have enacted laws to com- sitive to their needs decker et al 1984 pensate crop owners for wildlife caused dam- conover and decker 1991 crop owners con- age musgrave and stein 1993 these actions cerns about wildlife damage strongly affect have been initiated largely in response to con- how the agricultural community will respond stituent concerns over the economic impact of to environmental issues and whether federal or predatingdepredatingdeprecatingde wildlife particularly big game in state wildlife programs aimed at maintaining or cultivated crops

cpaitmentdepartnient of fisheries and wildlife utah state university logan UT 84322521084322 5210 2departinentepaitinent ofrorestu forestfoiesthorest resources utah state university logan UT 84322521584322 5215

254 199611996 WILDLIFE DAMAGE TO ALFALFA 255

crop owners in utah may destroy depredatdepredate common management practices used on their ing big game animals if the animals are not farms and ranches removed by the utah division of wildlife responses were stratified and analyzed by resources UDWR within 72 h of notification the number of hectares in alfalfa 0 40 41 80 chapter 183 utah code 1993a utah crop 81 200 201 400 and 400 and type of oper- owners also may receive monetary compensa- ation irrigated or dryland levere s tests were tion for damage caused by big game animals used to determine equality of variances by types chapter 307 utah code 1994b and ring and sizes of alfalfa operation SPSS 1995 necked pheasants phasianus colchicuscolchicum chap- we assumed that alfalfa growers on the hay ter 46 utah code 1971 list have the same values and perceptions as we surveyed utah alfalfa growers to deter- the population of utah alfalfa growers to determine their perceptions regarding wildlife dam- mine if the hay list was statistically representa- age to hay crops such surveys can provide tive of utah alfalfa growers we compared the wildlife managers with important information mean alfalfa farm size and regional distribu- regarding landowner wildlife damage manage- tions of farms on the hay list with acreage cate- ment needs and concerns conover 1994 gories reported by the UDA for all utah alfalfa farms gneiting 1994 using a kruskal wallis METHODS one way analysis of variance differences in these tests were considered significant if P we surveyed 334 alfalfa growers 4 of all 005oos0.05 alfalfa growers in utah whose names were on the utah department of agriculture s UDA RESULTS 1993 hay list the UDA maintains this list to provide information to individuals who contact alfalfa production the department about purchasing alfalfa hay in one hundred sixty four completed ques- utah the UDA updates this list each january tionnationnairesires 49149149.1 were returned of which 150 we included a 2 page wildlife damage sur- 91 were useable for analysis survey respon- vey in a UDA mailing sent to the growers in dents reported growing 16867 ha of alfalfa of addition to the survey growers received a cover which 14391 ha 85 was irrigated and 2486 letter the udsUDAUDNs s questionnaire and a business ha 15 was dryland alfalfa irrigated alfalfa reply envelope the cover letter stated that if farms ranged in size from 5 to 1062 ha dry- no response was received within 30 d the land alfalfa farms ranged in size from 3 to 320 grower s name would be removed from the hay ha all farms were family owned and operated list A followupfollow up letter was sent to nonrespon- since the UDA hay list is relatively dynam- dents 3 wk after the initial mailing those fail- ic it contains information regarding the grower s ing to respond to the and2nd mailing were removed mailing address telephone number and inter- from the hay list est in selling alfalfa hay but not the size and the survey contained questions about the type of operation information on alfalfa opera- growers experiences with wildlife in their alfalfa tions was obtained through the survey thus growers crops were asked to identify wildlife we were unable to determine if there were any species causing damage to hay crops type of significant differences between respondents damage their annual monetary loss from wild- and nonrespondents life damage specific damage control techniques although the responses received consti- employed on their farm to control wildlife dam- tuted 2 of all utah alfalfa growers N 7600 age whether they received any type of damage our sample was representative of the popula- compensation or assistance who they contacted tion based on mean farm size H 707.0to 7 df P for assistance and information and what type 00010.001 and regional distribution utah alfalfa of information and programs they found most acreage percentages reported by the UDA for useful in managing wildlife damage further northern central eastern and southern regions growers were asked to rate on a scale of 0 to 5 were 30 31 19 and 20 respectively 0 no cost through 5 high cost relative gneiting 1994 regional alfalfa acreage per- losses caused by different wildlife species to cencentagestages for our sample were northern 27 their alfalfa crops and the costs associated with central 34 eastern 21 and southern 18 256 GREAT BASIN naturalist volume 56

wildlife species present in damage losses by size for irrigatedirrigateddrylanddryland utah alfalfa fields alfalfa farms F 040.4 126 df P 0520.52 respondents with alfalfa farms 80 ha re- respondents reported 20 different species ported that rodents F 797.9 1107 df P of wildlife present in their alfalfa fields were in 00060.006 and ungulatesungulatedtes F 18218.2 1107 df P pocket ungula gophers and mule deer were the most 00010.001 caused higher monetary losses when abundant being reported present on 124 compared to smaller farms 80 ha no signif- 827 and 120 800 farms respectively icant differences in monetary losses due to water- other wildlife species reported by farmers as fowl were detected by alfalfa farm size F common in alfalfa fields included jackrabbitsjackrabbits 00060.006 1107 df P 09400.940 lepus sppapp n 89 59359.359 3 ground squirrels relative costs of wildlife spermophilus sppapp n 83 55355.355 3 prairie damage in alfalfa fields dogs cynomys sppapp n 69 46046.046 0 waterfowl anatidae n 66 44044.044 0 elk cervus elaphusclaphus respondents ranked on a scale of 0 5 0 n 62 41341.341 3 pronghorn antilocapra ameriamerlemenamen no cost through 5 high cost the relative cana n 54 36036.036 0 and voles microtus sppapp damage costs associated with common wildlife n 50 33333.333 3 wildlife species reported by species reported in their alfalfa fields as fol- farmers as being less common in alfalfa fields lows mule deer 29292.9 pocket gophers 24242.4 elk included marmottmarmots mannotaflaviventrismarmota flamventns bad- lgig161.6 prairie dogs 14141.4 ground squirrels 141.4 gers taxidea taxus red foxes vulpes vulpes jackrabbitsjackrabbits 13131.3 waterfowl loio101.0 pronghorn sandhill cranes grus canadensis canada geese 07070.7 and meadow voles 09og090.9 respondents branta canadensis cottontail rabbits syvila with irrigated alfalfa farms 200 ha reported tg gus sppapp deer mice peromyscus maniculatusmamculatus that elk F 797.9 156 df P 007007.007 and prong- 75 oos raccoons procyon lotorbotor ring necked pheas- horn F 757.5 148 df P 008008.008 caused signif- ants and muskrats ondatra zibethicazibethica icantly greater cost related problems than on smaller farms 200 ha respondents with monetary losses caused by wildlife dryland alfalfa farms 200 ha reported greater one hundred nine growers 72 reported significant cost related problems caused by losing 350000 x 3242 s 526 be- jackrabbitsjackrabbits F 14114.1 120 df P 00010.001 and mule 85 cause of wildlife damage in their alfalfa fields deer F 858.5 128 df P 00070.007 than on monetary losses averaged 2479ha247924 79haha smaller farms 200 ha survey respondents the average dollar loss reported by respon- indicated that alfalfa production problems dif- dents who grew only irrigated alfalfa was 3016 fered by specific wildlife species table 1 n 86 ssj 554 respondents who grew both farm and ranch management irrigated and dryland alfalfa reported an aver- practice comparisons age loss of 4388 n 21 sjrajrs 1525 those who ranked a scale of 0 5 0 grew only dryland alfalfa reported an average respondents on no cost through 5 high cost the relative cost loss of 3750 n 2 ssy 250 follows the highest losses berhaperha were reported by of the 7 farm management practices as respondents who grew both irrigated and dry- irrigation 38383.8 fertilization 34343.4 weed control gg land alfalfa 42 ha respondents who grew 29292.9 insect control 96262.6 fencing 23232.3 big only irrigated or dryland alfalfa reported losses game control 20202.0 and rodentrabbitrodent rabbit control berhaperha of 19 and 28 respectively lgig191.9 fertilization weed control and irrigation growers with irrigated alfalfa farms 200 were used on 82 81 and 80 of the farms ha in size reported significantly higher mone- respectively big game and rodentrabbitrodent rabbit con- tary losses than operations 200 ha in size F trol were used by 71 and 38 of the respon- 15515.515 5 1103 df P 00010.0010 001 although the dents respectively respondents also reported average monetary loss reported by larger alfalfa employing several techniques to control wildlife farms was 5078 n 50 compared to 1639 damage in alfalfa fields table 2 based on for smaller farms n 55 the average loss sizes and types of alfalfa operations the only berhaperha was higher on smaller 37 than larger significant cost differences reported by man- farms 21 F 24924.924949 9 1103 df P 00010.0010 ooiool001 agement practices were for irrigation on farms growers reported no significant difference in 200 ha F 505.0so 1124 df P 0030.03 199619961 WILDLIFE DAMAGE TO ALFALFA 257

TABLE 1 percentage of all respondents N 150 reporting problems caused by a specific wildlife species in utah alfalfa fields in 1993 and a breakdown of that percentage into s tibLib categories based on the most severe type of problem caused

percentagepercenpereen tage identifying a specifics lucificlecific problem as mostme st severe wildlife species reporting hay hay equipment increased causing damage problems quality quantity damage costs pocket gophers 687 140 207 260 80 ground squirrel 333 40 107 153 33 voles 107 27 67 13 00 Jackjackrabbitsrabbits 328 27 287 07 07 prairie dogs 233 07 80 133 13 elk 200 60 127 13 00 mule deer 640 87 540 13 00 antelope 93 13 80 00 00 waterfowl 173 27 147 00 00

wildlife damage management agencies are unable to perform this work on a assistance programs regular basis our experience suggests that wildlife agencies should consider using state fourteen respondents 9 reported receiv- agriculture department hay lists to conduct ing compensation for wildlife damage in their benchmark surveys to identify wildlife damage alfalfa fields of these 12 received compensa- for damage caused by mule deer another management concerns and needs most states tion parker 48 31 indicated they received some type of maintain hay lists R personal commu- technical assistance to control wildlife damage nicationni UDA 1995 most of this assistance 75 was provided to our results summarize perceived losses control damage caused by mule deer the relationship between perceived and actual one hundred twenty two respondents 80 losses is unclear and probably difficult to esti- reported seeking either information or assis- mate conover 1994 this relationship depends tance in dealing with wildlife depredation prob- in part on how conspicuous the damage appears lems conservation officers were cited by 53 and which wildlife species causes the damage growers 43 as being their primary contact wakeley and mitchell 1981 decker et al 1984 for information or assistance county agents and melsormelvormcivor and conover 1994b UDWR biologists ranked and2nd 22 and ard3rd most respondents reported problems with 18 respectively other sources of informa- pocket gophers and mule deer other species tion in order of decreasing importance were commonly causing problems included jackrab other landowners 7 farm and ranch stores bits ground squirrels prairie dogs waterfowl 5 and UDA agricultural representatives 3 elk pronghorn and meadow voles conover respondents preferred compensation and 1994 also found that these species in particu- incentive programs 42 to other types of pro- lar deer were perceived to cause most damage grams to manage damage caused by wildlife in to agricultural crops in the US alfalfa fields research 17 field demonstra- based on statewide averages in 1993 utah tions 13 workshops 13 facts sheets 13 alfalfa growers harvested 10510.5losios tonshakonsha with a and videos 14 were rated nearly equal in market value of tigg716671.66 a ton survey respon- usefulness dents produced 177104 tons of alfalfa on 16867 ha having a total value of 12691000 discussion the 350000 loss reported due to wildlife represents 282.8 of the crop value expanding this perceived relationship of damage to the total value of alfalfa produced in utah costs to wildlife management during 1993 results in a total perceived loss of surveys can be cost effective means of assess- 44444.4 million this is 9 times the amount the ing the magnitude and economic impact of wild- utah state legislature annually appropriates life depredation crabb et al 1986 unfortu- 500000 to reimburse crop owner depreda- nately due to the cost and time associated with tion claims and expenses chapter 307 utah conducting reliable surveys many wildlife code 1994b 258 GREAT BASIN naturalist volume 56

TABLETABLL 2 percentage of all respondents N 150 using a specific technique to control damage caused by wildlife species in utah alfalfa fields in 1993 and a breakdown of that percentage into subcategoriessubcategonessub categories based on the most effective technique used

percentageperce niagentage identifyingiderlderitifying a specificspeelflefiefic techniqueie as beingbeibeling most effectiveeffectivelve using wildlife speciessr ecieseclesdecies damage controlconitiolithol shooting poison causing damagedamage techniques trapping hunting baits fumigantsfurnigantsFumiFurni gants cultural fencing hazing pocket gopherxoi901goi heihel 417 67 00 330 20 00 00 00 gigroundaundound squirrelsqaq un i el 454 47 173 220 00 07 00 07 voles 133 20 27 73 00 13 00 00 jackjaekjackrabbitsjackijackl abbitrabbits s 393 00 360 27 00 00 07 00 prairiepi an ie dodogs 240 20 127 73 13 00 07 00 elk 213 00 120 00 07 00 73 13 mule deeldeer 467 00 227 00 07 00 160 73 antelope 97 00 40 00 07 00 27 13 watelwaterfowlfowlrowl 167 00 133 07 07 00 00 20

