Full Issue, Vol. 57 No. 4

Great Basin Naturalist

Volume 57 Number 4 Article 15

10-31-1997

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GREAT BASINBAS I1 N naturalistnaturalist

VOLUME 57 n2na 4 OCTOBER 1997

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 nmshbll1byueduNMS hbllibyuedu

associate editors J R CALLAHAN PAUL C MARSH museum of southwestern biology university of center for environmental studies arizona new mexico albuquerque NM state university tempe AZ 85287 mailing address box 3140 hemet CA 92546 STANLEY D SMITH BRUCE D ESHELMAN department of biology department of biological sciences university of university of nevada las vegas wisconsin whitewater whitewater WI 53190 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great basin naturalist the great basin naturalist founded in 1939 is published quarterly by brigham young university unpublished manuscripts that further our biological understanding of the great basin and surrounding areas in western north america are accepted for publication subscriptions annual subscriptions to the grgreateat basin naturalist for 1997 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 busi- ness should be directed to the editor great basin naturalist5 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

copyright 0 1997 by brigham young university ISSN 001736140017 3614 official publication date 31 october 1997 109710 97 750 23798 the great basin naturalist PUBLISHED AT PROVO UTAH BY BRIGHAM YOUNG university

ISSN 001736140017 3614

VOLUME 57 31 OCTOBER 1997 no 4

creatgreat basin naturalist 574 0 1997 appp 283 314

MAYFLY FAUNA OF NEW MEXICO

WP McCaffermccaffertylMcCaffertymccafferty1tyl1 CRC R lugo ortizl and GZG Z jacobiajacobi2

ABSTRACT an inventory and analysis of the mayfly insecta ephemeroptera fauna of new mexico based on the published literature and study of extensive matematerialsnalsnais from diverse collections indicate the pieplesencepresencesencesenee of 12 families 37 genera and 81 species of these species 25 presentrepresentle new state records acentrella turturbidasurbidabida ameletusfalsusamelettisAmelettis falsusbalsus A sparsatussparsatus babtisbaetis adonis B bicaudatus B flaviflavistngaflavistrigastriga baetodesBaetodes deficiensdefiniens caeniscanniscaems latipennislatzpenmslatiiatipennis callibaetis fluctuantfluctuans cinygmula par ephemera simsimulantsimulansulans hexageniahexagemahexagyniaHexageniaagema bilbilineatebilineataineata Isonychia sicca labiobaetis propinquuspropinquous lachlaniaLachlania saskatchewanensis lepto phlebiaaphlebia bradleyi leucrocuta pelersipetersipetersi neochoroterpes nanita paraleptophlebia dabilisdebilisdebilis P heteroneaheteronea procloeon conturba tum rhithrogena plana R robusta R vitta and thraulodes gonzgonzalesigonzalesmalesi baetodesBaetodes deficiensdefiniens representslepieleplesents a new USA record for 37 of the 56 previously reported and confirmed species 124 new county records are provided with respect to continental affinities 34 species are western 27 southwestern 13 widespread I1 is a southern USA species and I1 eastern of the major drainagediainage systems in the state the gila system is the most species rich with 48 species followed by the rio grande 46 pecos 39 canadian 28 and san juan 25 relationships between drainage systems and between new mexico and broadly adjoining states are discussed lachlaniaLachlania dencyannaedency annae the only endemic species in new mexico occuisoccurs in the gila system and is lareiarerare and endangered certaingertain other species from the gila system are also noted as being at risk flomfrom other drainages B adonis Epheephemerellamerella mollitiamollitia and L pelersipetersi also aiealeare of some concern at the national level additional species that are rare in new mexico and are of concern at least at the state level include ametropusAmetropus albrighti C fluctuantfluctuans H bilineatebilineatabilineata L bradleyi N nanita P conturbatumconturbacontuicongui batumtum and R hagenihagemhagenl

key words ephemeroptera new mexico species inventory

the first report of a mayfly from new mex- mainly piecemeal first reports of species were ieoicoleo was that of ephemerellaEphemerella sp by needham contributed by 21 authors since the descrip- 1905 which was taken from the upper pecos tion of R undundulataungulataulata needham 1927 traver river in san miguel county this record later 1935 alienallenailen and edmunds 1959 1961 1963 proved to be applicable to the common west- 1965 peters and edmunds 1961 koss 1966 ern mountain species drunella grandis eaton 1972 alienallenailen 1968 1978 koss and edmunds the next species was recorded when banks 1970 kilgore and alienallenailen 1973 allenalienailen and chao 1924 described ephorus undulatesundulatus now rhithchithbhith 1978a 1978b morihara and mccafferty 1979a rogena undulataundungulataulata banks from the jemez river pescador and peters 1980 alienallenailen and murvosh in sandoval county the discovery of additional 1983 Kondratieffkondratieffandand voshell 1984 waltz and species of maymayfliesflies in new mexico has been mccafferty 1987 provonsha 1990 henry 1993

ldepartmentepaitmentottntomologyof entomology purduepin due umveiuniversityumpei sitygitsgity west lafayette IN 47907 nvnonmcntal2environniental science and management new mexico highlands university las vegasvcgasvagas NM 87701

283 284 GREAT BASIN naturalist volume 57

durfee and kondratieff 1995 lugoortizlugo ortiz and whole with respect to faunal elements within mccafferty 1995c zloty 1996 based on reports it importantly our data have served as the of these workers 56 species of maymayfliesflies have bases for evaluating a number of new mexico been known from new mexico prior to this species that can now be seen to be at environ- study mental risk either within the state or nation- six species of maymayfliesflies were described orig- ally A fine account of the history of new mex- inally from new mexico and thus have their icoleoieo s hydrologic setting beginning with the type localities within the state these species influences of the 12th century pueblo dwell are ametropusAmetropus albhghtialbnghti traver cloeodesCloeodes macro ing native americans can be found in the lamellus waltz and mccafferty homoeoneunahomoeoneuria fishes of new mexico by sublette et al 1990 allem pescador and peters lachlanialachlamaLachlanialamaiamaiania den cyannae koss rhithrogena undundulataungulataulata banks STUDY AREA and thraulodes brunbrunneusbrunneousneus koss only L den cyannae has proven to be endemic to new new mexico fig 1 is the ath5th largest state mexico in the USA covering some 195685 km2 it our interests in the new mexico mayfly possesses a diverse geologic and topographic fauna first developed in the mid 1960s when landscape with elevations ranging from slightly one of us WPM became involved in exten- less than 1050 m at red bluff reservoir in the sive collecting excursions across the state with southeastern part of the state where the pecos arwin provonsha and dick koss all of the river enters texas to over 4550 m at wheeler material from those trips eventually came into peak in the sangre de cristocastocnstogristo mountains the possession of WPM and purdue university approximately 50 km south of the colorado in the 1970s those collections and another border although new mexico is the ard3rd most collection made by WPM arwin provonsha and state in the USA it does receive 108176 and dan bloodgood in the early 1980s have hectare meters of water annually either as pre- contributed significantly to previous published cipitation or inflow hamsharris 1984 there is sig- revisionary studies of maymayfliesflies as well as the nificantnificant precipitation in some higher elevations present study in the late 1970s another of us while lower elevations afearearc typical of deserts GZJ became involved with the new mexico see eisenhood 1979 for example the eastern environment department and later joined the slopes of the sangre de cristogristo mountains and faculty of new mexico highlands university jemez mountains in the north central part of GZJ his students and colleagues have sampled the state receive over 180 cm of precipitation maymayfliesflies in new mexico for several years often per year mainly as snow mount taylor in the in association with various ecological studies western sector the mogollon mountains in the another of us CRL developed extensive exper- southwestern sector and the sacramento moun- tise on the systematics of southwestern maymayfliesflies tains in the south central sector of the state while conducting research on the maymayfliesflies of receive 80 120 cm of precipitation and south- mesoamericaMesomesoamencaamerica and his familiarity with the mex- ern valleys such as the rio grande and tula icanlean fauna has been invaluable in the assess- rosa receive 40 cm or less ment of new mexico five major and 3 minor drainage systems in the aforementioned collections of maymayfliesflies the southwestern USA have headwaters or along with materials loaned or donated to us near headwater flow in new mexico an inm from brigham young university colorado state depth treatment of these systems has been university and the university of utah have given by sublette et al 1990 however the provided a large body of material upon which following brief discussion will serve the imme-imme we have based much of the present study we diate purposes of this study also review all published data relevant to the the canadian river drainage system head- new mexico mayfly fauna provide extensive waters are in the northern sangre de cristo new site records for those species previously mountains the canadian river flows east- known from new mexico and provide new ward through the canadian escarpment from state records for an additional 25 species we the las vegas and raton plateaus into the offeloffer commentary on drainage distribution of northern panhandle of texas in the same area each of the new mexico species and synthe- of new mexico but north of the canadian the size all specific data and analyze the fauna as a dry cimarron river not to be confused with 199719971 NEW MEXICO MAYFLIES 285

C 0 L 0 R A D 0 ZM navajo res jnafr0t R TAOS dry COL AX 0 c0ca I1 K uan I1 reeler pk UNI ON RIOARRIBA f SAN 1 JUAN 0 chmchaahm

MORA HARDING MCKINLEY lalIAL S

conchas I1 res go SAN MIGUEL SANTA fetFEI SANDOVAL J ute res r r bernamBERNAU LLO santa rnarma res QUAY VAL NCIA GUADALUPE

I1 TORRANCE sumherresSumberresherRes 7 CURRY z DE BAC catron M 0 SOCORRO N ROOSEVELT LI NCOLN L

0 0 CHAVES oe elephant hondo buttebuffe res L

RRA si J1 I GRA NT caballo res 0 LEA

4 brantley rm OTERO L DON ANA EDDY

LUNA red bluff T E X A S res H I1 DAL GO M E X IC 0 chihuahua

fig 1 new mexico with mainmaln riverslivers drainage systems and counties indicated the cimarron river in colfax county which is cristo mountains the pecos river flows south- a tributary of the canadian river within new ward for about 640 km through the pecos plains mexico flows eastward off the trinidad escarp- of new mexico into west texas exiting new ment and the north slope of capulin mountain mexico in eddy county and ending with its mainly the northern tier of union county confluence with the rio grande at the texas these 2 river drainage systems are part of the mexico border larger arkansas river drainage system a major the rio grande the ath5th largest river in the south central tributary of the mississippi river USA has its headwaters in south central col- drainage system orado it flows almost directly south for over headwater streams of the pecos river drain- 720 km through new mexico the rio chama age originate from the southern sangre de in rio arriba county is a major tributary of 286 GREAT BASIN naturalist volume 57

the rio grande drainage system inm nornorthernthein MLHIODSMETHODS AND presentation new mexico two aveisnveisrivers in northwestern new mexico are extensive collections of new mexico may part of the middle Coloiacoloradodo river drainage flies were examined and material was identi- system the 640 km long san juan river with fied to species whenever possible collections headwaters in southern colorado flows for are noted in the data section with the follow- about 190 km in new mexico mainly san juan ing acronyms BYU brigham young university county to the foulfour cornelscorners aleaarea of utah col- collection provo utah CSU colorado state orado new mexico and arizonaanzona and then university collection fort collins colorado through extreme southeastern utah to lake NMHU new mexico highlands university powell and the Coloiacoloradodo river the small zuni collection las vegas new mexico and rivelriver which originates in the zuni mountains PERC purdue entomological research col- in new mexico south of the san juan river lection west lafayette indiana flows mainly inm southern mckinley county in the species accounts section of this work west to the little Coloiacoloradodo river in northern species are presented alphabetically by family aiarizonaizona genus and species species that are newly two main liversrivers in southwestern new mex- portedreportedle for new mexico are indicated with ico aiealeare paitpaltpart of the lower Coloiacoloradodo river drain- an asterisk for each species numerous data age system the san francisco river drainage are given first useful descriptions of adults system originatesongmatesongmates in eastern arizona but a and larval stages of the species are referenced intjorkaoimaoi portion of it flows in southwestern new under the heading descriptions such data are mexico gatloncationcatron county before returning to important because keys for the species are arizonaanzona where it joins the gila river head- generally not available and such descriptions waters of the gila river drainage system are are part of the basis for species identification located near mogollon baldy a peak over 3500 in most cases the original description is noted in high in the mogollon mountains the gila and if a subsequent more comprehensive and rivelriver flows foiroirolforrotror about 160 linkinkm mainly in grant useful description is available it is referenced county to the arizona bolderborder and continues also as are published keys when pertinent through southern arizonaanzona until it joins the historical names used in such works if differ- Coloiacoloradodo river in the extreme southwestern ent from the current name are also indicated cornelcorner of arizona the mimbresmembres river is also so that descriptive data can be easily tracked in southwestern new mexico grant and luna if the species is newly reported for new counties but is internallyiteimtei nallynaily drained mexico the complete locale data upon which four physiographicphysiogi aphie divisions provinces and the record is based are given under the head- plateaus are found in new mexico see snead ing new records the collection source acro- 1979 the canadian cimarronCimairon dry cimarronCimanon nym is always indicated first and if more than and pecos riversi ivel s aiaree found in the great plains one collection is involved data are presented provincepi oviceovmce these rivers have dissected lava alphabetically by collection source if the species capped plateaus and buttes and have their was a previously reported species then the lowellower courses on broad alluvial plains the previous distributional data are given in short lowerlowet two thirdsthnahn ds of the rio grande is in the form under the heading previous records by mexican highlands section of the basin and noting only the county and body of water name range plateau where upper tributariestributa ries are in and the bibliographic reference if we have isolated block faulted mountains separated by discovered additional county records then libroadoad plains the upper one third of the rio these are given again in the same short form crandegrande is in the rocky mountain province described above along with an indication of thelethere tributary stistreamsearnsearms in the jemez and south the collection source under the heading new sangiesanglesangre de cristogristo mountains aiealeare separated by county records because these data are often lava plateaus and alluvial plains the san juan extensive the short form of new county records zuni san franeiscofranciscoflanFiancisco and gila rivers are part of is used to conserve space however if the reader the intermontaneintel montane plateau where landformslandforms is interested in the entire locale information it consist of canyon and plateaus of sedimentary can be accessed from curators of the various 01or igneous rock collections indicated 1997 NEW MEXICO MAYFLIES 287