utah code authorizes the UDWR to imme-imme life damage has increased in the last 30 yr our diately pay any approved damage claims survey results reinforce conover and decker s 5 500 claims or total amounts of claims sub- 1991 suggestion that programs necessary to mitted by a claimant in the fiscal year that are adequately address crop owner concerns have 500 are not paid until the total amount of not yet been implemented approved claims for the fiscal year is deter- role of state agencies in resolving mined if the amount claimed exceeds the wildlife damage management concerns appropriation the per claimant amounts paid in excess of 500 are prorated the current state wildlife management agencies are appropriation falls short of satisfying wildlife responsible for managing damage caused by damage compensation claims and expenses big game upland game and waterfowl mus- R valentine personal communication UDWR grave and stein 1993 state agriculture de- 1996 partmentspartments administer and enforce pesticide if 13 of utah alfalfa growers n 1000 control legislation that regulates the safe and submitted approved claims of 500 their proper use of pesticides for vertebrate pest claims would deplete the annual appropriation damage because of this role agriculture depart- although the alfalfa growers we surveyed pre- ments have jurisdiction over the control of ferred compensation and incentive payments unprotected wildlife species vertebrate pests over other types of wildlife damage manage- in utah these include pocket gophers field ment programs only 9 had ever received any mice muskrats ground squirrels jackrabbitsjackrabbits financial support raccoons skunks red fox and coyotes in the united states 2122.11 million farmers the UDWR recognizes that private lands control 400 million ha of our 937 million ha within utah provide habitat for wildlife and land base their actions largely influence the that under some circumstances wildlife may quality and quantity of the existing wildlife cause economic losses to the landowner with habitat base gerard 1995 landowners per- this understanding the UDWR cooperates with ceptions and concerns about wildlife damage the UDA and the US department of agricul- are important because they influence their atti- ture animal plant health and inspection ser tudes and behavior toward wildlife conover viceanimalvice Animal damage control ADC program 1994 suggested that wildlife damage has to conduct predator bird and rodent control reached levels that discourage private land- activities and compensate landowners for cer- owners from managing for wildlife on their tain losses caused by wildlife using funds property our results suggest that utah alfalfa appropriated by the legislature growers also peicpeiceiveperceiveeiveelve wildlife damage in alfalfa in 1994 the utah legislature enacted an fields as a serious concern although wildlife alternative compensation program that allows professionals working for federal or state wild- landowners to receive permits to harvest antler- life and agricultural agencies believe that wild less animals as mitigation for damage caused 199611996 WILDLIFE DAMAGE TO ALFALFA 259 by big game chapter 176 utah code 1994a sate smaller nonparticipating landowners adja- in 1995 the UDWR southern region issued cent to their operation for crop damage caused 1200 mitigation permits of which 50 were by big game animals filled in 1996 both the number of tags issued wildlife and number of animals harvested declined as the habitat authorization program landowners lost interest in the program N mckee personal communication UDWR the WHA program requires persons 14 yr 1996 of age or older to purchase a wildlife habitat to better address landowners concerns authorization prior to purchasing certain hunt- given fiscal and legal constraints we suggest ing or fishing licenses or permits the funds that agencies and organizations responsible for generated from this authorization are placed managing wildlife resources and wildlife dam- into a restricted account to be used for wildlife age on utah agricultural lands collaborate to habitat improvements several other great basin develop strategies that allow profitable agricul- states operate similar programs designed to ture and wildlife to coexist utah s posted hunt- generate funds to do habitat work ing unit PHU chapter 288 utah code 1993b we recommend that state wildlife agencies and wildlife habitat authorization WHA pro- consider using habitat funds to implement and grams chapter 75 utah code 1995 may offer evaluate enhancement projects and programs additional mechanisms to achieve this goal on public and private land that are designed specifically the posted hunting unit program to reduce big game depredation on private land habitat funds could be used to es- the UDWR also recognizes that wildlife can tablish big game lure crops situate interceptor be a significant benefit to the landowner the strips or modify migration corridors as a means PHU program provides landowners with mon- of abating localized depredation problems etary incentives through an allocation of hunting permits to include wildlife small game wildlife damage education needs waterfowl and big game in farm and ranch crop owners also need additional informa- management plans landowners who partici- tion on techniques used to manage wildlife pate in the program are required to improve damage several respondents reported using wildlife habitat but are ineligible to receive fumigantsfumi gants and poison baits to control damage compensation for crop losses caused by wildlife caused by ungulaungulatesungulatedtes lagomorphslagomorpha and birds the most successful of utah s PHU programs these practices are illegal as no products are involves big game animals in 1994 47 big currently registered in the US to control dam- game PHU programs encompassing over age caused by these species 400000 ha of private land provided additional we recommend that state wildlife agencies economic returns for hundreds of landowners agriculture departments and federal ADC pro- and hunting experiences for thousands of grams cooperate in the development of public hunters current program guidelines limit par- outreach extension education and research ticipation to landowners or landowner groups activities intended to inform crop owners about who own at least 4000 ha chapter 288 utah techniques that can be used to manage wildlife code 1993b the size limitation was estab- damage these programs also should provide lished to create more manageable herd units information on conservation technologies non in our survey respondents reported that big lethal strategies and opportunities that can be game animals caused the greatest damage we used to control wildlife damage and benefit suggest that big game PHU guidelines be wildlife resources while maintaining or enhanc- modified to accommodate farm or ranch units ing agricultural profitability 4000 ha in size this modification would pro- in conclusion previous studies conducted vide the stimulus necessary to alleviate many in the great basin focused on evaluating the crop owners wildlife damage concerns and effects of big game depredation tebaldi and provide an additional incentive to include wild- anderson 1982 austin and urness 1987a life in farm and ranch management plans in 1987b 1989 1993 and sandhill cranes melvormcivor addition we suggest that big game PHU oper- and conover 1994b on agricultural production ators be encouraged to incorporate provisions our study adds to this research by providing in their wildlife management plans to campencompen important insights regarding crop owners 260 GREAT BASIN naturalist volume 56 perceptions about wildlife damage and their science report 4 US department of interior needs and preferences in managing damage national biological service washington DC 28 appp our results suggest that alfalfa grow- GNEITING D J 1994 utah agriculturalagncultural statistics utah utah agricultural statistics service and the utah depart- ers perceive wildlife damage as a serious con- ment ofagricultureofagriculture salt lake city 138 appp cern this concern should be shared by wild- LEWIS S R AND J M OBRIEN 1990 survey of rodent life managers and rabbit damage to alfalfa haybay in nevada pages in addition to informing landowners of their 166 117 in proceedings of the 14th vertebrate pest conference university of california 320 concern over wildlife damage wildlife davis appp man- LUCE D G R M CASE AND J L stubbendieck 1981 agers should demonstrate it by addressing damage to alfalfa fields by plains pocket gophers potentials for increasing damage on private journal of wildlife management 45 258 260 lands when developing wildlife habitat man- MCIVOR D E AND M R CONOVER 1994a perceptions of farmers agement plans conover 1994 wildlife man- and non farmers toward management ofprob- lem wildlife wildlife society bulletin 22 211 219 also agers should incorporate strategies in man- 1994b impact of greater sandhill cranes faragforagfoimol ag- agement plans to benefit wildlife and reduce ing on corn and barley crops agriculture ecosys- depredation potentials on private land tems and environment 49 233 237 MUSGRAVEMUSGBAVE R S AND M A STEIN 1993 state wildlife laws handbook center for wildlife law institute of public acknowledgments law university of new mexico albuquerque 840 appp NATIONAL agricultural statistics SERVICE 1995 1994 we acknowledge R parker for assistance in crop statistics US department of agriculture wash- distributing the survey we thank M conover ington DC S barras and A hall for reviewing earlier PACKAM C J AND G CONNOLLY 1992 control methods research for animal damage pages drafts ofthisorthisof this priorities control manuscript 12 16 in proceedings of the 15th vertebrate pest conference university of california davis 300 appp literature CITED PIPER S E 1909 the nevada mouse plague of 1907 08 farmers bulletin 352 1 23 AUSTIN D A AND P J URNESS 1987a consumption of RENNISON B D AND A P BUCKLE 1988 methods for fresh alfalfa hay by mule deer and elk great basin estimating the losses caused in rice and other crops naturalistnatuialist4747 100 102 by rodents pages 69 80 in rodent pest management 1987 guidelines for evaluating annual crop CRC press inc boca raton FL 238 appp losses due to depredating big game utah division of SAUER W C 1978 control of the oregon ground squirrel wildlife resources publication 87587 5 42 appp pages 99 109 in proceedings of the ath7th vertebrate 1989 evaluating production losses from mule pest conference university of california davis 323 deer depredation in apple orchards wildlife society PP bulletin 1716117 igiigl161 165 statistical PROGRAMS FOR SOCIAL SCIENCES 1995 micro- 1993 evaluating production losses from mule deer soft windows release 50 microsoft corporation depiedepredepredationdation in alfalfa fields wildlife society bulletin redmond WA 2139721 397 401 TEBALDI A AND C C ANDERSON 1982 effects of deer CONOVER M R 1994 perceptions ofgrassof grassrootsgrass roots leaders of use on winter wheat and alfalfa production wyoming the agiagriculturalicultural community about wildlife damage on fish and game department job final report FW 3 their farmsearms and ranches wildlife society bulletin 22 R 26 78 appp 94 100 UTAH CODE 1971 chapter 46 section 2317523 17 5 damages CONOVER M R AND D J DECKER 1991 wildlife dam- for destroyed crops limitations appeal age to crops perceptions of agricultural and wildlife 1993a chapter 183 section 2316323 16316 3 damage to professionals in 1957 and 1987 wildlife society bul- cultivated crops by big game animals notice to letin 19 46 52 division crop owner authorized to kill animals CRABB A C T P SALMON AND R E MARSH 1986 surveys 1993b chapter 288 section 2323123 23 1 posted hunt- as an approach to gathering animal damage informa- ing units tion pages 2 4 in vertebrate pest control and man- 1994a chapter 176 section 231632316352323163.516 353 5 damages agement materials americanamerlean society for testing and to livestock forage fences or irrigation equipment on materials STP 974 philadelphia PA 12 appp private land DECKERDEC KER D J G F MATTFIELD AND T L BROWN 1984 1994b chapter 307 section 2316423 16 4 damages for influence of deer damage on farmers perceptions of destroyed crops limitations appraisal deer population trends important implications for 1995 chapter 75 section 23194223 19 42 wildlife habi- manamanagersgeisgels proceedings of the first eastern wildlife tat authorization damage control conference 1 191 195 WAKELEY J S AND R C MITCHELL 1981 blackbird dam- DUNN J P J A CHAPMAN AND R E MARSH 1982 jack age to ripening field corn in pennsylvania wildlife rabbits pages 124 145 in wild mammals of north society bulletin 9 52 55 america biology management and economics john hopkins university press baltimore MD 1147 appp received 11 september 1995 GERARD P W 1995 agricultural practices farm policy and accepted 10 april 1996 the conservation of biological diversity biological great basin naturalist 563 0 1996 appp 261 266

SPATIAL relationships AMONG YOUNG cercocarpc7sCERCOcercocarpusCARPUS ledifoliusLEDIFOLIUS CURLLEAF MOUNTAIN MAHOGANY

1 paul 3 brad W schuitzSchultzschultzlschultz1schultzel robin J tauschetausch2 1 and T Tueller

ABSTRACT this study analyzed spatial location patterns of Cercocercocarpuscarpus ledifohusledifoliuslediledlfolius nutt curlleafcurlleancurlleaf mountain mahogany plants classified as current year seedling established seedling juvenile and immature individuals at a central nevada study site most current year seedlings were located in mahogany stands in which large mature individuals hadbad the greatest abundance these stands had greater litter covelcover and a thicker layerlayel of litter than areas with few cur rent year seedlings most established young Cercocercocarpuscarpus were located in adjacent artemisia ttltntrltridentata sspasp vastyanavasevaseyanayana mountain big sagebrush communities or in infrequent canopy gaps between relatively few large mature Cercocercocarpuscarpus we discuss potential roles of plant litter root growth characteristics nurse plants and herbivoreherbherbivoryivory in the establishment and renewal of Cercocercocarpuscarpus communities

key words Cercocercocarpuscarpus litter mountain mahogany seedling recruitment spatial relationships maturity class

Cercocercocarpuscarpus ledifoliusledifolius nutt curlleafcurlleancurlleaf moun- 1975 rapid root growth should benefit cerco tain mahogany hereafter Cercocercocarpuscarpus is a carpus seedlings in the great basin where a desirable browse species in the intermountain semiarid climate predominates previous stud- west smith 1950 smith and hubbard 1954 ies indicate land managers require additional hoskins and dalke 1955 attempts to revege- information about 2 processes in Cercocercocarpuscarpus tate wildlife habitat with Cercocercocarpuscarpus have had communities 1 the dynamics of current year little success common problems have been Cercocercocarpuscarpus seedlings in relationship to the competition from annual weeds holmgren rest of the vegetative community and 2 con- 1954 sensitivity to frost and drought plum- ditions that permit current year seedlings and mer et al 1957 1968 slow growth plummer established young Cercocercocarpuscarpus to be recruited et al 1957 and impaired germination liacos into the population structure and nord 1961 young et al 1978 schultz et al 1991 presented the first pre- Cercocercocarpuscarpus does not sprout from root dictive relationships about the structure of crowns following removal of the canopy Cercocercocarpuscarpus stands their study in western and ormiston 1978 austin and urness 1980 central nevada found that mean Cercocercocarpuscarpus reproduction must occur from seed limited crown volume had a significant FP 0050.05 in- research has addressed the structure of cerco verse relationship rar2 0780.78 with density of carpus stands scheldt 1969 duncan 1975 Cercocercocarpuscarpus in established seedling juvenile davis 1976 davis and brotherson 1991 or and immature maturity classes schultz 1987 how stand structure may influence regenera- also found that Cercocercocarpuscarpus canopy cover and tion except for duneanduncan s 1975 work in mon- mean Cercocercocarpuscarpus crown volume had signifi- tana past studies concluded that most stands cant P 0050.05oos positive correlations with den- have few young Cercocercocarpuscarpus and that older sity of current year Cercocercocarpuscarpus seedlings this individuals have the greatest abundance these dichotomy along with other patterns observed studies scheldt 1969 duncan 1975 davis by schultz 1987 may offer valuable insight 1976 davis and brotherson 1991 also found into the regeneration of Cercocercocarpuscarpus stands few seedlings low seedling survival and irreg- additionally schultz 1987 observed that 1 lo- ular seed production plummer et al 1968 cations with large canopy gaps between widely the few current year Cercocercocarpusgercocarpuscarpus seedlings scattered mature individuals generally had that emerge apparently have rapid elongation more Cercocercocarpuscarpus in established seedling of their taproot 0970970.97 in after 120 days dealy juvenile and immature maturity classes than

ibiologicalabiologicalbiological sciences centergenter desert research institute university of nevada system box 60220 reno NV 89506 correspondingcongongoo expending authoauthor 2usdaUSDA forest service intermountainintel mountain forestfoiest and range experiment station reno NV 89512 313cpartmentdepartment of environmental and resource sciences university ofot nevada reno reno NV 89512