all species have a remarks section associated koss WP mccafferty and AXAV provonsha with them this generally includes a review of exuviae larvae the overall distribution of the species with REMARKS this species was described from particular attention to states surrounding new arizona by mcdunnough 1938 and previous mexico distribution of the species within new to this study has been known from only 3 mexico is also noted with respect to the drain- counties in that state see zloty 1996 new age systems involved counties and drainage mexico specimens of A falsusfalsnsfaluns agree with the systems may be located using figure 1 other alienallenailen and chao 1981 description ameletusofameletzisof pertinent taxonomic or environmental infor- sp C and it is possible that larvae of A sp B mation may also be included in the remarks allenalienailen and chao also belong to this species the faunistics section contains an analysis zloty 1996 ameletus sp B was reported from of the fauna in terms of its broad affinities and new mexico by alienallenailen and chao 1981 from inter and intrastate relationships species of rio arriba county in the upper rio grande special note because of their lantytarityrarity and enviedvienvi- drainage in the southern rocky mountain ronronmentalmental susceptibility are also noted here province thus in new mexico A falsusbalsus apparently occurs in tributariestributaries of the upper pecos SPECIES ACCOUNTS rio grande and headwaters of the river drainage system in the southern sangre de ameletidaeamelctidae cristogristo mountains ameletus doddsianusdoddsianus meletusamelefttsA sparsparsatussparsatnssafussatus zloty 1996 mcdunnough 1931 descriptions adult traver 1935 as A descriptions adult mcdunnough subnotatus eaton see also zloty 1996 larva 19311931aa see also zloty 1996 larva mcdun- alienallenailen and chao 1981 as A velox dodds nough1935nough 1935 PREVIOUS RECORDS zloty 1996 taos co NEW RECORDS NMHU otero co agua NEW COUNTY RECORDS CSU sierra co chiquita nr woods canyon 6 VIII 1980 GZG Z N fork palomas cr NMHU santa fe co rio jacobi and L smolka larvae enenmedioegmediomedio PERC grant co cherry cr REMARKS this western species was re- REMARKS zloty 1996 provided no details ported from alberta british columbia col- of his report of this species in taos county orado idaho and montana by zloty 1996 new mexico however one of us GZJ has lugoortizlugo ortiz and mccafferty 1995a reported collected material of this species from the it from arizona records of A aequivocusaeqwoocus same county at a rio hondo lake fork mcdunnough in colorado mccaffeitymccafferty et al beaver pond zloty also reported the species 1993 are also applicable to this species in from arizona colorado and utah certain new mexico A sparsatussparsatus has been taken only specimens reported as A subnotatus col- from the far southeastern part of the state in in pecos orado by mccafferty et al 1993 belong to the lower river drainage system this recently described species and the histor- ametropodidae ical confusion of what is now A doddsianusdoddsianus and A subnotatus was discussed by zloty 1996 ametropusAmetropus albrighalbnghtialbrightiti the occurrence of this species in new mexico traver 1935 in the gila river and upper rio grande descriptions adult alienallenailen and edmunds drainage systems lepresentsrepresents the southernmost 1976 larva traver 1935 see also alienallenailen and part of its known range edmunds 1976

1 PREVIOUS RECORDS san ameletusfalsusameletus falsusbalsus traver 1935 juan mcdunnough 1938 co san juan R REMARKS this psammophilous species was descriptions adult mcdunnough 1938 originally described from near farmington see also zloty 1996 larva alienallenailen and chao new mexico by traver 1935 since then it 1981 as A sp C has been reported only from the green river NEW RECORDS PERC san miguel co in southwestern wyoming and northeastern pecos R nr cowles 1 VIII 1944 larvae 222.22 2 utah edmunds and musser 1960 and the mi N of pecos pecos R 13 vilVII 1969 RWR W yampa river in northeastern colorado ailenalienalienallen 288 GREAT BASIN naturalist volume 57 and edmunds 1976 both of which are part of from all USA states bordering new mexico the upper colorado river drainage system in see mccafferty et al 1993 lugoortizlugo ortiz and new mexico it is known from the san juan mccafferty 1995a 19950 mccafferty et al river drainage system in the extreme north- 1997 however it has not been taken in mex- western corner of the state which is associ- ico in new mexico A turbidasurbida has been found ated with the middle colorado river drainage in the northern areas of both the rio grande system it remains to be seen if the species and pecos river drainage systems still exists in new mexico baetisbabtis adonis baetidaeBaetidae traver 1935 acentrella insignificantmsigmftcansinsignificans descriptions adult traver 1935 larva mcdunnough 1926 unknown NEW RECORDS PERC miguel co 5 mi descriptions adult mcdunnough 1926 N of pecosecos pecos R at dalton fishing site as babtisbaetis insignificantinsignificans mcdunnough oftecosofT larva 7300 ft 13 VII 1969 RW koss WP morihara and mccafferty 1979a as B mccaf- insigeinsig ferty and AV provonsha male and female ans AX nificansmficansnific adults PREVIOUS RECORDS morihara and mccaf- REMARKS poorly known is a ferty 1979a as baetisbabtis insignificanttnsigmficansinsignificans grant co this species small sized member of the rhodanirhodanic group that gila R mora co mora R san juan co san was previously known only from the san juan R durfee and kondratieff 1995 gabriel catron co taylor cr mountains in california traver 1935 its discovery in new mexico in the northern NEW COUNTY RECORDS CSU dona ana pecos co radium springs NMHU colfax co river drainage system therefore repre- a significant cieneguilla cr lincoln co rio ruidosoRuidoso sents range extension san miguel co pecos R rio arriba co rio baetisbabtis bicaudatus chama sierra co rio grande PERC catron dodds 1923 co cottonwood cr gila R taylor cr tula rosa R san francisco R san miguel co pecos descriptions adult dodds 1923 larva R sapello R dodds 1923 see also morihara and mccaf- REMARKS this relatively common west- ferty 19791979aa NEW RECORDS mora co s ern species is known from all mexican and NMHU jack USA states adjoining new mexico excluding cr 3065 m 18 VI 1991 MD hatch larvae oklahoma see mccafferty et al 1993 lugo- PERC catron co 42 mi N of silver city rt ortiz and mccafferty 1995a 1995b mccaf- 527 gila R at jet with little cr 15 VII 1967 ferty and lugoortizlugo ortiz 1996a in new mexico R and D koss female adult tularosaTularosa R 2 mi it is now known from throughout most of the above aragon ll11 VI 1974 B stark larvae state and all major drainage systems it is very lincoln co sierra blanca ski lodge rio rui common in catron county dosoboso 27 VIII 1976 MW sanderson larvae san miguel co Panchpanchuelauela cr 9 VII 1944 Acentrella turbidasurbida larvae santa fe co big tesuque cr big mcdunnough 1924 tesuque 10 VI 1974 B stark and TA wolff descriptions adult mcdunnough 1924a larva little tesuque cr nr hyde park 2960 as pseudocloeon turbiturbidumdum mcdunnough see m 20 IV 1973 B stark and T wolff larva also mccafferty et al 1994 larva mccaf- taos co rio trampas above el valle 9 VI ferty et al 1994 1974 B stark and TA wolff larva red NEW RECORDS NMHU taos co costilla river 3 mi E of questa 9 VI 1974 B stark cr 17 VIII 1989 GZG Z jacobi larvae PERC and TA wolff larvae san miguel co pecos R 29 VI 1939 larvae REMARKS this relatively widespread west- 5 mi N of pecos pecos R at dalton fishing ern species is found in mountainous streams site 7300 ft 13 vilVII 1969 RW koss WP and it reaches the southernmost limits of its mccafferty and AXA V provonsha female adults range in new mexico although common in pecos R I VIII 1936 female adult colorado mccafferty et al 1993 it has not REMARKS acentrella turbidasurbida is a wide- been taken in arizona mexico or texas in spread continental species that was recently new mexico it has been collected from all treated by mccafferty et al 1994 it is known major drainage systems except the san juan 199711997 NEW MEXICO MAYFLIES 289 river in the extreme northwestern part of the PREVIOUS RECORDS monharamondaramorihara and mccaf- state with the exception of alaska and parts of ferty 1979a as baetisbabtis sp B grant co cherry canada babtis bicaudatus is most commonly cr confined to higher elevations NEW COUNTY RECORDS NMHU colfax co canadian R sierra co main diamond baetisbabtis caecaelestiscoelestislestis cr torrence co canon de tajiquetadiqueTaj PERC catron alienallenailen and murvosh 1983 ique co glenwood gila R pueblo cr lincoln co descriptions adult unknown larva eagle cr rio ruidosoRuid oso morihara and mccafferty 1979a as B sp A REMARKS this western species was origi- PREVIOUS RECORDS morihara and mccaf- nally described as babtisbaetis sp B by monharamondaramorihara ferty 1979a as baetisbabtis sp A catron co san and mccafferty 19791979aa based on larvae from penasco francisco R otero co R new mexico and it was later named by mccaf- NEW COUNTY RECORDS BYU grant co ferty and waltz 1986 known from western sapillocapilloSapillo cr nebraska to panama lugoortizlugo ortiz and mccaf- REMARKS baetisbabtis caecoelestiscaelestislestis is a relatively ferty 1993 1996a it appears to be particularly common southwestern species presently common in arizona new mexico and col- known from arizona lugoortizlugo ortiz and mccaf- orado see also mccafferty and waltz 1986 ferty 1995a baja california alienallenailen and mur- durfee and kondratieff 1993 we have recently vosh 1983 chihuahua lugo and mccaf- lugoortizortiz identified larvae of this species collected by ferty 1996a and texas mccafferty and davis kondratieff and baumann in march 1993 from 1992 in new mexico where it was originally the san sabo river in menard county south- found by morihara and mccafferty 1979a it west central texas baitisbaetisbaetzs magnus is one of the is known from the gila lower pecos and san few species of maymayfliesflies now known to occur in francisco drainage systems in the south- river both the nearctic and neotropical biogeo- ern part of the state graphic regions in new mexico it is now baetisbabtisbaetisflavistrigaflaviflavistrigastriga known from all major drainage systems except mcdunnough 1921 the san juan river part of the middle colorado drainage system however has descriptions adult mcdunnough 1921 it been taken from the drainage larva ide 1937 see also morihara and mccaf- colorado system in ferty 19791979aa arizona and colorado NEW RECORDS PERC san juan co san baetisbabtis notos juan R river mile 165 1820 m 29 VI 1960 alienallenailen and murvosh 1987 larva REMARKS this relatively widespread north descriptions adult durfee and kon- american species is known mainly from the dratieffdiadratieff 1995 larva morihara and mccaf- eastern half of the continent but also from the ferty 1979a as B sp C black hills of south dakota mccafferty 1990 PREVIOUS RECORDS mouhartmouharamorihara and mccaf- and several localities in colorado see mccafferty 1979a as baetisbabtis sp C catron co gila ferty et al 1993 it was recently found in the R durfee and kondratieff 1995 catron co colorado drainage area of west central utah taylor cr mccafferty and macdonald 1994 mccaf- NEW COUNTY RECORDS NMHU colfax ferty and davis 1992 reported it from texas co Clenecieneguillacleneguillaguilla cr grant co gila R clene and lugoortizlugo ortiz and mccafferty 1994 found guillagmllacrcr PERC grant co gila R it in the adjoining mexican state of chi- REMARKS this southwestern species is huahua in new mexico B flaviflavistrigastriga has also known from arizona Monmoriharahara and mccaf- been taken only in the san juan river drain- ferty 1979a colorado ward and stanford age system 1990 and texas mccafferty and davis 1992 it was recently discovered in veracruz by lugo- baetisbabtis magnus ortiz and mccafferty 1994 and thus is ex- mccafferty and waltz 1986 pected to occur in northern mexico also dur- descriptions adult durfee and kon- fee and kondratieff 1995 reared this species dratieffdratieff 1993 larva morihara and mccaf- flomfrom taylor creek in catron county new ferty 1979a as B sp B mexico it has been collected in new mexico 290 GREAT BASIN naturalist volume 57

only in the cliaclidgilagild canadian and san francisco baetodesBaetodes edmundedmundsiedmundsosi river drainage systems koss 1972 babtisbaetis tncaudatustricaudatus descriptions adult koss 1972 larva dodds 1923 koss 1972 PREVIOUS RECORDS koss 1972 grant co descriptions 1923 adult dodds larva gila R dodds 1923 see also morihara and mccafmccaa NEW COUNTY RECORDS NMHU san mi- feity1979afertyberty 19791979aa guel co canadian R PREVIOUS RECORDS peters and edmunds REMARKS in addition to new mexico 1961 as baitisbaetisbaetzs intermedius san juan co san baetodesBaetodes edmundedmundsiedmundsosi has been reported from juan R morihara and mccafferty 1979a arizona koss 1972 texas edmunds 1950 san mora mora cationcatrongationgatlon co francisco R co and the mexican states of sinaloa and sonora penasco R otero co rio rio arriba co alienalienallenailen and murvosh 1987a new mexico it pecos in rio brazosBi azos san miguel co R has been taken only from the gila and cana- grant NEWNLW COUNIYCOUNTY RECORDS BYU co dian river drainage systems in the far south- gila in turkeytulkey cr R sapillocapillo cr lincoln co west and northeast regions of the state rio hondo NMHU colfax co canadian R respectively cieneguilla cr eddy co rio penasco giantglantgrant co black canyon cieneguilla cr guadalupe callibaetisferrugineuscallibaetis ferrugmeus hageni co pecos R lincoln co rio ruidosoRuidoso san- eaton 1885 doval co rio cebollabebollaCebolla san antonio cr santa descriptions adult eaton 1885 as C grande fe co rio sierra co main diamond hageni eaton larva dodds 1923 as cfuscacruscaC fusca cicr south diamond cr taos co red R rio dodds grant costilla PERC colfax co cimarron R PREVIOUS RECORDS peters and edmunds co cherry cr lincoln co eagle cr rio 1961 as callibaetis nigritus san juan co san ruidosoRuidoso sandoval co jemez R santa fe juan R goose co rio grande tesuque cr taos co NEW COUNTY RECORDS CSU catron co cicr la junta cr pueblo cr red R santa bar wall lake NMHU taos co beaver pond on baiarbara R rio hondo lake fork PERC eddy co sit- REMARKS this species is known from ting bull falls throughout most of north america including REMARKS this relatively widespread west- mexico lugoortizlugo ortiz and mccafferty 1994 ern subspecies ranges from alaska southward and it is probably the most widespread and to arizona lugoortizlugo ortiz and mccafferty 1995a ubiquitous baitisbaetisbcietis on the continent likewise and new mexico and it is one of the most it is now known from all major drainage sys- common maymayfliesflies in colorado where it was tems in new mexico treated under the synonym C americanosamericanusamericamencanusanus banks by mccafferty et al 1993 exten- baetodesBaetodes definiensdeficiensdeficzens the of the can be found in cohen and allenalienailen 1972 sive synonymy species in mccafferty and waltz 1990 and mccaffeitymccafferty descriptions adult unknown but sub 1996 callibaetis ferrugiferruginousneus hageni is not imagosimages were described by flowers 1987 known from texas and therefore arizona and larvalaiva cohen and alienallenailen 1972 new mexico apparently represent its south- NEW RECORDS BYU grant co sapillocapillo cr eastern range limits it has been taken in jet with gila R 1555 m 26 V 1985 B jensen ponds and lakes in wide ranging areas of new larvaelaivae mexico as well as from the san juan river REREMARKSMARKS this is essentially a mexican callibaetis fluctuans species it has been known from the mexican callibaetisfluctuansfluctuant 1862 states of guerrero jalisco molmoielosMoimorelosmoreioselos sonora walsh and veracruz see mccafferty and lugoortizlugo ortiz descriptions adult walsh 1862 as cloe 1996 its discovery in new mexico is impor- fluctuantfluctuans see also traver 1935 larva burks tant because it represents a new USA record 1953 in new mexico it has been taken only from NEW RECORDS BYU and PERC eddy the gila river drainage system in the inter- co castle cr black river village 141198714 1 1987 montane plateau baumann sargent and kondratieff larvae 199711997 NEW MEXICO MAYFLIES 291