261 262 GREAT BASIN naturalist volume 56

did locations with small canopy gaps 2 loca- METHODS tions with small canopy gaps and hence greater Cercocercocarpuscarpus canopy cover and crown initial measurements describing the struc- volume hadbad a greater abundance of young ture of Cercocercocarpuscarpus stands occurred on the Cercocercocarpuscarpus in adjacent artemisia tridentata shoshone range and leavinepeavine mountain in sspasp vastyanavaseyanavaseyana mountain big sagebrush com- june and july 1985 relevant results are pre- munimunitiesties 3 established Cercocercocarpuscarpus in the sented in table 1 measurements describing artemisia community were often rooted under the spatial location of individuals in current the protective canopy of another shrub or year cercocarwCercocercocarpuscarpus seedling established seedling shrub skeleton and 4 most current year cer juvenile and immature maturity classes were carpuscocarpusco seedlings were found where thick made on the shoshone range in early august plant litter had accumulated under mature 1985 abundant rainfall in central nevada dur- Cercocercocarpuscarpus table I1 summarizes differences ing june and july allowed current year cerco patterns in Cercocercocarpuscarpus stand structure from carpus seedlings to survive until we initiated locations in western leavinePepeavineavine mountain and this study similar data could not be collected central shoshone range nevada table 2 de- from leavinepeavine mountain in western nevada fines the maturity classes mentioned through- because a dry spring and summer resulted in out this study the early desiccation and disappearance of cercocarpus based on observations about the spatial most Cercocarpus seedlings seven 1 location of current year Cercocercocarpuscarpus seedlings I X 40 m belt transects BT were located at 4 of the 13 Cercocercocarpus and established Cercocercocarpuscarpus in the youngest carpus stands in the shoshone range maturity classes we implemented a brief de- measured by schultz et al 1990 1991 none of the BTs placed scriptive study on the shoshone range in cen- were in study plots sampled by al tral nevada to quantify the spatial distribution schultz et 1990 1991 also described in schultz 1987 of current year Cercocercocarpuscarpus seedlings and cer because those study plots were located the cocarpus in established seedling juvenile and in interior of the stands not near the with the adja- immature maturity classes we integrate data ecotone cent artemisia community 4 stands sam- from this study the schultz et al 1990 1991 the pled were selected because 1 they were near studies about stand structure which were con- access roads and time was limited and 2 their ducted at the same location as this study and respective topographic positions allowed at other relevant literature to describe possible least I1 transect of the 7 to be located at each processes mechanisms factors influ- or that cardinal aspect ence survival of current year cercocamwCercocercocarpuscarpuscarpas seed- the following criteria were used to select lings and their subsequent recruitment into transect locations 1 a Cercocercocarpuscarpus stand dom- established seedling juvenile and immature inated by mature individuals was present 2 a maturity classes our goal is to stimulate thought sharp ecotone existed between the Cercocercocarpuscarpus that can guide research about the regeneration stand and adjacent artemisia community 3 of this desired browse species the transect remained on the same landfordlandformlandform

TABLE 1 mean values for structural characteristics of Cercocercocarpuscarpus communities from 2 mountain ranges in western and central nevada data from schultz 1987 schultz et al 1990 mean values in the same column followed by the same letter are not significantly different P 005oos0 05

established seedling Cercocercocarpuscarpus mountain current year immature and mature crown Cercocercocarpuscarpus litter bare range seedlings juvenile Cercocercocarpuscarpus volume cover cover groundlground1groundly m 2 ha ha mpmplantimplantm3plantlant 1 leavinepeavine olaoia0 la 922a 233a 58a 56a 67a loaioa shoshone lgb lllbellb111b 344b 395b39 ab5b 79b 76b loaioa includeslincludes graddgravdyiavel 199611996 CERCOcercocarpusCARPUS regeneration 263

TABLE 2 Cercocercocarpmcercocarpuscarpus maturity classes descriptions were TABLE 3 elevation slope and aspect of each belt tran- developed from a reconnaissance of Cercocercocarpuscarpus stands sect in which count data were obtained nealnear reno NV elevation slope aspect current year germinated during the current growing transect Wm N degrees seedling season usually has 4 leaves 1 2688 41 80 established plants I1 year of age 2 7 hunnunmm basal 2 2688 41 80 seedling diameter smoothsmouth bark may be up to 3 2688 41 80 30 cm tall 8 or u e leaves 4 2400 29 290 5 2758 34 0 juvenile young plants 7 mm basal diameter 6 2758 34 0 smooth bark plants to 60 cm tall 7 2758 25 168

immature young plants 1251.251 25 cm basal diameter smooth bark plants to 151 5 rn tall cocarpus community or the adjacent artemisia young mature cracked bark 151.5lsis1 5 303.03 0 rn tall crown community and 2 the number of established broadened may be multimultistemmedstemmed from seedling juvenile and immature Cercocercocarpuscarpus base not suppressed by adjacent larger rooted under and not under the canopy of a mountain mahogany plants live or dead shrub the wilcoxon signed rank mature cracked bark wide full crown few dead test was used to determinedeterwineermine if there was a sig- branches may have several stems from nificantnificant difference in the distribution of indi- base 3 m tall viduals in the Cercocercocarpuscarpus and artemisia com- munitiesmunities respectively for each maturity class overmatureover mature cracked bark may be multistemmedmultisternmedmultistemmed numerous dead branches may be 3 m the significance level is P 005oos0.05 unless other- tall frequently suppressed by adjacent wise noted larger mountain mahogany plants RESULTS and had the same aspect throughout its length current year Cercocercocarpuscarpus seedlings were not and 4 all transects located in the same stand distributed evenly between Cercocercocarpuscarpus stands were 40 in or more apart table 3 describes the and adjacent artemisia communities table 4 elevation slope and aspect of each transect significantly more current year seedlings were Cercocercocarpuscarpus in the shoshone range are largely rooted in the Cercocercocarpuscarpus community restricted to the foxmountFoxmount soil series carol jett at least 81 of established seedling juve- personal communication which is a gravelly nile and immature Cercocercocarpuscarpus were rooted loam specifically a loamy skeletal mixed topic in the adjacent artemisia community table 4 cryboroll this soil is well drained and moder- for established seedling and juvenile maturity ately permeable depth to a paraparalithiclithic contact classes the difference in spatial distribution was averages 60 100 cm significant the significance level for immature all transects were located such that 20 in Cercocercocarpuscarpus was P 0060.06 occurred in the Cercocercocarpuscarpus stand and 20 in in more established seedling juvenile and the adjacent artemisia community each tran- immature Cercocercocarpuscarpus were rooted under the sect was divided into forty I1 X 1 m quadratsquadquadransrats protective canopy of a live or dead shrub than every Cercocercocarpuscarpus rooted in each quadrat was in the open table 5 only I1 transect had more classified by maturity class for Cercocercocarpuscarpus inm plants without a protective canopy but the sig- established seedling juvenile and immature nificancenificance level was FP 00100.10olo10 maturity classes we determined whether the plant was rooted under the protective canopy discussion of a live or dead shrub distribution of current year seedling estab- spatial distribution of current year cerco lished seedling juvenile and immature cerco carpus seedlings and established young cerco carpus was summarized for 10 classification carpus had an inverse relationship tables 1 categories populations these were 1 the 4 current year seedlings were most abundant number of Cercocercocarpuscarpus in current year seed- in Cercocercocarpuscarpus stands dominated by large ma- ling established seedling juvenile and immature Cercocercocarpuscarpus and least abundant in adja- ture maturity classes rooted in either the cer cent artemisia communities young established 264 GREAT BASIN naturalist volume 56

rableFABLETABLE 4 number ofofcunentcurrent searyear seedling established seedling juvenile and immature mahogany rooted in cercocar pus CER stands dominated by mature individuals and in adjacent artemisia ART communities within each maturity class total values between community types with different letters are significantly different P 0050 05 currentcurrengurrenit year estahestablishedlisheddished seedseedlingling seedseedlinghaghmg juvenilejuve nile immatureimmatureacture transect CER ART CER ART CER ART CER ART

1 20 0 1 11 1 5 0 1 2 72 15 1 15 3 3 0 5 3 75 53 0 16 0 6 5 6 4 31 39 0 2 0 7 0 4 5 337 25 0 11 0 19 0 0 6 506 28 1 11 0 4 0 0 7 33 0 1 9 0 2 0 5 total 1074a 160b 4aaa 75b 4aaa 46b 5aaa 21al pelpeipercentcent 87 13 5 95 8 92 19 81

isilsigiflSisiriiifiltntlydifitgifgil eiiitlydiffentittplent it P 006

Cercocercocarpuscarpus were virtually absent from mature 1 presence or thickness of plant litter 2 root Cercocercocarpuscarpus stands but had a greater abun- growth characteristics 3 presence of nurse dance in adjacent alreaireATteattenisiaarteimsiaattemisiamisiamisla communities tables plants and 4 herbivoreherbherbivoryivory 1 4 young Cercocercocarpuscarpus were also abundant in moderate levels of litter can favor seed ger- stands with low Cercocercocarpuscarpus crown cover or minationmi and seedling establishment by de- relatively few large Cercocercocarpuscarpus table 1 the creasing soil temperature and increasing soil low density of current year seedlings in adja- moisture evans and young 1970 thick litter cent artemisia communities table 4 has 2 however can reduce seedling establishment and possible interpretations 1 viable Cercocercocarpuscarpus survival by preventing or restricting contact seeds were not dispersed into the artemisia between soil and seed or soil and root fowler community or 2 germination of Cercocercocarpuscarpus 1986 seed was impaired because data about seed high litter cover table 1 and a thick layer densities are lacking a definitive conclusion of litter personal observation were common cannot be made Cercocercocarpuscarpus seed however is in Cercocercocarpuscarpus stands in the shoshone range primarily wind dispersed USDA 1948 there- litter cover and litter thickness were not mea- fore it is unlikely that few seeds were present sured in adjacent artemisia communities how- in the artemisia community particularly since ever litter cover in high elevation 2200 m all data were collected within 20 m of the cer artemisia communities ranges from 15 to cocarpus stands most likely over 85 fewer 50 tueller and eckert 1987 extensive and Cercocercocarpuscarpus seedlings were in the artemisiaartermsia deep litter in Cercocercocarpuscarpus stands may promote community table 4 because seed germination seed germination but decrease seedling sur- was substantially lower than in the cercocar vival because roots from Cercocercocarpuscarpus seedlings pus stands seldom make contact with the mineral soil the inverse relationship for distribution of less litter in the artemisia community may re- current year seedlings and established young duce Cercocercocarpuscarpus seed germination but enhance Cercocercocarpuscarpus indicates that locations with a survival of seeds that germinate root growth high abundance of current year seedlings are characteristics may play an important role not necessarily locations with the best seedling rapid root growth that current year cerco survival populations perpetuate when seedlings carpus seedlings experience dealy 1975 should survive and advance into successively older enhance survivorship of Cercocercocarpuscarpus seedlings maturity classes eventually producing new during seasonal drought a common phenome- seedlings the pattern for spatial distribution non in the great basin root systems that of current year seedling established seedling undergo rapid elongation should be able to fol- juvenile and immature Cercocercocarpuscarpus derived low a retreating zone of soil moisture down- from this study and that conducted by schultz ward better than root systems that elongate et al 1990 1991 indicates that 4 factors may slowly we excavated several Cercocercocarpuscarpus seed- influence survival of current year seedlings as lings rooted in thick plant litter and found that well as plants in the youngest maturity classes root growth was extensive 20 cm but not 199619961 CERCOcercocarpusCARPUS regeneration 265

TABLE 5 the number of established seedling juvenile lishmentlishment of woody species marquis 1974 and immature Cercocercocarpuscarpus rooted under and not under mcauliffe 1986 including Cercocercocarpuscarpus another shrub or shrub skeleton significance level is P scheldt 010 and tisdale 1970 the presence of protective nurse plants therefore may be for transect rooted under not rooted under important regeneration of Cercocercocarpuscarpus seedlings 1 16 3 Cercocercocarpuscarpus stands in the shoshone range 2 23 4 had a mean shrub canopy cover of 11 3 20 13 schultz al 4 8 5 et 1990 total shrub canopy cover was not 5 6 24 measured in adjacent artemisia communities 6 9 7 however it generally ranges from 41 to 50 7 15 2 tueller and eckert 1987 total 97a 58b thus shrub cover percentage 63 37 in adjacent artemisia communities is 353.53 5 to 4 times greater than that in Cercocercocarpuscarpus stands since more established seedling juvenile and immature Cercocercocarpuscarpus were rooted under a downward toward or into the mineral soil shrub or shrub skeleton than not table 5 the root growth was largely lateral following ger- difference in shrub canopy cover between cer minationmination in early spring available moisture in cocarpus stands and adjacent artemisia com- both mineral soil and plant litter is probably munitiesmunities may influence survival of current year high since cool temperatures and abundant seedlings established seedlings juvenile and precipitation are common houghton et al immature Cercocercocarpuscarpus artemisia and other 1975 because moisture is not limiting early in short statured shrubs may serve as nurse plants the growing season root growth probably fol- and protect small Cercocercocarpuscarpus including cur lows the path of least resistance when thick rent year seedlings from herbherbivoresivores until their litter resides on top of mineral soil the path of photosynthetic surface is large enough to cope least resistance would be laterally through the with frequent browsing since shrub cover is litter not downward through the mineral soil low in Cercocercocarpuscarpus stands more young cerco loamy soil cercocarpus the that Cercocarpus stands inhabit carpus are probably exposed to herbherbivoresivores than undoubtedly stores and retains more water in aileAtleariemmaatlemisiaartemmamisiamisla communities this may help explain than plant litter does and thus should desic- the near absence of young Cercocercocarpuscarpus in cer slowly cate more if thick plant litter prevents cocarpus stands and their greater abundance in regards in or retards roots of current year Cercocercocarpuscarpus adjacent artemisia communities seedlings from reaching or penetrating moist mineral soil seedling mortality should be high conclusions when litter desiccates rapidly later in the summer we observed high mortality for current abundance of current yeaiyealyear Cercocercocarpuscarpus year Cercocercocarpuscarpus seedlings in august in cerco seedlings is greatest in Cercocercocarpuscarpus stands that carpus stands with thick accumulations of lit- have high Cercocercocarpuscarpus canopy cover large mean ter less litter on leavinepeavine mountain table 1 Cercocercocarpuscarpus crown volume and an extensive and in the artemisia community see tueller layer of plant litter these stand attributes also and eckert 1987 may enable root systems of result in a low density of plants in established Cercocercocarpuscarpus seedlings at these locations to seedling juvenile and immature maturity grow downward into mineral soil immediately classes established young Cercocercocarpuscarpus are following germination this should increase most abundant where gaps occur in the cerco survivorship of current year seedlings which carpus canopy or in adjacent artemisia com- may account at least partially for the greater munimunitiesties survival of current year seedlings abundance of established seedling juvenile appears best at locations that permit roots of and immature Cercocercocarpuscarpus on sites with less seedlings to make contact with mineral soil surface litter survival of current year seedlings and progres- herbivoreHerbherbivoryivory may also play a role in seedling sion of individuals hornfromhormhomm established seedling survival current year Cercocercocarpuscarpus seedlings maturity class into successively older maturity have an average leaf surface area of only 4 cm2 classes appear to be enhanced by the presence dealy 1975 which herbivoresherbivores can easily con- of a shrub canopy that protects small cercocar sume herbivoreHerbherbivoryivory can adversely affect estabbestab pus from herbherbivoresivores 266 GREAT BASIN naturalist volume 56

literature CITED PLUMMER A P R L GENSEN AND H D STAPLEY 1957 job completion report for game forage revegetation AUSIINAUSTIN D A AND PPJJ URNESS 1980 response of curlleafcurlleancurlleaf project W 82 11 2 utah state department of fish game mountain mahogany to pruning treatments in north- and ern utah journal of range management 33 275 277 PLUMMER A P D R christensen AND S B MONSEN DAVIS J N 1976 ecological investigations in Cercocercocarpuscarpus 1968 restoring big game range in utah utah divi- ledifohusledifoliuslediledlfolius nutt communities of utah unpublished sion of fish and game publication 69369 3 master s thesis brigham young university provo UT SCHELDT R S 1969 ecology and utilization of curl leaf DAVIS J N AND J D brotherson 1991 ecological rela- mountain mahogany in idaho unpublished master s tiontionshipsships of curlleafcurlcurlleanleaf mountain mahogany cercocar thesis university of idaho moscow pus ledifoliusledifohtislediledlfolius nutt communities in utah and impli- SCHELDT R S AND E W TISDALE 1970 ecology and uti- cations for management great basin naturalist 51 lizalizationtion of curlleafcurlcurlleanleaf mountain mahogany in idaho uni- 153 166 versity of idaho forest wildlife and range experi- DEALY J E 1975 ecology of curlleafcurlcurlleanleaf mountain mahogany ment station note 15 Cercocercocarpuscarpus ledifoliusledifohuslediledlfoliuskolluskolius nutt in eastern oregon and SCHULTZ B W 1987 ecology of curlcurlleafcurlleanleaf mountain adjacent areas unpublished doctoral dissertation mahogany Cercocercocarpuscarpus ledifoliusledzfoliuslediledlfolius in western and oregon state university corvallis central nevada population structure and dynamics DUNCAN E 1975 the ecology of curlleafcurlcurlleanleaf mountain mahog- unpublished master s thesis university of nevada any in southwestern montana with special reference reno iliiiilllpp111 appp to mule deer unpublished master s thesis montana SCHULTZ B W R J TAUSCH AND P T TUELLER 1991 state university bozeman size age and density relationships in curlleafcurlleancurlleaf moun- EVANS R A AND J A YOUNG 1970 plant litter and the tain mahogany Cercocercocarpuscarpus ledifoliusledifoltuslediledlfolius populations in establishment of alien annual weeds in rangeland western and central nevada competitive implica- communities weed science 18 697 703 tions great basin naturalist 51 183 191 FOWLER N L 1986 microlitemicrositeMicrosite requirements for germina- SCHULTZ B W PR T TUELLER AND R J TAUSCH 1990 tion and establishment of three grass species ameri- ecology of mountain mahogany Cercocercocarpuscarpus ledi can midland naturalist 115 131 145 folius in western and central nevada community HOLMGREN R C 1954 A comparison of browse species and population structure journal of range manage- for the revegetation of big game winter ranges of ment 43 13 20 southwestern idaho USDA intermountain forest SMITH A D 1950 feeding deer on browse species during and range experiment station research paper 33 winter journal of range management 3 130 132 HOSKINS L W AND P D DALKE 1955 winter browse on SMITH A D AND R L HUBBARD 1954 preference rat- the pocatello big game range in southeastern idaho ings for winter deer forages from northern utah journal of wildlife management 19 215 225 ranges based on browsing time and forage consumed HOUGHTON J G C M SAKAMOTO AND R 0 ciffordclffordGIFFORD journal of range management 7 262 265 1975 nevada s weather and climate nevada bureau TUELLER P T AND R E ECKERT 1987 big sagebrush of mines and geology special publication 2 artemisia tntridentata dasevastyanavaseyanavaseyandvaseyandyondyana and longleaf snow- LIACOS L G AND E C NORD 1961 Cercocercocarpuscarpus seed dor- berry symphoncarpossymphoricarpos oreoreophilusophilus plant associations mancy yields to acid and thioreathiourea journal of range in northeastern nevada great basin naturalist 47 management 14 317 320 117 131 MARQUIS D A 1974 the impact of deer browsing on USDA 1948 Cercocercocarpuscarpus H B K mountain mahogany allegheny hardwood vegetation USDA forest ser- pages 132 133 in woody plant seed manual USDA vicevlee research paper NE 308 miscellaneous publication 654 mcauliffe J R 1986 herbivore limited establishment YOUNG J A R A EVANS AND D L NEAL 1978 treat- of a sonoran desert tree cercidium microphyllum ment of curlleafcurlleancurlleaf cercocercocarpuscarpus seeds to enhance germi- ecology 67 276 280 nation journal ofwildlife management 42 614 620 ORMISTON J H 1978 response of curlleafcurlleancurlleaf mountain mahogany to top pruning in southwest montana pro- received 17 may 1995 ceedingsce of the first international range congress accepted 25 march 1996 denver CO great basin naturalist 563 C 1996 appp 267 271

POTENTIAL FOR controlling THE SPREAD OF CENTAUREA MACULOSA WITH GRASS competition

john L lindquistl2lmdquist12 bruce D Maxwellmaxwelll1 and T weaverlweavellweaver1

ABSTRACT spotted knapweed centaurea macumaculosemaculosalosa lam is a major rangeland and roadside weed of the northern rocky mountains it is often found in plant communities dominated by pseudoroegneriapseudoroegnena spicaspicatumtum or festuca idahoensis but it rarely invades roadsides dominated by bromus inermiainermis leyssdeyss aboveground biomass of the 3 grass species grown in mixture with centaurea was compared to growth in monoculture at a range ofofnitiogennitrogen input levels the results suggest that bromus is capable of suppressing the growth of centaurea with the degree of suppression increasing with increasing nitrogen levels the 2 native grasses had no impact on centaurea under the controlled environment conditions of this study

key words competition weed control centaurea macumaculosemaculosalosa bromus ermisinermisinermiainermis agropyron spicaspicatumtum festuca idahoenidahoan sis exotic plants

centaurea macumaculosamaculoselosa lam spotted knap- our objective was to measure the competi- weed is a major weed associated with spring tive ability of 3 grass species against centaurea wet summer dry areas of the northern rocky in 2 way interaction experiments in sand cul- mountains forcella and harvey 1981 tyser ture mixture and monoculture treatments and key 1988 weaver et al 1989 centaurea were tested for 12 wk at 5 positions on a nitro- dominates waste places invades disturbed gen gradient to determine whether competi- rangeland and sometimes invades undisturbed tive relations were influenced by differences in range tyser and key 1988 in contrast it rarely nitrogen availability A plant s ability to com- invades roadsides dominated by bromus inennisinermisinermia pete is related to its growth rate or ability to leyssdeyss weaver et al 1989 this suggests that gain biomass relative to associated species it may be excluded from waste places that are harper 1977 we compared aboveground bio- planted to bromus before centaurea invades mass of each species grown in mixture with alternatively because planting exoticsexotica violates centaurea to its growth in monoculture the charge of national park managers one may ask whether centaurea might also be excluded MATERIALS AND METHODS from disturbed areas by planting native grasses that naturally dominate either relatively dry the rhizomatous exotic pasture grass bromus pseudoroegneriapseudoroegnetla spicatumspicatum pursh scribner and inennisinermisinermia leyssdeyss and 2 native bunchbunchgrassesgrasses nor- smith agropyron spicatumspicatum or more moist mally dominating relatively dry foothills eshupieupseu festuca idahoensis elmer foothill habitats doroegneria spicatumspicatum or moister grasslands weed suppression may be accomplished by immediately above and below the conifer zone 1 preempting resources with more competi- festuca idahoensis were grown in 2 species tive plant species or 2 using biocontrols or mixtures replacement series with C macumaculosamaculoselosa herbicidesherbicides that selectively increase weed mor- experiments consisted ofaof3of 3 competition treat- tality decrease vigor or prevent reproduction ments monoculturesmonocultures of both grass and centau- lindquist et al 1995 this study considers rea and 5050 mixture combined with 5 nitro- management of centaurea macumaculosamaculoselosa by compe- gen addition treatments each treatment com- tition rather than by common herbicide and bination had 10 replicates within each experi- biocontrolbiocontrol methods this approach deserves ment pots were arranged in a completely ran- attention because it may be less expensive and domized design on a greenhouse bench and more effective than herbicherbicidesides in the long rotated weekly to minimize position effects term each experiment was subject to different light

department of biology and department of plant soil and environmental sciences montana state university bowmandowmanbozemandozemandomman MT 59717 present addressaddless department of agronomy university of nebraska lincoln NE 68583091568583 0915

267 268 GREAT BASIN naturalist volume 56

conditions because of its position in the green- calculated to determine whether competitive house A square planting pattern was used relationships varied across relative nitrogen with 4 plants spaced 5 cineincm apart in each pot in levels RCI is calculated as the mixture treatments plants of the same species were located on the diagonal RCI bonobanobpno13mono bmix0monoixmonoix mono 221 seeds were planted at a depth of 101.0iolo1 0 cm in cm3 1000 pots filled with coarse washed sand where bmonoamono and amixbmixB are the aboveground dry pots 1 were watered daily for 1 wk to allow biomass g plant for a species grown in mono- seedling establishment excess seedlings were culture and mixture respectively A negative thinned and remaining seedlings allowed to RCI value indicates that the species performs grow for an additional week prior to the addi- better in mixture than in monoculture RCI may tion of nutrients the basic nutrient solution be the best measure for determining species was balanced with respect to all essential nutri- displacement under competitive conditions ents but could be varied to allow the establish- across a resource gradient grace 1995 analy- ment of nitrogen levels from 0 1 10 sis of variance was used to test for differences 30 and 100 of a standard level machlis and in RCI within a species across nitrogen treat- torrey 1956 sufficient nutrient solution 200 ments student s t was used to compare RCI ml was applied to saturate the pot twice weekly between species at each nitrogen addition level and water 200 ml was added once each week regular watering with nutrient solution and RESULTS alternate washing with tap water held the soil solution near the applied level and prevented A hyperbolic relationship between individ- any concentration of the soil solution due to ual plant biomass and relative nitrogen level evapotranspiration experiments were conducted was found in all mixtures and monomonoculturescultures during march april and may 1988 when figs la f estimates of iiI1 biomass at inter- greenhouse temperatures ranged from 14014 to cept differed between mixtures and monoculmonocur 32 c25cmeanC 25 C mean tures only for centaurea grown in mixture with twelve weeks afterahter emergence plants in bromus table 1 estimates ofaof ai maximum each pot were clipped at the soil surface sepa- biomass differed for bromus and centaurea rated by species dried at 45 C for 5 d and table 1 weighed relative competition intensity was signifi- nonlinear regression procedures SAS 1988 cantly negative for bromus at all nitrogen addi- gauss newton least squares estimation method tion levels it varied from negative values at were used to fit a rectangular hyperbola equa- low nitrogen to positive values at higher nitro- tion 1 cousens 1985 to mixture and mono- gen levels for centaurea in competition with culture data for each species bromus fig 2 however RCI did not differ from zero in the experiments where P spica iiI1 N tum and FE idahoensis were in competition with B 1 centaurea data not shown iiL N 1 aiA discussion where B abovegroundaboveground dry biomass g growth responselesiesponse of bromus to nitrogen was plant 1 aiA maximum aboveground biomass greater in mixture with centaurea than in of species ti g plant 1 N relative nitrogen monoculture as indicated by the regression addition level and iiJ biomass of species i as lines fig la and the negative RCI values relative nitrogen addition level approaches zero across all nitrogen addition levels fig 2 in to determine the relative success of centau- contrast growth response of centaurea was rea in competition with each grass species lower in mixture with bromus than in mono- estimates ofaof ai and ii1 were compared between culture fig 1 the increase in centaurea RCI mixtures and monoculturesmonocultures using the extra at high relative nitrogen level indicates that sum of squares procedure ratkowsky 1983 bromus is a better competitor in the high lindquist et al 1996 in addition relative nitrogen treatments fig 2 results suggest competition intensity RCI grace 1995 was that bromus is capable of suppressing the 199619961 competitive suppression OF CENTAUREA 269

CIO 5 E 5 535253.53clo b ssW a CL Q 5 4 5 4

n cz C 3 E 3 0 0 co 2 jbCQ 2 w en s a2aa Q A 1 0 0 CQ 08 0.202 040.4 06 08 101.0to 00 020.2 040.4 06 080.8 10 0 00 02

151.5 ascu 151.5 C CL d cn 121.2 121.21 2

cd og 090.9osog E 090.9os 0 CO 060.6og 06

030.3 030.3

CL 000.0oo vf 000.0oo L og 06og 08 00oo 04 0.606 0.8ob08 101.0to 000.0oo 020.2 040.4 060.6 080.8 101.0loio 0 000.0 02 040.4 06 08

30 3 303.0 r z 303.0 f CL Ceg CL 5 252.5 252.5 W cn in wn on 202.0 czco 202.0U E E S 0 1.515 12.2 15 ez 15 CD a v to10 J 101.0iolo 101.0 8 8 C 050.5 i ii 05 J h i 1M 000.0oo S 00 tf og 00oo 02 04 0.606og 0.808 1.0to10 00 020.2 040.4 060.6 08 101.0to u 000.0 020.2 040.4 06 08 10 0 relative nitrogen addition level