REMARKS this species is known from turkey cr gila R lincoln co tucson mts across the continent but is most common in mckinley co thoreau sandoval co gahsgalis the midwest it has been taken as far west as teo cr san miguel co pecos R santa fe co california and oregon in the north but with Galigalistelgalisteosteo cr respect to areas surrounding new mexico it REMARKS this species occurs throughout has been reported only from colorado mccaf- western north america as far east as south ferty et al 1993 and texas lugoortizlugo ortiz and dakota mccafferty 1990 and texas mccaf- mccafferty 1995b the isolated sample of ferty and davis 1992 and south to costa rica this species from the extreme southern part of lugoortizlugo ortiz and mccafferty 1996b it is appar- the pecos river drainage system in new mex- ently one of the most common species of cal ico represents the southernmost limits of its libietislibaetislibaetis in new mexico where it has been taken western range in southern utah C fluctuantfluctuans in the gila river pecos river rio crandegrandegi ande and has historically been misidentified as C mon- zuni river drainage systems tanus eaton camelobaetidittscamelobaetidius musserimussen callibaetis montanus traver and edmunds 1968 eaton 1885 descriptiondescriptionss adult unknown laivalarva descriptions adult eaton 1885 larva traver and edmunds 1968 as dactylobaetis lugoortizlugo ortiz and mccafferty 1996b musserimussen traver and edmunds see also lugo- PREVIOUS RECORDS traver 1935 san juan ortiz and mccafferty 1995c co san juan R socorro co rio grande PREVIOUS RECORDRECORDSS lugo ortiz and NEW COUNTY RECORDS PERC catron co mccafferty 1995c catron co san francisco gila R dona ana co rio grande R W fork gila R gila R grant co E fork REMARKS callibaetis montanus has been gila R correctly reported from arizona and new REMARKS this species is widespread in mexico south to nicaragua eaton 1885 1892 mexico and central america traver and banks 1900 traver 1935 mccaffertymccaffeity and edmunds 1968 lugoortizlugo ortiz and mccafferty lugoortizlugo ortiz 1992 records of the species north 1995c lugoortizlugo ortiz and mccafferty 1995c of arizona and new mexico ege g edmunds have recently shown that C sakeussalinussaknus alienallenailen and 1954 newell 1970 rahel and kolar 1990 are chao which had been taken in the USA in evidently identificationsmismlsmisidentifications of C fluctuantfluctuans arizona and nevada is a junior synonym ofofcofaC and records of it in texas are probably all or musserimussen in new mexico the species appears mostly attributable to C punctilusus mccaf- to be restricted to the gila and sailsallsan francisco ferty and provonsha see mccafferty and pro- river drainage systems in the southwestern vonsha 1993 lugoortizlugo ortizandortiz and mccafferty 1994 portion of the state in new mexico it has been taken from diverse camelobaetidius warremwarrenl areas of the gila river san juan river and traver and edmunds 1968 rio crandegrande drainage systems descriptions adult traveltraver and edmunds callibaetis dictuspictus 1968 as dactylobaetis varreniwarremwarreni traveltraver and eaton 1871 edmunds larva traver and edmunds 1968 descriptions adult eaton 1871 as as D varreniwafwarreniwarwarremreni see also lugoortizlugo ortiz and mccaf- babtisbaetis dictuspictus eaton larva seemann 1927 as ferty 1995c C pacificus seemann see also lugoortizlugo ortiz and PREVIOUS RECORDS allenalienailen and chao 1978a mccaffeitymccafferty 1996b as dactylobaetis nauisnadisnavisnauls san juan co san juan PREVIOUS RECORDS traver 1935 as calli R as dactylobaetis tntritrltnvialistrivialistrivialismvialis colfax co cim- babtisbaetis pacificus sandoval co jemez springs arron R traver 1935 as callibaetis sigsignatusnatus socorro NEW COUNTY RECORDS CSU and PERC co Sasabinalsabmalpabinalbinalbinai cr catron co W fork gila R gila R san fran- NEW COUNTY RECORDS BYU grant co cisco R NMHU catron co san francisco turkey cr rio arriba co valerosVacvaceroseros canyon R grant co E fork gila R CSU sierra co circle seven cr PERC REMARKS this is the most common and catron co gila national forest chaves co widespread species of camelobaetidiuscamelohaetidius in the roswell grant co cherry cr sapillocapillo cr westeinwestern USA known from arizona california 292 GREAT BASIN naturalist volume 57

Coloiacoloradodo idaho and utah in addition to new labiobaetislabiobaetts apache mexico and it also ranges through mexico mccafferty and waltz 1995 sur guer- baja california chiapas chihuahua descriptions adult durfee and kondra rero oaxaca and sonora and central amer- tieff1997tieff 1997 larva mccafferty and waltz 1995 icaiealea lugoortizlugo ortiz and mccafferty 1995c it has PREVIOUS RECORDS peters and edmunds been known by many synonyms in the past as 1961 as baetisbabtis propinquuspropinquous walsh san juan shown by lugoortizlugo ortiz and mccafferty 1995c co san juan R wherein C navis allenailenalienallenalien and chao and C triv REMARKS this distinctive species was taltslalistaitslails alienalienallenailen and chao which had been reported recently described from the colorado drain- previously from new mexico were synonym age system of northern arizona and western izediced with it in new mexico C warrent has utah by mccafferty and waltz 1995 we have been taken from the canadian gila san juan acquired and studied the original material on and san francisco river drainage systems which peters and edmunds 1961 based their record of babtisbaetis propinquuspropinquouspropmquus now L propin Cloeodes macrolamellus cloeodes buusquus from the san juan river in new mexico waltz and mccafferty 1987 and it proved to be L apache records of L descriptions adult unknown larva propinquuspropinquousproptnquus from douglas creek near langelyrangelyRangely waltz and mccafferty 1987 colorado in the colorado drainage system mccafferty et al 1993 are also attributable to PREVIOUS RECORDS waltz and mccafmccaa L apache according to and kondratieff feity1987fertyberty 1987 grant co cherry cr durfee 1997 san juan river drainage system in REMARKS cloeodesClo eodes macrolamellus was the in new mexico is part of the middle colorado originally described from new mexico bbyy drainage system and thus L apache remains more remains waltz and mccafferty 1987 recently it known only from the greater colorado drain- was shown also to occur in the northern mexi- in age system in north america can states of chihuahua and durango lugo- ortiz and mccafferty 1994 and in the USA in labiobaetis propinquuspropinquous arizona lugoortizlugo ortiz and mccafferty 1995a walsh 1863 in new mexico it is known only from the cilacliagilaglia descriptions adult walsh 1862 as cloe river drainage system dicinaulcina walsh neenecneenec hagen see also morihara and mccafferty 1979b fallceonFallceon quillenquilleri as babtisbaetis propinquuspropinquous walsh 1940 as babtisbaetis dodds 1923 larva berner spin esusosus mcdunnough see also morihara and descriptions adult dodds 1923 as mccafferty 1979b as B propinquuspropinquous baetisbabtis quillenquilquliletiietileri dodds see also traver 1935 NEW RECORDS PERC colfax co bonilponil larva morihara and mccafferty 1979a as B cr 454 5 mi E cimarron 14 VII 1969 RWR W koss quillenquilleri see also lugoortizlugo ortiz et al 1994 WP mccafferty AXAV provonsha larvae otero penasco PREVIOUS RECORDS morihara and mccaf- co rio 12 VII 1969 RWR W koss WP ferty 1979a as baetisbabtis quilleriquilleri catron co gila mccafferty AXA V provonsha larvae R grant co cherry cr sapillocapillo cr REMARKS this primarily central and eastern USA evidently its western- NEW COUNTY RECORDS BYU eddy co species attains most range limits eastern new where castle cr CSU sierra co palomas cr in mexico we have seen it from the canadian and pecos PERC chaves co penasco ana rio dona river drainage systems labiobaetis co small tributary of grande eddy co propinquuspropinquous rio was reported from texas by mccafferty and castle cr lincoln co ruidosoRuidoso rio davis 1992 but is not known from mexico REMARKS variable is this highly species is published records of this species by peters widespread in central america mexico and and edmunds 1961 from the san juan river southwestern central and eastern USA lugo- in new mexico are attributable to L apache ortiz et al 1994 in new mexico it has thus see above far been taken only from the southern part of procloeon conturbatumconturbatum the state in the gila river pecos river and mcdunnough 1929 rio crandegrande drainage systems it is however known from the colorado riserrivelriver drainage sys- descriptions adult mcdunnough 1929 tem in colorado mccafferty et al 1993 as centroptilum conturbaconturbatumtum mcdunnough 199719971 NEW MEXICO MAYFLIES 293