1 fig 1 plot of observed 0 o and predicted 0 o aboveground dry biomass plantpiantpianti on relative nitrogen addition level when grown in monoculture and mixture a bromus grown in monoculture and in mixture with centaurea b centaurea grown in monoculture and in mixture with bromus c pseudoroegneriapseudoroegnena grown in monoculture and in mix-mix ture with centaurea d centaurea grown in monoculture and in mixture with pseudoroegnenapseudoroegneria e centaurea grown in monoculture and in mixturemixtuie with festuca f festuca grown in monoculture and in mixture with centaurea 270 GREAT BASIN naturalist volume 56

TABLETABLL 1 estimates of parameter values followed by asymptotic standard errors for maximum aboveground biomass g 1 plantpiantpianti A biomass as relative nitrogen level approaches zero 1I7 and the coefficient of determination rar2 obtained from fitting equation 111 to monoculture and mixture data of each species variation in I1 and A between competition treatments was tested using the extra sum of squares principle with P value indicating the significance level for the comparisoneomcompanson ofparameterof parameter goeff values from the monoculture and mixture regressions species coeffboeff competitionn treatment P value monoculture mixture brornus A 1 884018840 115 3819036138190.3613 8190 361 0000 I1 8 7691876918.7691 141 128932197128932.19712 8932 197 0283 r2ra 090 084 centaurea with A 1306008213060.0821 3060 082 043800560.43800560 4380 056 0000 bromus I1 10621186110 6211 861 4571735474571735.4745 71735 47 0023 r2ra 085 014 pseudoroegneriapseudoroegnena A 1289023712890.2371 2890 237 0827008708270.0870 8270 087 0081 I1 1758032317580.3231 7580 323 249004422.49004422 4900 442 0410 r2ra 080 082 centaurea with A 0636004806360.0480 6360 048 0805010708050.1070 8050 107 0307 pseudoroegneriapseudoroegnena I1 6770l639677016396 7701 639 4617l454461714544 6171 454 0665 r2ra 077 060 festuca A 0595002905950.0290 5950 029 049100560.49100560 4910 056 0303 I1 167473892167473.89216 7473 892 23921149252392114.92523 92114 925 0857 r2ra 083 039 centaurea with A 126201141 2620 114 1522017215220.1721 5220 172 0514 festuca I1 6418j2956 4181 295 100612870100612.87010 0612 870 0535 r2ra 077 066

growth of centaurea the degree of suppres- nutrients as well as water are similar grime sion increasing with increasing nitrogen levels 1979 fitter and hay 1987 in addition one growth response of pseudoroegneria and may hypothesize based on the resource ratio festuca to nitrogen when growing in mixture theory tilman 1982 that bromus is a superior with centaurea did not differ from their response competitor for nutrients other than nitrogen in monoculture likewise growth response of relative to centaurea by increasing nitrogen centaurea did not differ between monoculture both species should be limited by essential and mixtures with pseudoroegneria or festuca nutrients other than nitrogen and the species therefore these results suggest that these with the lowest R the superior competitiorcompetition native grasses are not likely to increase or sup- for the other nutrients should displace the press growth of centaurea regardless of nitro- species with the higher R for the same nutri- gen addition level this result is contrary to ents tilman 1990 the observed invasion of centaurea into com- the ability of bromus to out compete cen- munitiesmunities dominated by these grasses one ex- taurea in nutrient culture provides one expla- planation may be that disturbance especially nation for the observed population dynamics of grazing in the field creates gaps in the grass centaurea in the field roadsides seeded with community where centaurea can establish itself bromus are rarely invaded by centaurea weaver even though it is not a superior competitor for et al 1989 both field and laboratory observa- resources tions suggest that disturbed sites seeded simul- competitive interactions were greater be- taneously with centaurea and the exotic bro- tween each grass species and centaurea at the mus will be dominated by bromus the effec- high end of the nitrogen gradient this may be tivenesstiveness of bromus in suppressing centaurea a function of rapid growth thus in nitrogen may be increased with fertilization further- rich environments fast growing plants may more it may be expected that established bro- rapidly occupy space and usurp resources to mus plants will suppress the growth of centau- the exclusion of slow growing species grime rea seedlings the results of this study suggest 1979 radosevich and holt 1984 similar com- that at the seedling stage bromus may be used petipetitivetive effects may be expected to occur on to competitively exclude centaurea this method other soil resource gradients assuming adapta- of weed management merits trial in the field tions for acquisition of nitrogen and other mobile on the other hand the regional dominantsdominants 199611996 competitive suppression OF CENTAUREA 271

A C A

C C 0 0 AB u Q u0 A A A A A C bromus centaurea U cc 151.5 0 oolooi0010.01 01oi0.1 030.3 101.0loio relative nitrogen addition level

fig 2 relative competition intensity RCI of bromus and centaurea across 5 relative nitrogen addition levels letters above bars indicate whether RCI varies Dunduncanscarscaiseais multiple range test FP 0050.05oos within species across nitrogen level an asterisk indicates that RCI differs P 005 among species at that nitrogen level pseudoroegneria and festuca probably would control in a corn zea mays soybean glycine max not sufficiently suppress centaurea to decrease rotation weed science 43 269 275 LlLINDQUISTNDQUIST J L D A MORTENSEN S A CLAY R SCHMENK the potential for invasion J J KELLS K HOWATT AND P WESTRA 1996 stability advantages of the competitive method over of corn velvetleaf interference relationships weed herbicidesherbicides and biocontrolbiocontrol treatments used to science 44 309 313 manage centaurea are its long duration and MACHLIS L AND J TORREY 1956 plants in action free- san CA 282 appp low environmental impact given these advan- man francisco radosevich R S AND J S HOLT 1984 weed ecology tages exclusion of centaurea with bromus implication for vegetation management john wiley merits trial in environments where the danger & sons inc new york 265 appp of invasion exists RATKOWSKY D A 1983 nonlinear regression modeling a unified practical approach marcel dekker inc new york 276 appp literature CITED SAS 1988 SASSTAT user s guide release 603 SAS institute carscarygary NC 1028 appp COUSENS R 1985 A simple model relating yield loss to TILMAN D 1982 resource competition and community weed density annals ofappliedofapplied biology 107239107 239 252 structure princeton university press princeton NJ FITTER A H AND R K M HAY 1987 environmental 1990 mechanisms of plant competition for nutri- press physiology of plants and2nd edition academic ents the elements of a predictive theory of competi- inc san diego CA 423 appp tion pages 117 141 in J B grace and D tilman FORCELLA FE AND S HARVEY 1981 migration and distri- editors perspectives on plant competition academic bution of 100 alien weeds in NW USA 1881 1980 press inc new york herbarium montana state university bozeman 117 TYSER R AND C KEY 1988 spotted knapweed inm natural PP area fescue grasslands an ecological assessment GRACE J B 1995 on the measurement of plant competi- northwest science 62 151 160 tion intensity ecology 76 305 308 WEAVER T D GUSTAFSON J lichthardt AND B WOODS GRIME J P 1979 plant strategies and vegetation pro- 1989 distribution of exotic plants in the northern cesses john wiley and sons london 222 appp rocky mountains by environmental type and distur- HARPER J L 1977 population biology of plants academic bance condition MSU biology report 41 bozeman press inc san diego CA 892 appp MT 91 appp LINDQUIST J L B D MAXWELL D D BUHLER AND J L GUNSOLUS 1995 modeling the population dynamics received 26 september 1995 and economics of velvetleaf abutilon theophrastitheophrasti accepted 29 april 1996 creatgreat basin naturalist 563 C 1996 appp 272 275

indicators OF RED SQUIRREL tamiasciurusTAMIASCIURUS hudsonicusHUDSO NICUS ABUNDANCE IN THE WHITEBARK PINE ZONE

david J MattMattsonsonI1 and daniel P RemhartReinhartreinhartdremhart2reinhart22

ABSTRACT we investigated occupied squirrel biddensmiddens and squirrel sightings and vocalizations as indicators of red squirrelsaumsqum el hanlahuniatamtatamiasciuruswaniasciurusTamiaWania sciurus hudsonicushudsomcushudso nicus abundance in the high elevation whitebark pine pinus albialblalbicauhsalbicauliscaulis zone data were collected 1984 1989 hornfrom line transects located on 2 study sites in the yellowstone ecosystem we evaluated the performance of each measure on the basis ofofpiecisionprecision and biological considerations we concluded that of the 3 measures active mid dens weiewelewere the best indicator ofiedobiedof red squirrel abundance we also observed that the density of active biddensmiddens dropped by 48 66 between 1987 and 1989 following a severe drought and extensive wildwildfiieswildfireswildfifiresiesles that burned one of the study sites duringduidul ing 1988

key words transect fourier seriesserles midden vocalization sighting wildfire

whitebark pine finuspinus albialblalbicaulisalbicauhscaulis seeds are and the other near cooke city montana an important bear food that affects the survival immediately northeast of the park 4500n and fecundity of grizzly bears ursus arctos in these sites spanned the whitebark pine zone the yellowstone ecosystem use of pine seeds from 2360 to 2870 m elevation the whitebark by grizzgrizzliesgrizzlierlies is almost entirely contingent upon pine zone borders upper timberline and is the availability of cones cached in biddensmiddens iei e accordingly cold average annual temperatures larder hoards by red squirrels tamiasciurusTamiasciurus OC often windy and subject to deep 1 2 m hudsohudsomcushudsonicusnicus management of whitebark pine winter snow accumulations weaver 1990 habitats for grizzgrizzliesgrizzlierlies has thus become contingent upon management of red squirrel popula- MATERIALS AND METHODS tions mattson and reinhartremhart 1994 we studied red squirrels in the whitebark broad study objectives affected our transect pine zone using data collected from line tran- design we mapped the study area by habitat sects because these data included counts of type cover type strata based upon ground biddensmiddensmiddens animals and vocalizations we were truthed interpretation of 120000 aerial pho- able to evaluate the relative efficacy of these 3 tographyto the result was a fine scale mosaic indicators of squirrel presence we were inter- with individual map polygons forest stands ested in identifying a well behaved and rele- sometimes as small as individual squirrel terri- vant indicator of density to facilitate our inves- tories to minimize effects of edge between tigationti of relationships between squirrel different habitat types we placed transects so abundance and environmental factors such as as to maximize the number of right angle inter- midden use by grizzly bears we were also sections with stand boundaries as well as the interested in providing managers with an amount of intersection with stand interiors approach they could use to indicate squirrel because of this consideration and because for- abundance short of using intensive methods est and meadow were variously intermixed that relied upon marked animals transect lines were of unequal length we surveyed transects in the same order STUDY AREA eachyeareach yearyean beginning after 10 august and end- ing prior to 28 september two observers walked our study area consisted of 2 sites one permanently marked transect lines with one ob- located on the mt washburn massif in north server primarily responsible for observations and central yellowstone national park 4447n44047n the other primarily responsible for recording

I1 national biological seiselserviceSci vicevicc department offiiliofFiof fishyllyilili and wildlife resol11ccsre&ouices university of idaho moscow ID 83843 2usausS national paikpalkpark serviceSc ivleeivice resource management yellowstone national park WY 82190

272 199611996 RED SQUIRREL indicators 273

data and keeping on line at least one of the annual densities from this measure for this observers the junior author was the same area during all years at both sites vocalizations and total distance to line frequency distribu- observed squirrels or biddensmiddens both active and tions for each of the 3 measures did not differ inactive by criteria of finley 1969 were between the mt washburn and cooke city recorded along with their estimated perpen- study sites mantel haenszel x2xa2 for ordinal dicular distances from the transect line indi- categories P osl0510.51 0610.61 and 0350.35 for active vidual cone caches were not considered to be a mid 11 biddensmiddensdens vocalizations and sightings respec- middeimiddermidden and were easily distinguished from tivelyti perpendicular distributions of sightings these larger more permanent features and active biddensmiddens peaked in the nearest 10 we used the computer program TRANSECT m distance category although the distribution al burnham et 1980 to estimate densities of sightings more closely resembled a negative individual transects constituted sample units exponential and the distribution of biddensmiddens a for density calculations As recommended by negative sigmoidal function the majority 65 al burnham et 1980 we used the fourier and 78 respectively by year and study site series with 1 4 terms to estimate distance to of both these distributions were adequately fit line probability detection functions axexlgxlgx the xax22 test FP 010olo0.10 by a single term fourier distance at which we specified the limits of function distributions of vocalizations peaked detectability for our measures ie the cut point in the and2nd 11 20 m distance category and exerted considerable influence on the fit of the were characteristically 94 fit by a 2 or fourier function to the observed detection dis- higher term fourier function in 3 18 in- tributiontribution accordingly we varied cut points to stances we could not achieve an adequate fit achieve the best fit to each year or site spe by any model eifle data set because data were collected relationships among annual density esti- from only 35 transects on the mt washburn mates from the 3 measures were varied fig massif during 1984 and from 15 transects in 1 on the mt washburn site mean sighting the cooke city area during 1984 and 1989 com- and vocalization densities were weakly corre- pared to 57 and 21 transects respectively for lated r 07220.722 but tended to have overlap- all other years we also calculated densities ping 95 confidence intervals only 2 of 9 con- solely from these original 35 and 15 transects fidence intervals for the observed estimates all for all years so as to allow comparison with years for both the 1984 and inclusive samples results from 1984 did not contain the line describing perfect correspondence fig id in all but a single in- RESULTS stance cooke city 1984 mean midden densities were greater than mean densities of the we sampled the study 5 area yr 1984 1987 other 2 measures and were more strongly cor- and 1989 during 1988 wildfireswildfires burned 562000 related with sightings than vocalizations r ha of the yellowstone including area 52 of 09810.981 versus r 08310.831 respectively for tran- the mt washburn transects 47 severely sects 1 57 mt washburn fig lc however transects on the mt washburn area totaled in this case only two of nine 95 confidence 18918.9 km during 1984 mean transect length X intervals for midden and sighting densities in- 539 245 m s and 29829.8 km during the cluded the possibility ofperfect correspondence remaining 4 yr X 523 258 m similarly during 1984 and 1989 transects on the cooke conclusions city area totaled 16416.4 km X 1091 427 m and 21121.1 km during the remaining years X from these results we concluded that densi- 1005 405 m we recorded 124 squirrel sight- ties calculated from active biddensmiddens were more ings 641 vocalizations and 300 active biddensmiddens useful than densities calculated from the other on the mt washburn study site and 54 sight- 2 measures for indicating red squirrel abun- ings 528 vocalizations and 201 active biddensmiddens dance our conclusion followed from the greater on the cooke city study site during the 5 study apparent detectability of biddensmiddens compared to years the small number of sightings from the the squirrels themselves the consistency with cooke city site prevented us from estimating which a single term fourier function described 274 GREAT BASIN naturalist volume 56

a MT WASHBURN b COOKE CITY

1201.20 1201.20

v loo100 Z Z

080 0800.80

0600.60ogo 0600.60 U U 1 LL 0 4 0 0 0 40 0ajl7jl 020 U ZU Z Z

0000.00ooo 0000.00 Z 0 1 0000.00ooo00 0200.200 20 0400.400 40 0600.600ogo60 0800.800 80 1001.00loo00 1201 20 0000.00ooo0 00 0200.200 20 0400.400 40 0600.600ogo60 0800.800 80 1001.00loo1 00 1201 20 DENSITY OF vocalizations nha DENSITY OF vocalizations nha

c MT WASHBURN d MT WASHBURN

1201.20 loo1001.00

1001.00loo U 080 Z Z 0 0800.80 0600.60ogo ogo 0600.60 0 U 0 0400.40 0400.40 L 0 0 020 40 020 Z Z

000ooo 000 000ooo0 00 0200.200 20 0400.400 40 0600.600 60 0800 80 1001.00loo1 00 1201.201 20 000 020 0400.400 40 060 0800.800 80 1001.00loo1 00

DENSITY OF SIGHTINGS nha DENSITY OF SIGHTINGS niunimnha

fig 1 relationships between annual estimates of densityofdensity foiforbbr active biddensmiddens compared to vocalizations a for mt washburn and b for cooke city c sightings compared to active biddensmiddens for mt washburn and d sightings compared to vocalizations mt washburn 1984 1987 and 1989 error bars correspond to 95 confidence intervals solid circles to results fromblom all transectstianseetssects and open chelescircles to results from the fewer transects established and first surveyed in 1984 diag- onals representi epi esentbesent perfect correspondence between estimates

the probability detection distribution for mid ence of squirrels sightings and vocalizations dens and the resulting consistently smaller could suggest there were none standard errors for the density estimates in because red squirrel biddensmiddens are nonmobile addition scatter plots showed that active mid- often numerous relatively easily observed and den densities tended to be 0 when sighting typically associated with only one squirrel kil- and vocalization densities were not by impli- ham 1954 A smith 1968 wolff and zasada cation vocalization and especially sighting 1975 vahle and patton 1983 they are logical densities were more likely to underestimate indicators of squirrel abundance furthermore true squirrel densities iei e at the same time they do not suffer from sampling problems that active biddensmiddens clearly indicated the pres associated with weather season and time of day 199619961 RED SQUIRREL indicators 275

TABLE 1 estimated mean n hatihahaci1 and standard error sifsicsacsjc for densities of active biddensmiddens on the mt washburn and cooke city study sites 1984 1987 and 1989 percent coefficient of variations for annual variation 1984 1987 and percent decline in density from 1987 to 1989 results are given for the transects established and surveyed during 1984 1 35 and 1 15 and for the larger sample of transects surveyed during all other years except for 1989 in the cooke city area 1 57 and 1 21 mt washburn cooke city

trans 1 35 trans 1 57 trans 1 15 tramtrans 1 21 year mean STs i mean STs mean STs j mean sirsicsacsjcs 1984 0447 0083 0428 0077 1985 0557 0110 0632 0084 0682 0116 0635 0095 1986 0219 0042 0426 0078 0548 0126 0540 0098 1987 0453 0093 0838 0143 0544 0103 0790 0170 1989 0234 0062 0285 0098 0262 0137 CV 1984 1987 354 326 189 192 decline 1987 1989 483 660 518 to the same extent as do sightings and vocaliza- FERRON J J P OUELLET AND Y LEMAY 1986 spring and the tions cf C smith 1968 pauls 1978 ferron et summer time budgets and feeding behaviour of ledred squirrel tamlatamiasciurmtamiasciurusTamiasciurus hudsohudsonicushudsomcusnicus canadian jour- al 1986 these expectations were corroborated nal of zoology 64 385 393911 by our analysis biddensmiddens also have a direct tie FINLEY R B JR 1969 cone caches and biddensmiddens of tami to management of resources such as bears asciurus in the rocky mountain region university of that are of common concern in this zone kansas museum of natural history miscellaneous publications 51 233 273 of middens in our studstudyy densities active biddens KILHAM L 1954 territorial behaviour of red squirrel 1 area averaged between 020.2 and 080.8 hatlhatiha and journal of mammalogy 35 252 253 on both study sites were lowest during 1989 MATTSON D J AND D P REINHART 1994 bear use of following the drought and wildwildfiresfires of 1988 whitebarkwhitebaikbalk pine seeds in north america pages 212 220 compilers table 1 although annual variation tended to inw W C schmidt and FKEKFE K holtmeier proceedings international workshop on subalpine be on the mt washburn site compared greater stone pines and their environments the status of to the cooke city site this difference was not our knowledge US foiestforesthorest service general tech- statistically significant df 44 F 1311.31 P nical report INT GTR 309 050.5os both sites exhibited similar annual pat- PAULS R W 1978 behavioural strategiesstiategies relevant to the terns of variation including relatively low den- energy economy of the red squirrel tamiasciurusTamiasciurus journal of zoology 56 sities during 1984 and 1986 and a substantial hudsohudsonicushudsomcusnicus canadian 1519 1525 decline in active midden densities between SMITH C C 1968 the adaptive nature of social organiza- 1987 and 1989 tion in the genus of tree squirrels tamiasciurustamicisciurustamlaTamiasciurus ecological monographs 38 31 63 acknowledgments SMITH M C 1968 red squirrel responses to spruce cone failure in interior alaska journal of wildlife management 32 305 317 study was funded by the US national this VAHLE J R AND D R PATTON 1983 red squirrel cover park service through the interagency grizzly requirements in arizona mixed conifer forests jour- bear study team A hubbard D campopi- nal of forestry 81 14 15152222 ano D dunbar III111 and G green provided WEAVER T 1990 climates of subalpine pine woodlands pages 72 79 W C schmidt and K J mcdonald invaluable field assistance we also thank L S in compilerscompileis proceedings symposium on whitebark mills J peek R G wright D johnson C pine ecosystems ecology and management of a smith C benkman and an anonymous re- high mountain resource USU S forest service gen- viewer for their thoughtful reviews of this eral technical report INT 270 manuscript and its earlier versions WOLFF J 0 AND J C ZASADA 1975 red squirrel response to clearcut and shelterwood systems in interior alaska USU S forest service research note PNW literature CITED 255

BURNHAM K P D R ANDERSON AND J L LAAKE 1980 received 9 october 1995 estimation of density from line transect sampling of accepted 8 may 1996 biological populations wildlife monographs 72172 1 202 great basin naturalist 563 C 1996 appp 276 278 THERMAL characteristics OF MOUNTAIN LION DENS vernon C Bleichbleichl2bleich1212 becky A PiercelPiercepiereepiercellpiercel2pierce12122 jeffrey L davisdavisi and vicki L davisi

absrracrABSTRACr we used radiotelemetryradio telemetry and searched with a trained hound to locate the dens of 3 recently parturient mountain lions reilsbehsfelis concolor these dens were located in dense riparian vegetation along the same stream in the bot- tom of a steep canyon we monitored the circadian temperatures of 2 dens at 1 h intervals and compared them to ambient temperatures recorded simultaneously we found mountain lion dens to effectively moderate high ambient temperatures but these dens failed to provide a thermal advantage at the lowest ambient temperatures recorded in this investigation we conclude that mountain lion dens provide effective protection fromhrombrom thermal maxima for young immobile kittens

key words felis concolor mountain lion temperature california den behavior

female mountain lions feitsreitsreilseellsfelisfelts concolor select hound bruce 1918 to locate the dens and kit- protected locations in which to bear young tens we estimated the ages of these kittens shaw 198919897 7 beier et al 1995 but little infor- according to criteria summarized by anderson mation is available on den site characteristics 198343 and currier 1983 for this elusive felid here we describe some we examined the thermal characteristics of characteristics of 3 dens used by different the dens by placing a recording thermograph females and their litters and quantify the ther- model RTM ryan instruments inc kirk- mal characteristics of 2 of those dens land WA on the floor of each den and an identical instrument on the ground 100 inm description OF STUDY AREA away on a north exposure supporting sagebrush and pinyon pine because of the shrubs our study area is located inm mono co cali- and trees present on these north facing slopes fornia approximately 35 km NW of bishop thermothermographsthermographygraphs were not exposed directly to the 11825w 3720n inyo co california sun for most of each day hourly temperatures this area is on the western edge of the great were recorded at den 2 from 4 september to 4 basin immediately east of the crest of the october 1994 and at den 3 from 11 august to sierra nevada the dominant vegetation type 16 september 1995 we did not have access to in the general area is sagebrush artemisia tntritrl thermographsthermographythermographs during the period that den 1 was dentata scrub with pinyon pine pinus mono active we made ocular estimates of tree height phylla forest at higher elevations dense vege- and canopy closure as well as horizontal cover tation dominated by willows salix sppapp and at each den wild rose rosa sppapp occurs along the major we used analysis of variance and analysis of water courses covariance to explore the effects of day and time on temperature simple linear regression METHODS to examine the relationship between day of the study and daily temperature and t tests to during august and september 1994 and compare den temperatures with ambient tem- 1995 telemetry indicated that several adult peraperaturestures zar 1984 females in our investigation of mountain lion ecology had restricted their daily movements RESULTS these females returned repeatedly to the same locations suggesting that they had established three dens containing kittens were located natal dens beier et al 1995 we searched along the owens river den I1 contained I1 male these 3 areas and after detecting vocalizations and I1 female den 2 contained 3 males and I1 of neonatal mountain lions we used a trained female den 3 contained 2 males and I1 female

lcaliffildialilili toimi department offisliof fish and gamegarne 407 W line st bishop CA 93514 2111stituteniititute of arctic biology and department of biology and wildlife university of alaska fairbanksfanran banks AK 99775

276 199611996 MOUNTAIN LION DENS 277 we estimated the kittens at dens 1 and 3 to be when date was used as a covariate to control 20 days of age and those at den 2 to be 10 for daily solar radiation the mean temperature days old differential also varied on an hourly basis at all 3 dens were located in dense groves of both dens den 2 F 112271112.271 df 23 719 P willows that ranged in height to approximately 00010.001 den 3 F 329936329.9363299365329.9365 df 23 863 P 4 m wild rose was abundant at all 3 sites and 00010.001 hourly ambient temperatures were each den was located 50 m from the river greater than corresponding den temperatures canopy closure at each den was nearly 100 at both locations den 2 t 3228532.285 df 743 and direct sunlight did not reach the substrate FP 00010.001 den 3 t 2566225.662 df 887 FP during any of our midday visits n 2 den 1 n 00010.001 this difference was especially pro- 5 den 2 n 3 den 3 horizontal cover at nounced at high ambient temperatures 31 each location was sufficiently dense that even C HAT fig 1 at HAT the temperature dif- while standing we were totally obscured from ferentialferential at den 2 Tx 219221.92 4494.49 QC was each other s view at 3 m the substrate of all 3 times that at moderate ambient tempera- 3 dens was littered with deciduous leaves as tures 31 C MAT x 6036.03 4464.46 QC and well as tree trunks branches twigs and bark the temperature differential at den 3 afx5f we were able to reach the kittens only by 135613.56 8378.37 QC at HAT was nearly 5 times crawling into the dense vegetation present at that at MAT af5fx 2872.87 3503.50 QC at den 2 the each site mean range of daily ambient temperatures x we found significant differences between 289628.96 7817.81 QC was nearlydearly double that of ag the dens in mean ambient temperature xgedenyden 23 daily den temperatures x 157915.79 5265.26 QC 131813.18 9949.94 s C xdnddnx den 3 204720.47 logi106110.61 t 158315.83 df 30 FP 00010.001 similarly at C F 202584202.584 df 1 1630 P 000150.001500010.001 den 3 the mean range of daily ambient temper- agn 4 mean den temperature xenXxgnden 2 601goi6.01 5775.77 atures xY 325432.54 3713.71 QC was isls151.5 times Y C x den 3 152215.22 7087.08 C F 807949807.949 df that of daily den temperatures x 208920.89 1 1630 P 00010.001 and mean daily tempera- 3503.50 C t 152415.24 df 36 FP 00010.001 for ture differential ambient temperature den both locations combined den temperatures gos temperature edeydex 2 7167.16 6056.05 C agxgX den 3 were less than ambient temperatures for all but 5255.25 6096.09gog C F 4022440.224 df 1 1630 P 2 02 020.2 of the paired hourly observations 00010.001 at den 2 there was a significant effect of day on ambient temperature F 38143.814 df discussion 30 713 P 00010.001 den temperature F 31913.191 df 30 713 P 00010.001 and tempera- these mountain lion dens effectively mod- ture differential F 43204.320 df 30 713 P erated high ambient temperatures consistent 00010.001 at den 3 however there was no such with the hypothesis of shaw 1989 that dens effect on ambient temperature F 04210.421 df play an important role in protecting young 36 851 P 09990.999 den temperature F defenseless kittens from thermal maxima at 05350.535 df 36 851 P 098909890.989 or temperature HAT mean temperature differentials were 3 5 differential F 04880.488 df 36 851 P times greater than at MAT fig 1 there were 09950.995 As the study progressed there was a significant effects of time of day both dens significant decline at den 2 in ambient temper- and day length den 2 on temperature differ- ature r 03400.340 P 00010.001 den temperature ential and hence the moderating influence of r 01120.112 P 00010.001 and temperature dif- the dens nevertheless den temperatures ferentferentialial r 02280.228 P 00010.001 lesser de- were less variable than were ambient tempera- clines in ambient temperature r 01000.100 P tures we found no evidence that these dens 00010.001 den temperature r 00510.051 P provided a thermal advantage ie den tem- 00010.001 and temperature differential r 00480.048 peraperaturestures greater than ambient temperatures P 00010.001 occurred at den 3 at the minimum ambient temperatures we at both dens there was significant diel vari- recorded dens may however provide protec- ation in ambient temperature den 2 F tion for kittens when temperatures fall below 103382103.382 df 23 720 P 00010.001 den 3 F those that we encountered 618443618.443 df 23 864 and den temperature few descriptions of mountain lion dens are den 2 F 9100891.008 df 23 720 P 00010.001 available but females may select caves rocky den 3 F 431275431.275 df 23 864 P 00010.001 areas or dense thickets in which to bear young 278 GREAT BASIN naturalist volume 56