see also lowen and flannagan 1991 as CG where it has been taken only in the lower pecos conturbatumconturbatum larva lowen and Flflannaganarmagan river drainage system was expected 1991 as C conturbatumconturbatum NEW RECORDS PERC san juan co san ephemerellidae juan R t32n r6wraw sec 27 river mi 165 Attenella margaritamarganta 1820 m 29 VI 16 VII 1960 WL peters male needham 1927 adults REMARKS specimens from the san juan descriptions adult mcdunnough river drainage system of northeastern new 1931b as ephemerellaEphemerella margaritamarganta needham mexico agree with the redescription of this larva needham 1927 as E matmarmargaritaganta see species by lowen and flannagan 1991 this also alienallenailen and edmunds 1961 as E mar species is known from western canada see garitaganta lowen and flannagan 1991 and in the USA PREVIOUS RECORDS alienallenailen and edmunds from california and wyoming traver 1935 1961 as ephemerellaEphemerella margaritamarganta san miguel co pecos and utah edmunds 1954 it represents a sig- R nificantnificant addition to the new mexico mayfly NEW COUNTY RECORDS NMHU taos co fauna costilla cr REMARKS western populations of this caenidaeCaenmaenidaeidae species range from british columbia to new caeniscannis bajaensis mexico alienalienallenailen and edmunds 1961 argyle and edmunds 1962 has alienallenailen and murvosh 1983 it not however been taken from arizona or texas in new mexico it is now descriptions adult provonsha 1990 known from the upper pecos river and upper larva alienallenailen and murvosh 1983 see also rio grande drainage systems this northern provonsha 1990 region in new mexico represents the south- PREVIOUS RECORDRECORDSS provonsha 1990 ernmost limits of the range of this species catron co gila national forest grant co cherry cr drunella coloradensis NEW COUNTY RECORDS NMHU mora co dodds 1923 wolfwolfcrwolfercr descriptions adult dodds 1923 as REMARKS caeniscannis bajaensis is also known ephemerellaEphemerella coloradensis dodds larva dodds from arizona colorado northern mexico and 1923 as E coloradensis see also alienallenailen and nebraska provonsha 1990 larvae of this edmunds 1962 as E coloradensis species are apparently restricted to sandy PREVIOUS RECORDS needham 1927 as bottomed areas of streams with slow to moder- ephemerellaEphemerella coloradensis san miguel co ate current in new mexico it is now known pecos R alienallenailen and edmunds 1962 as ephem from the canadian and gila river drainage erella coloradensis sandoval co rock cr taos systems co santa barbara R REMARKS caeniscannis latilatipennispennis this relatively common west- banks 1907 ern mountain species ranges from alaska to arizona and new mexico alienallenailen and edmunds descriptions adult banks 1907 see 1962 it apparently is restricted in new mex- also provonsha 1990 larva provonsha 1990 icoleoieo to the northern mountain regions of the NEW RECORDS NMHU eddy co black pecos river and rio grande drainage systems R at higby hole 7 VIII 1991 ST pierce larvae eddy co delaware R above pecos R drunella doddsi confluence 5 villVIII iggi1991 GZ jacobi larvae needham 1927 REMARKS the range of this species descriptions adult needham 1927 as extends across canada and the USA to south- ephemerellaEphemerella doddsi needham larva need- ern mexico provonsha 1990 lugoortizlugo ortiz and ham 1927 as E doddsi see also alienallenailen and mccafferty 1996a with respect to areas adja- edmunds 1962 as E doddsi cent to new mexico C latilatipennispennis has been PREVIOUS RECORDS traver 1935 as reported from colorado mccafferty et al 1993 ephemerellaEphemerella doddsi santa fe co santa fe R oklahoma traver 1935 and arizona and texas alienallenailen and edmunds 1962 as ephemerellaEphemerella provonsha 1990 its presence in new mexico doddsi san miguel co pecos R 294 GREAT BASIN naturalist volume 57