30 1 regulatorymomoregulatory purposes might be lessened under i these circumstances fewer movements by kit- 0 00oo 0 0 0 tens may decrease the probability of discovery

0 0 by potential predators thereby enhancing the 0 0 survival of young defenseless mountain lions 0 00 t Z 0 1 0 W 20 0 0 0 0 00oo 1 0 0 r 0 acknowledgments W 0 0 0 .0 U 0 0 L j 00 0 we thank S parmenter D and C W 0 0 becker 0 so 000ooo eo T 00 0 milliron for us the & g lending recording thermo 0 t V 0 10 0 0 00oo N E 0 graphs G andrew and P beier for critical cc 0 0 0 0 W 0 comments on an early draft of the manuscript CL 1 0 2 0 and M W oehler sr for assistance in the field uj financial support was provided by the mule 0 p 0 foundation university of callcailcaliforniaCalifomia white 1 deer 0 0 10 20 30 40 50 mountain research station california department of fish and game CDFG national rifle AMBIENT temperature C association safari club international and the fish and game advisory committees of inyo 1 mean fig temperature differential ambient den is and mono counties this is a contribution from more than 3 times greater at high ambient temperatures plan 31 QC than at moderate ambient temperatures 31 C the CDFG deer herd management data from dens 2 and 3 combined F 1 24107241941 07 df 1 implementation program 1630 P 00010.0010 ooiool001 mountain lion dens in dense vegetation effectively moderate extreme high temperatures and afford literature CITED young helpless kittens protection from ambient maxima consistent with the hypothesis of shaw 1989 ANDERSON A E 1983 A critical review of literature on puma felis concolor colorado division of wildlife special report 54 1 91 bruce 1918 young and goldman 1946 mcbride BEIER P D CHOATE AND R H BARRETTBARRETr 1995 move- 1976 russell 1978 shaw 1989 we hypothe- ment patterns of mountain lions during different size that thermal characteristics vary among behaviors journal of mammalogy 76 1056 1070 BRUCE J C 1918 lioness tracked to lair california fish types of dens and that mountain lions inhabit- and game 4 152 153 ing particular environments select den sites CURRIER M J P 1983 felis concolor mammalian species based in part on the thermal advantages 2001200 1 7 they provide MCBRIDE R T 1976 the status and ecology of the moun- stanle mex in an area with a warm mediterranean cli- tain lion felis concolor stanleyanayana of the texas ieoicojeo border unpublished master s thesis sul ross mate beier et al 1995 reported 2 dens that state university alpine TX were located in a small canyon with very heavy RUSSELL K R 1978 mountain lion pages 207 225 in J L cover of brush similar to those we investi- schmidt and D L gilbert editors big game of north gated dens located in thick woody vegetation america ecology and management stackpole books conceal harrisburgHarnsburg PA may young that are vulnerable to pre- SHAW H 1989 soul among lions johnson publishing com- dation but they also provide protection for kit- pany boulder CO tens from extreme temperatures associated YOUNG S P AND E A GOLDMAN 1946 the puma myste- with direct insolation such locations provide rious american cat american wildlife institute wash- important thermal benefits for kittens at high ington DC ZAR J H 1984 statisticalBiobiostatistical analysis prentice hall engle- ambient temperatures and a more stable ther- wood cliffs NJ mal environment than exists outside the den throughout the range of ambient temperatures received 30 december 1995 we recorded movements by kittens for ther accepted 5 april 1996 great basin naturalist 563 0 1996 appp 279 280

JAMES WILLIAM BEE 1913 1996

wilmer W tannerlnannerltanner1

tion pertaining to indian winter camps summer camping areas and burial grounds and an insight into the role of utah lake and the surrounding mountains as providers of abundant fish and game their travels near this lake and in the mountains brought them in contact with numerous birds each spring flocks of birds entered the valley some remained and others moved on this phenomenon stimulated a great interest so much so that james lishisbishrs father and various friends became amature ornithologists their ornithological work encompassed life history studies observation of arrivals in the spring and investigation of nests and nesting ulti- mately this interest in birds lead to the assem- bly and preparation of eggs for those species nesting in the valleys and mountains of central utah thus was born a naturalist whose contribu- tions are invaluable and most of which could james W bee not now be assembled the archaeological collections are presently at the museum of peo- ples and cultures brigham young university james W bee professor of zoology and emer- james and his father contributed 812 sets of itus university of kansas lawrence kansas bird eggs and 112 single eggs representing 234 died at seattle washington 18 april 1996 he species james contributed 7918 mammal 245 was born 25 september 1913 in provo utah bird and 504 amphibian and reptile specimens his family including parents robert G and to the M L bean life science museum also at mary culbertson bee and brother and sister brigham young university in the bean museum max and mary were residents of provo utah library are field records 27 volumes from where they received their early education it james and 20 volumes of his father s all well was from this setting in utah valley that james documented and done with great care these was introduced at an early age to the sciences were written in the field as the data were of archaeology and ornithology by his father obtained and represent field records of a time who loved natural history and the little known when some pristine conditions still existed history of utah valley its lake and its early james entered brigham young university in inhabitants 1932 and received his BA degree in 1937 As a youth and young man he accompanied world war II11 interrupted his studies for the his father on many collecting trips that resulted MA but this he finished in 1947 As an under- in assembling artifacts of the past these graduate he became interested in and re- archaeological finds provided valuable informainforms searched mammals thus his master s research

1mam L bean museum bnghambrighambangham young university provo UT 84602

279 280 GREAT BASIN naturalist volume 56 was the mammals of utah county while in the publications armed forces 1941 1946 he was trained as a hospital administrator and served as a sergeant BEE JAMES W major organizing 50 key men as a cadre to 1957 biological survey of the virgin islands national park procedures and establish a new hospital supervised several observations recommen- he dations 33 pages 2 maps submitted to national new hospital departments and for a year and a park service half served in field hospitals for airborne in units 1958 birds found in the arctic slope of northern alaska in india assam burma and china during university of kansas publications museum of these years he met annette P malseed RNR N natural history 105 163 211 2 plates I1 figure they were married 15 october 1945 in kun in text ming yeaman china 1970 vasco M tanner a diversified career great in september 1948 james entered the uni- basin naturalist 30 216 217 versity of kansas to continue his research in 1994 rough grouse siting the trumpeter san juan mammalogy with a desire to complete his audubon society 1443144 3

study on the genus microtisMimicrotuscrotis he completed his BEE JAMES W DUMITRU MURARIUMUBARIU AND ROBERT S studies at KU and spent a summer at friday HOFFMANA harbor washington he was a noted field 1980 histology and histochemistry of specified integu- zoologist and spent many years collecting re- mentary glands in eight species of north ameri- search material and field data for the museum can shrews mammalia insectivora travauxtramaux du of natural history at the university of kansas museum d histoireHistoire naturelleNaturelle grigor antipas bucharest romania 22 547 569 students doing research in vertebrate zoology at brigham young university or at the univer- BEE JAMES W AND E R HALL sity of kansas will find numerous specimen 1951 an instance of coyote dog hybridization trans- tags labeled collected by james bee after 37 actions of the kansas academy of science 541 years he retired from KU and built a new 73 77 4 figures 1 table home on lopez island washington james and 1954 occurrence of the harbor porpoise at point annette were the parents of three children barrow alaska journal of mammalogy 351 122 james robert annette christine kenagy and 123 mary pauline bee kaufman 1956 mammals of northern alaska on the arctic slope university of kansas museum of natural his- it was my pleasure to have spent several tory miscellanceous publications 8 1 309 4 summer field trips with james A highlight was plates 122 figures 51 tables the summer of 1939 when we studied the ver- 1960 the red fig eating bat stenodermaStenoderma rufuerufum des tetebrates of western utah county during this harertmarert found alive in the west indies mam- time we prepared and assembled museum malia museum dhistoire naturelleNaturelle parispans 141 67 75 2 figures in text specimens of importance to me was finding a in nesting colony of the western slinkskink and secur- BEE JAMES W AND HOWARD LEVENSON additional of hypsigalena we ing specimens bald eagle use of kansas river riparian habitat both participated in the new discoveries and it in northeastern kansas kansas ornithological was obvious that jim was at his best min prepar- society bulletin 34 28 33 ing precise field data I1 learned much from him that summer and appreciated his dedication to a complete understanding of the natural world we were investigating james had a very likeable personality that was reflected in his family which he held in high esteem great basin naturalist 563 0 1996 appp 281 282

BROOK stickleback CULAEA inconstantinconstansCONSTANSIN KIRTLAND 184118411 A NEW ADDITION TO THE UPPER COLORADO RIVER BASIN FISH FAUNA

timothy moddel and G bruce haineslhamesthames1

key words brook stickleback range extension nonnativenormative

brook stickleback culaeaculaba inconstans is a a quatrefoil light trap at the outlet of old small gasterosteid fish native to arctic and charley wash river kilometer RK 402 on the atlantic drainages in north america the green river RK measured from the conflu- species native range extends west from nova ence of the green and colorado rivers four scotia to british columbia and south from the adult fish 41 46 48 54 mm TQTL were col- northwest territories to southern ohio drain- lected between 1 october and 12 october 1995 ages including the mississippi missouri river from old charley wash a wetland on the above the confluence of the illinois river ouray national wildlife refuge that connects scott and crossman 1973 hubbs and lagler to the green river during high spring flows 1958 reported brook stickleback from the fish were collected when the wetland was illinois river in illinois and the missouri river drained modde in press all were found in in kansas historical accounts exist relictualrevictualofrelictualof reilerelictualtuai low or no velocity habitats populations in the platte river system but tyus et al 1982 cited the establishment of cross 1967 noted its absence from kansas 42 nonnative fishes in the upper colorado river an isolated and presumably relict population compared to 13 native species brook stickle- occurs in the canadian river drainage of new back is an additional transplanted species prob- mexico koster 1957 brook stickleback has ably the result of human introduction rather been collected outside its native range in than a natural range extension brook stickle- alabama boschung 1992 kentucky burr and back introductions elsewhere in the united warren 1986 tennessee etnier and starnes states were presumably through bait bucket 1993 the rio grande river drainage in new transfers or contaminated game fish stockings mexico sublette et al 1990 colorado zuck- zuckerman and behnke 1986 sublette et al erman and behnke 1986 and the klamath 1990 river california peter moyle university of california davis personal communication literature CITED between july and october 1995 we collected 5 brook stickleback from the middle BOSCHUNG H T 1992 catalogue of freshwater and maimalmarineme green river uintah county utah the ist rec- fishes of alabama alabama museum of natural his- ord for the species in utah catalog number tory bulletin 14114 1 266 LFL 24871 larval fish laboratory colorado BURR B M AND M L WARREN JR 1986 A distributional atlas of kentucky fishes kentucky nature pre- state university brook stickleback was first serves commission scientific and technical seriesserlessenes reported elsewhere in the upper colorado no 4 river drainage in 3 small tributatributariesries of the elk CROSS F B 1967 handbook of fishes of kansas state bio- river south coleman and deep creeks in logical survey and the university of kansas museum northwestern colorado in 1993 jake bennett of natural history lawrence colorado division of wildlife personal com- ETNIER D A AND W C STARNES 1993 the fishes of ten- press municationmunication nessee the university of tennessee knoxville HUBBS C L AND K F LAGLER 1958 fishes of the great one brook stickleback juvenile 27 mm lakes region the university of michigan press total length TL was collected 18 july 1995 in ann arbor

icoloradolcolorado riverrivel fish project USU S fish and wildlife service 266 west 100 north suite 2 vernalveinalmernal UT 84078

281 282 GREAT BASIN naturalist volume 56

kosterKOSTLR W J 1957 guide to the fishes of new mexico and status pages 12 70 in W H miller H M tyus university of new mexico press albuquerque and C A carlson editors fishes of the upper col- MODDE T in press juvenile razorback sucker in a man- orado river system present and future american aged wetland adjacent to the green river great fisheries society bethesda MD basin naturalist ZUCKERMAN L D AND R J BEHNKE 1986 introduced SCOTT scotr W B AND E J CROSSMAN 1973 fishes of canadaofcanada fish in the san luis valley colorado pages 435 452 fisheries Reseaichsealch research board of canada bulletin 184 in R H stroud editor role offish culture in fishery SUBLEsuijlcnei re J E M D HATCH AND M SUBLETTE 1990 management american fisheries society bethesda the fishes of new mexico university of new mex- MD icoieoleo press albuquerque TYUS H M B D BURDICK R A VALDEZ C M HAYNES received 4 january 1996 T A LYTLE AND C R bebryBERRYBLRRY 1982 fishes of the accepted 12 april 1996 upper colorado river basin distribution abundance

gleatgreat basin naturalist 563 0 1996 p 282

ERRATA

correction to lence of red coloration in california humming- bird flowers american naturalist 100 85 98 sutherland steven D and robert K vickery and 2 K A grant and V grant 1968 hum- jr 1993 on the relative importance of mingmingbirdsbirds and their flowers columbia uni- flower color shape and nectar rewards in versity press new york 115 appp among other attracting pollinatorspollinators to mimulus great references in fact the grants point out just basin naturalist 53 107 117 the opposite ie that experimental investigation shows that hummingbirds have not evolved the article states hummingbirds are com- a preference for red or any other color actu- monly said to have evolved a preference for ally sutherland and vickery s article comes to red or orange red flowers citing 1 K A this conclusion also grant 1966 A hypothesis concerning the preva great basin naturalist 563 D 1996 appp 283 285