NEWnl w COUNJYCOUNTY RECORDS NMHU catron altana has been collected from the canadian co gila R rio arriba co rio brazosBi azos san river pecos river and rio grande drainage miguel co gaigallinasGalhnas R sapello R taos co systems and we provide the first records of it rio Fernfernandocindolundocundo de thostaos PERC taos co rio from the gila river drainage system in ari- hondo canyon zona it has been collected from the middle remai1ksrl MARKS new mexico is the southern- colorado river drainage system inm the north- most langeiangerange limit of this common western eastern part of that state and therefore it may species allenailenalienalienallen and edmandsedmnndsedmunds 1962 it is not eventually be found in all major drainages in known flomfrom alionaallonaarizona or texas in new mexico new mexico it has been taken fromfi om the canadian river gila Epheephemerellamerella riverriven pecos rivelriver and olandecrandeglandegrande diadradrainage inermiainermis rivel rio inage 1884 systems eaton descriptions adult eaton 1884 larva drunella grandis grandis traver 1935 see also alienallenailen and edmunds eaton 1884 1965 and johnson 1978 DESCdl sc ribRIPTIONsRIP I1 lonsIONS adult needham 1927 as PREVIOUS RECORDS traver 1935 san- epheinerellaephemerellaEphemerella grandis eaton larva eaton 1884 doval co jemez springs san miguel co as epheinerellaephemerellaEphemerella sedis incerincertaetae nymph 11 pecos R peters and edmunds 1961 san see also alienallenailen and edmunds 1962 as E gran- juan co san juan R alienallenailen and edmunds dlyadly grandis 1965 taos co la junta cr PREVIOUSPRLVIOUS RECORDSreoRLC ORDS needham 1905 as NEW COUNTY RECORDS BYU hondo co ephemerellaEphemerella sp r san miguel co pecos R hondo R NMHU colfax co cieneguilla needham 1927 as epheinerellaephemerellaEphemerella grandis santa cr lincoln co rio ruidosoRuidoso PERC catron fe co santa fe R kilgore and alienallenailen 1973 co gila R colfax co cimarron R grant co as ephemerellaEphemerella grandis lincoln co rio rui gila R lincoln co eagle cr rio ruidosoRuidoso dosoboso mora co mora R rio arriba co rio bra- nlNEWw COUNTY RECORDSRLCORDS BYU colfax co zos san miguel co pecos R meimelverlyeriveimeoVeiveri meoriejofiejo R NMHU colfax co cieneguilla REMARKS ephemerellaEphemerella inermisinerrnisinermia is a rela- cicr sierra co main diamond cicr taos co tively common species that occurs throughout costilla cicr red R PERC taos co rio much of western north america excluding pueblo de taos mexico alienallenailen and edmunds 1965 using only REMARKSRLMARKS this subspecies of this com- the keys of alienallenailen and edmunds 1965 one mon western species occurs in arizona col could easily confuse specimens of the mainly orado01 ado new mexico utah and wyoming ailenalienalienallen eastern species E dorothea needham which and edmunds 1962 in new mexico it occurs we have seen in texas with this strictly west- in mountain tributatnbutanestributariesriesrles mainly of the cana- ern species there are however unpublished dian river pecos rivelriver and rio grande drain- discriminating characters in the larvae that age systems in westeinwestern diadladrainagesmages it is known can be used to separate the 2 species it is for only fromfi om gila river drainage headwaters this reason that impending records of E mer- mismls from texas further substantiation ephemerellaEphemerella altana mis require in new mexico it is known from all alienallenailen 1968 major drainage systems descriptionsSCRJPHONS adult alienallenailen 1968 laivalarva dl Epheephemerellamerella infrequentinfrequens alienallenailen 1968 mcdunnough 1924 PREVIOUSPRI vious RECORDSRLCORDS alienallenailen 1968 colfax co molenomoreno cicr moiamolamora co mora R san descriptions adult mcdunnough miguel co pecos R taos co rio grande 1924b larva traver 1935 see also alienallenailen rio hondo rio pueblo and edmunds 1965 and johnson 1978 NEWnl w COUNIYCOUNTY RECORDSRLCORDS PERC catron PREVIOUS RECORDS alienallenailen and edmunds co gila R grant co gila R 1965 taos co red R la junta cr rlri MARKS this species has been known REMARKS this western species does not flomfrom arizonaanzona and new mexico alienallenailen 1968 occur in arizona or mexico and its only records and lugoortizlugo ortiz and mccafferty 1994 reported in new mexico are from the northern part of it hornfromhormhomm baabaja califoiniacalifornia in new mexico E the state in the upper rio grande drainage 199719971 NEW MEXICO MAYFLIES 295 system we have not seen material of this PREVIOUS RECORDS alienallenailen andcind edmunds species from new mexico and thereforetherefoieboiebole can- 1963 as ephemerellaEphemerella tibialis taos co red R not be sure that previous reports were indeed REMARKS this species occurs throughout of E infrequentinfrequens rather than E inermis with the western USA and canada allenailenalienalienallen and which it has commonly been confused in the edmunds 1963 but it has rarely been taken past see johnson 1978 in arizona and new mexico in new mexico S tibialis has been known only from the upper ephemerellaEphemerella mollitiamollitia grande drainage system the seemann rio in rocky 1927 mountain province descriptions adult seemann 1927 timpanogaTimpanoga hecuba hecuba seemann 1927 see also alienallenailen and larva eaton 1884 edmunds 1965 PREVIOUS RECORDS traver 1935 san descriptions adult mcdunnough 1935 juan co san juan R as ephemerellaEphemerella hecuba eaton larva eaton REMARKS this species has been known 1884 as ephemerellaEphemerella sedis incertaeincertncertaefaetae nymph only from california and the northwestern cor- 4 see also alienallenailen and edmunds 1959 as E ner of new mexico san juan river drainage hecuba hecuba system alienallenailen and edmunds 1965 ques- PREVIOUS RECORDS alienallenailen and edmunds tioned the validity of traver s 1935 record 1959 as ephemerellaEphemerella hecuba hecuba san stating that she may have actually based the miguel co pecos R record on larvae of E inermisinermia we have not NEW COUNTY RECORDS NMHU rio arriba seen the material and therefore can neither co rio chama confirm nor refute the record at this time REMARKS this western species which is conspicuously absent from most of the great serratella mimichcheneneneri basin and arizona alienallenailen and edmunds 1959 traver 1934 reaches the southernmost part of its range in descriptions adult traver 1934 as northern new mexico where it is now known pecos ephemerellaEphemerella michmicheneneneri traver larva traver from the upper river and upper rio 1934 as E mimichcheneneneri see also alienallenailen and grande drainage systems edmunds 1963 as E mimichcheneneneri ephemeridae PREVIOUS RECORDS peters and edmunds 1961 as ephemerellaEphemerella chenenmimichenermichenerimichenenmicheneri san juan co ephemera simsimulantsimulansshnulansulans san juan R alienallenailen and edmunds 1963 as walker 1853 ephemerellaEphemerella chenenmimichenermichenenmichenerimicheneri rio arriba co san descriptions adult walker 1853 see juan R kilgore and alienallenailen 1973 as ephem also traver 1935 larva ide 1935 see also erella chenenmimichenermichenenmichenerimicheneri catron co negrito cr col- mccafferty 1975 fax ponilpoml grant co bonilpomi cr co little cr NEW RECORDS NMHU rio ambaarriba co rio NEW COUNTY RECORDS NMHU lincoln chama hwybwy 846484 64 bridge 12 VI 1981 GZG Z co rio ruidosoRuidoso sandoval co rio cebollabebollaCebolla jacobi female subimago santa fe co rio grande REMARKS this widespread north ameri- REMARKS this western species is known can burrowing species was shown by mccaf- from arizona baaraabaja california california col- ferty 1975 to occur from the east coast to orado new mexico oregon washington and idaho and utah it is not known from arizona wyoming alienalienallenailen and edmunds 1963 but it is mexico or texas the new state record from absent from idaho nevada and utah essen- the northern portion of new mexico in the tially the great basin in new mexico it upper rio grande drainage system was some- occurs in all major drainage systems what unexpected given the distribution pattern of this mccafferty 1994 serratella tibialis species mcdunnough 1924 hexagyniahexageniaHexagenia bilineatebilineatabilineata descriptions adult mcdunnough say 1824 1924b as Epheephemerellamerella tibialis mcdunnough DEdescriptionsSCRIPTIO N S adult say 1924 as baetisbabtis larva walley 1930 as E tibialis see also bilineatebilineatabilineata say see also spieth 1941 larva alienallenailen and edmunds 1963 as E tibialis clemens 1913 see also mccafferty 1975 296 GREAT BASIN naturalist volume 57