BOOK REVIEW snakes of utah douglas C cox and wilmer would be more useful if a caption were shown W tanner mark philbrick photography by the other photographs throughout the text monte L bean life science museum brig- eg the photo opposite page I1 and those shown ham young university provo UT 1996 on pages 3 4 5 8 the herpetologist will 1795179517.95 softcover probably recognize these without caption but as stated it s likely these specialists will not be snakes of utah anticipated for some time the primary users of the text identification of is finally available for distribution this book- snakes by these photographs may not be obvi- let 92 total pages includes all known species ous to most readers most photos show colors and subspecies of snakes found in the state and patterns of snakes but a few such as the with brief descriptions habits and habitats full view of the upper basin garter snake on along with colored photographs of each while page 59 do not show these identifiable fea- most people will likely shudder at the thought tures it s interesting that the only snake not of snakes especially while viewing photographs represented by a photo of the entire body is the enthusiast will recognize the value of the the sonoran lyre snake on page 67 one won- illustrations and other published information ders why perhaps it s because this snake is generally the booklet is written in nonscien- considered to be rare however the dixie tific language but it also includes some scien- college natural science museum contains tific notations for instance scientific names records of 7 specimens 2 having been found in and authorities of the 33 species and subspecies what is now considered downtodowntownwr st george along with common names are included for I1 specimen as recently as 1980 it seems likely each of interest perhaps only to the special- that with a little effort one of these rare snakes ist is the fact that only 2 binobinomialsmials are found might have been found the photo of the utah among all utah snakes 31 are trinotrinomialsmials it blind snake on page 17 is a surprise of the might be concluded that because of subspeci several dozen blind snakes observed by this atlon only 27 kinds of snakes are found in utah writer representing localities from the red to the general public a night snake is a night cliffs recreation area near leeds washing- snake a garter snake is a garter snake and a ton county to the extreme northwest corner rattlesnake is a rattlesnake herpetologists have of arizona not I1 specimen even approached named subspecies for practically all snakes this dark phase they have all been a pale tan compounding one s knowledge of these animals color frequently showing a suffusion of pink technically where closely related subspecies another important contribution of this book- show sympatric distribution there should be let is the distribution maps included with each intergradation between the 2 types most indi- species along with the general and sometimes viduals using this booklet will probably not specific distribution of the snake within the recognize differences between related sub- state while it is difficult to show accuracy on species found especially in these sympatric a small map some maps are erroneous for regions if interintergradesgrades are not present then instance the distribution of the painted desert these should be elevated to species and not kept glossy snake is in the extreme southeastern as subspecies little information is found in the sector of the state adjacent to northeastern booklet on intergradation of characteristics arizona7 page 40 the map however shows it an important contribution of this booklet is is found more south central than southeastern the colored photographs while not captioned an inconsistency from text to map is also most photographs are obvious because they are observed with the california king snake page shown on the page opposite the name and other 46 if this snake occurs from the southwest information on that snake this publication corner east to the colorado river why does

283 284 GREAT BASIN naturalist volume 56 the distributional map extend considerably the writer wonders at the importance of beyond the colorado river along the san juan the full page of illustrations page 13 showing river nothing in the text is speculative of a scalation with so little reference to most of these range extension the maps of the utah moun- features in descriptions some of these features tain king snake page 48 and the utah milk are referenced most are not snake page 50 do not accurately depict their while full pages of color separate groups of known distributions in washington county snakes does this mean that joshua trees are on page 60 of the western blackneck garter characteristic of the distribution of the utah snake the text states its northernmost habitat blind snake although the illustration on page is associated with streams in the regions of 18 may be typical of the habitat of the rubber southeastern utah the map shows its distri- boa in utah and on page 72 of the habitat of bution into east central utah reference is some of the rattlesnakes does the illustration made to a ground snake having been collected on page 22 depict the typical distribution of in carbon county far from its known range the colubridscolubrids perhaps these division pages and this area is shown on the map might this were added merely for color nevertheless they specimen have been one that escaped or was are attractive released from captivity reports have been the authors of the booklet include a number made of individuals transporting this snake of interesting anthropomorphisms perhaps from the st george area sheiewheiewhere it is common intentionally some of these are noted 1 in to elsewhere in the state there is speculation the introduction the statement is made page that the utah blackhead snake may occur fur- 5 that the snake employs rocks and brush to ther north in emery and carbon counties snag the skin and hold it while the snake crawls the proposed expansion is not shown on the out one wonders if the snake does this inten- map why might it not then be found in tiotionallynally 2 denning is a behavior pattern that wayne county and perhaps even san juan and provides the snake with an opportunity to grand counties if the midget faded rattle- come in contact with other snakes of the same snake is found at flaming gorge why does the species page 6 3 of the rubber boa it will map not show distribution in that area often cling like a bracelet and seem to enjoy it while it would add to the length of the text as much as the perpersonsor page 20 4 the it would have been better had the authors given statement is made about the western yellow- complete distribution ranges for all species belly racer page 28 that it will attempt to bite and subspecies rather than just a few A snake if it feels at all threatened 5 another exam- doerndoesndoesrt t recognize a political boundary as being ple is that rattlesnakes use the rattle as a its limits however it could be reasoned if the warning device to intimidate other animals distribution extends to the utah boundary the that may harm the snake page 75 occurrence of that snake would also be in the miscellaneous errors or inconsistencies in neighboring state narrative grammatical or otherwise are found the full page map of the state of utah page the introduction for instance discusses tall 11 is a good addition to the text however tales and folklore of the american west this with the number of snakes found only in utah s booklet is of course about snakes of one region mojave desert this feature might have been of the american west but tall tales and folk- identified along with the others in the geo- lore even some of the same stories heard in graphical and ecological descriptions of utah the american west are repeated wherever pages 9 10 considerable discussion is given snakes are found about montane regions some at high eleva- on pages 4 and 5 the statement is made tions yet little is written about the low hot that the mouth is the most universally used desert or the higher cold desert although the weapon employed by snakes in self defense authors admit to the richness of reptile fauna the emphasis is obvious because the accom- especially in the low hot desert the south- panying text is about self defense but snakes western region of the state use their mouths more often as a means of in addition to these other features snakes obtaining food also in the introduction the of utah includes both glossary though not statement is made that these studies and our inclusive of all technical words used in the museum program help them to understand text and index page 6 emphasis added later in the text 199619961 BOOK REVIEW 285

page 9 reference is made to brigham young an inconsistency is noted about the utah university s monte L bean life science mountain king snake and the utah milk snake museum the complete identification of the page 48 states if a specimen has a white nose museum should have been made when it was it is most likely a mountain king snake if how- first referenced on page 6 it could be pointed ever it has a black nose it is probably a milk out too that other schools and museums snake these characteristics are not completely might have the same purpose to help them reliable emphasis added page 50 states that 11 to understand about snakes the milk snake differs in that it has a black while the following is not necessarily in nose error it reflects a writing style on page 12 the on pages 68 and 70 the habits of the mesa following statements are made these snakes verde night snake and the desert night snake do not pose any threat to man but they do pro- are described as nocturnal secretive and sel- vide a mild venom to help immobilize their dom seen furthermore it is stated that the prey their prey includes worms insects frogs former feeds primarily on the lizard ufauta stans lizards and small mammals in writing re- aurianabunanaburiana uniuniformisuniformistformis and other small lizards peated words and phrases should be avoided while the latter feeds primarily on the side in consecutive sentences or within the same blotched lizard uta stansburianastansbunanastansbstansburunanaianalana stansburianastansbunanastansbstansburunanaianalana sentence it could better have been written to one wonders about this inasmuch as lizards help immobilize their prey which includes are primarily diurnal and snakes nocturnal of worms insects course snakes could feed at night while lizards in the introduction to the tropical wormlike are inactive snakes the statement is made that they feed while reference is made in the booklet about on insects and worms especially termites and the influence of soil on the ground color of ants found in the soil the emphasis in this some snakes there is no mention of this occur- statement suggests that termites and ants are ring in the mojave desert sidewinder page kinds of worms this should have been written 78 of the hundreds of sidewinderssidewinders observed 11 they feed on worms and insects especially by the author in the past 50 years the influ- termites and ants in reference to the utah ence of soil color on the ground color of the blind snake the statement is made also on snake is most obvious page 15 that vasco M tanner had seven despite these criticisms snakes of utah specimens to examine and the name is based should contribute importantly to our knowl- on no 662 in the BYU type collection name edge of these reptiles within a limited political is inappropriately used although specimen region As noted the booklet is written for lay- no 662 might have been published as the men and its distribution is more appropriate type specimen in national and state parks and monuments than one of the most frequently made grammat- in the scientific community it is a must for ical errors in writing is the inconsistency of backpackers individuals and families spend- singsingularsulars and plurals within a sentence on ing time in the out of doors where snakes might page 20 this type of error is made the rubber be encountered the authors the photographer boa is a delightful animal to have around and the publisher are to be commended for their wrist inasmuch as their is plural the finally making this booklet available plurality of wrists must also be used reference is made twice on pages 30 and andrew H barnum 44 that the snakes occur on the margins of professor emeritus deciduous forests small groups of deciduous dixie college trees may occur in riparian areas or where st george UT 84770 trees are cultivated but technically deciduous forests do not occur in the state of utah the redundant statement is made about the western leafleafnosenose snake that the rostral scale looks leaflike THE FUTURE OF ARID grasslands identifying ISSUES FINDING SOLUTIONS 9 13 OCTOBER 1996 TUCSON ARIZONA

A solution oriented conference for everyone interested in the future of grasslands in the american southwest and northern mexico this four day conference will focus on understanding problems facing those grasslands and practical tools for grassland management preservation and restoration attendees will be a mix of private and public land managers and owners scien- tists representatives of nonprofit groups and concerned citi- zens two full days will be spent in the field studying examples of grassland management in southern arizona the other two days will include keynote speakers and panelists as well as elltrpbep small group discussion and information sessions the final day will focus on methods for preservation ranging from coordi- nated monitoring systems land use and taxation tools to public involvement techniques

most of the speakers and panelists will be invited but abstracts are welcome for a few open sessions dealing with grasslands management interrelationships between grasslands and humans or wildlife and specific methods for preservation especially success stories

researchers are encouraged to submit abstracts for poster sessions which will be incorporated into the program featuring on the ground examples of problem solving to protect or restore grasslands both successful and unsuccessful examples are sought to illustrate what has and has not worked and why

the conference is organized by the audubon research ranch and is co- sponsored by numerous government agencies educational institutions and nonprofit groups

for more information grasslands conference tucson audubon society 300 E university 128 tucson AZ 85705 or university of arizona water resources center 520 7929591792 9591 information FOR AUTHORS the great basin naturalist welcomes previously VOUCHER SPECIMENS authors are encouraged to unpublished manuscripts pertaining to the biologi- designate properly prepare label and deposit cal natural history of western north america high quality voucher specimens and cultures docu- preference will be given to concise manuscripts of menting their research in an established permanent up to 12000 words simple species lists are dis- collection and to cite the repository in publication coucouragedraged references IN THE TEXT are cited by author and SUBMIT manuscripts to richard W baumann date eg martin 1989or1989 or martin 1989 multiple editor great basin naturalist 290 MLBM PO box citations should be separated by commas and listed 20200 brigham young university provo UT in chronochronologicallogical 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ponds journal of wildlife management por scientific style and format the CBE manualforManumanualalforalnorainorfor 44695 700 publishers authors editors and ath6th edition sousa W PE 1985 disturbance and patch dynamics council of biology editors inc41ineinclne 11 south lasalle on rocky intertidal shores pages 101 124 in street 1400 chicago IL 60603 USA PHONE suite S T A pickett and PE S white eds the ecolo- FAX 201 0214 we do 3122010101312 201 0101 3122010214312 however gy of natural disturbance and patch dynamics differ in our treatment of entries in literature cited academicA press new york authors may consult vol 51 no 2 of this journal coulson R N and A witter 1984 forest ento- format these J for specific instructions on instruc- mology ecology and management john wiley manuscripts SUBMITTED tions guidelines FOR and sons inc new york 669 appp TO THE GREAT BASIN naturalist are printed at the back of the issue also check the most recent issue TABLES are double spaced on separate 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mail address importance to be used for indexing 3 copies of the manuscript and WordwordperfectPerfect TEXT has centered main headings printed in all diskette capital letters second level headings are centered conformity with instructions in upper and lowercase letters third level head- photocopies of illustrations ings begin paragraphs issn0017ISSN 001736140017 3614 GREAT BASIN naturalist vol 56 no 3 july 1996 CONTENTS articles biogeographic significance of low elevation records for neotoma cinerea from the northern bonneville basin utah donald K grayson stephanie D livingston eric rickartRidkart and monson W shaver illIII111 191 synopsis of the mosses of wyoming PEMM eckel 197 variation in bitterbrushbitterbrush purshia dentatatritridentatatridentate pursh crude protein in southwestern montana carl L wambolt W wyatt fraas and michael R frisina 205 dam forming cacti and nitrogen enrichment in a pinon juniper woodland in northwestern arizona molly thomasrhomas hysell and charles C criergrier 211 distribution and ecological characteristics of lewisia longipetalalongipetala piper clay a high altitude endemic plant anne S halford and robert S nowak 225 larger ectoparasites of the idaho ground squirrel spermophilus brunbrunneusbrunneousneus eric yensen craig R baird and paul W sherman 237 roost sites of the silver haired bat lasionyctelasionycteristisris noctivagantnoctivagans in the black hills south dakota todd A mattson steven W buskirk and nancy L stanton 247 perceptions of utah alfalfa growers about wildlife damage to their hay crops implications for managing wildlife on private land terry A messmer and sue schroeder 254 spatial relationships among young Cercocercocarpuscarpus ledifoliusledifolius curlleafcurlleancurlleaf mountain mahogany brad W schultz robin J tausch and paul T tueller 261 potential for controlling the spread of centaurea macumaculosamaculoselosa with grass competition john L lindquist bruce D maxwell and T weaver 267 indicators of red squirrel tamiasciurusTamiasciurus hudsohudsonicusnicus abundance in the whitebark pine zone david J mattson and daniel P reinhart 272 thermal characteristics of mountain lion dens vernonvemonmemon C bleich becky M pierce jeffrey L davis and vicki L davis 276 james william bee 1913 1996 wilmer W tanner 279 note brook stickleback culaeaculaba inconstans kirtland 1841 a new addition to the upper colorado river basin fish fauna timothy modde and G bruce haines 281 errata 282 book review snakes of utah douglas C cox and wilmer W tanner andrew H barnum 283