NEW RECORDS eddy co black R 7 mi W ephorusepeorus albertae malaga 9 VII 1992 SAS A sanders male adult mcdunnough 1924 S gonzales 7 VIII 1992 SAA sanders C GXG Z descriptions adult mcdunnough 1924a jacobi larvae as iron albertae mcdunnough larva edmunds REMARKS the previous obscure record of and alienallenailen 1964 1941 this species by spieth from the red PREVIOUS RECORDS alienallenailen and chao river ostensibly in new mexico which was 1978b rio arriba co rio chama further recorded by mccafferty 1975 was REMARKS ward and berner 1980 indi- apparently not valid this conclusion is based cated that this western species tended to on the fact that the only red river in new replace other species of ephorusepeorus at lower ele- mexico is in taos county and that stream does vationsvations ephorusepeorus albertae is relatively common not have siltmarlsilt marl substrate that would be in colorado mccafferty et al 1993 but it has required by hexageniahexagemahexagyniaHexageniaagema larvae see keltner and not been taken in arizona mexico or texas mccafferty 1986 furthermore the material in new mexico it has been taken only in the on which spieth 1941 based his new mexico northern part of the state in the upper rio record was from the oklahoma natural his- grande drainage system tory survey and the red river along the okla ephorusepeorus deceptiousdeceptivus homa texas border in the eastern parts of those states does possess substrate types required mcdunnough 1924 by hexageniahexagemahexagyniaHexageniaagema larvae nonetheless we do pro- descriptions adult mcdunnough 1924a vide the first valid record of this species in new as iron deceptivusdeceptious mcdunnough larva mexico the male adult from eddy county edmunds and alienallenailen 1964 new mexico lower pecos river drainage sys- PREVIOUS RECORDS alienallenailen and chao tem clearly possesses the beaklike penes typ- 1978b san miguel co pecos R ical of H bilblibihbilineatabihneatabilineateneataineata however both the larvae NEW COUNTY RECORDS NMHU sierra and adult are very lightly pigmented which is co south diamond cr PERC taos co rio atypical of H hllhiibilbihbilineatabihneatabilineateneataineata although H bilbihbibbilineatabihneatabilineateneataineata hondo is known from texas and oklahoma it has not REMARKS edmunds and alienallenailen 1964 been found north west or south of new mex- indicated that this species was confined to the ico with the exception of a record from the intermountain western USA and those authors mexican state of san luis potosi by mccaf- were the first to indicate its occurrence in ferty 1968 the most widespread species of new mexico but they did not provide any hexageniahexagemahexagyniaHexageniaagema in north america H limbatalimbaga ser- specific locale data alienallenailen and chao 1978b ville has not been found in new mexico but recorded it from gila county arizona in new may eventually because it does occur in col- mexico we know of this species from the orado and texas upper pecos river and upper rio grande drainage systems as well as the gila river drainage heptagenndaeheptageniidae system in the southwestern part of the state cinygmulacmygmula par ephorusepeorus longimanuslongilongimanousmanus eaton 1885 eaton 1885 descriptions adult eaton 1885 as descriptions adult eaton 1885 as iron cinygma par eaton larva not published longimanuslongilongimanousmanus eaton larva dodds 1923 as iron NEW RECORDSIPECORDS NMHU san miguel co nymph 1 see also edmunds and alienallenailen 1964 elk mtnman elev 3540 m below springs VII PREVIOUS RECORDS alienallenailen and chao 198019805 LRL R smolka male and female adults 1978b rio arriba co Canjicanjilonlon cr san REMARKS this species occurs in the higher miguel co pecos R winsor cr taos co red elevations throughout much of western north R rio pueblo rio chiquito durfee and america including arizona eaton 1885 and kondratieffkondratieff19951995 catron co taylor cr colorado dodds 1923 it is not known from NEW COUNTY RECORDS NMHU lincoln mexico or texas in new mexico C par is the co rio ruidosoRuidoso PERC lincoln co eagle only species of the genus thus falfaifarhar taken and it cr has been found only in mountain headwaters REMARKS ephorusepeorus longimanuslongilongimanousmanus is the most of the pecos river widespread of the western species of ephorusepeorus 199719971 NEW MEXICO MAYFLIES 297

edmunds and alienallenailen 1964 it is very common heptageniaHeptagenia solitariosolitariasolitaria at higher elevations in colorado and utah but mcdunnough 1924 somewhat less common in arizona and new descriptions adult mcdunnough 1924a mexico in new mexico it has been taken larva bednarik and edmunds 1980 from the gila river upper pecos and river PREVIOUS RECORDS peters and edmunds upper grande drainage systems rio 1961 san juan co san juan R ephorusepeorus margarita NEW COUNTY RECORDS PERC catron co edmunds and alienallenailen 1964 gila R colfax co canadian R cimarron R grant co gila R mora co coyote san- descriptions cr adult edmunds and alienallenailen doval co rio grande 1964 edmunds and alienallenailen 1964 larva REMARKS mccafferty et al 1993 noted PREVIOUS RECORDS durfee and kondra that this co in colorado western species was found tiefftieff19951995 catron taylor cr primarily on the western slope of the colorado NEW grant COUNTY RECORDS PERC co plateau is one of the gila co it most common species R rio arriba rio brazos of heptageniaHeptagenia in the west and has also been REMARKS we have numerous samples of found in the gila river and middle colorado epeorus from ephorus margarita catron county col- drainage systems in arizona lugoortizlugo ortiz and lected in the 1960s and 1980s the species is mccafferty 1995a it is not known from mex- epeorus by far the most common species of ephorus ico or texas in new mexico it has been taken gila found in the river drainage system and in the canadian river gila river rio grande except for the rio brazos a headwater stream and san juan river drainage systems in the upper rio grande drainage system it has not been taken from the other major drain- leucrocuta pelersipetersi age systems in the state ephorusepeorus margarita alienalienallenailen 1966 was recently reported from numerous locali- descriptions adult alienallenailen 1966 as hep ties in the mexican state of chihuahua by tageniahagenia pelersipetersi alienallenailen larva bednarik and ed- lugoortizlugo ortiz and mccafferty 1996a and the munds 1980 as H pelersipetersipetersi species has been known from a few localities NEW RECORDS NMHU colfax co six in arizona edmunds and alienallenailen 1964 and mile cr nr hwybwy 64 2530 in 9 IX 93 GZ baja california allenailenalienalienallen and murvosh 1983 jacobi larvae PERC catron co 2 mi N sil- heptageniaHeptagenia elegantulaelegantula ver city rt 527 gila R at jet with little cr eaton 1885 15 VII 1967 R and D koss female adults gila cliff dwellings national monument gila descriptions adult eaton 1885 as rhi R at jetjctact with little cr 15 VII 1967 R and D ththrogenarogena elegantulaelegantula eaton larva needham koss male and female adults N silver city and christenson 1927 as R elegantulaelegantula see middle fork gila R 2525.25 mi above jet with W also bednarik and edmunds 1980 fork 9 VII 1969 RW koss WP mccafferty PREVIOUS RECORDS peters and edmunds and AXAV provonsha female adult grant co 1961 san juan co san juan R alienallenailen and 65 km N silver city rt 527 E fork gila R 15 chao 1978b colfax co cimarron R bonilponil 16 VII 1967 R and D koss larva cr REMARKS this species was previously REMARKS this species is known from known only from the green river upper col- throughout much of western north america orado drainage in southwestern wyoming and including arizona and colorado eaton 1885 northeastern utah alienallenailen 1966 new records and utah edmunds 1954 it has not been from new mexico are therefore significant in found in mexico oklahoma or texas larvae new mexico L pelersipetersi has been taken in the of heptageniaHeptagenia elegantulaelegantula can be distinguished canadian and gila river drainage systems it from the somewhat similar H solitariasolitariosolitaria see is possible that this species presently occurs below with the use of the key by bednarik only in new mexico because of its possible and edmunds 1980 the species prefers silted extirpation in utah and wyoming streams at lower elevations and in new mexico it is known only from the canadian and nixe criddlcriddleiei mcdunnough 1927 san juan river drainage systems in the northern part of the state descriptions adult mcdunnough 1927 298 GREAT BASIN naturalist volume 57 as heptageniaheptagemaHeptageniaagema criddlcnddleicriddleiei mcdunnough larva utah needham and chnstensonchristenson 1927 and it bednarikbednarlk and edmunds 1980 as H criddlcnddleicriddleiei is relatively common inm southern areas of col- PREVIOUSPRLVIOUS RECORDS peters and edmunds orado ege g mccafferty et al 1993 in arizona 1961 as heptageniaheptagemaHeptageniaagemaogema sp san juan co san and new mexico R hagenihagem is known from juan R alienallenailen and chao 1978b as heptaceheptage only a single locality in each state alienalienallenailen and niama criddlcnddleicriddleiei catron co san franciscofraneiscoFianpianciscoelseo R col- chao 1978b in new mexico it is known from fax co cimarronCimanon R molenomoreno cr grant co the upper rio grande drainage system sapillocapilloSapillo cicr lincoln co bonito cr cr at pine lodge moiamolamora co mora R rio arriba co rhithrogena morrisonimotmornsonmornsommorrisoni wolf cicr taos co rio hondo valencia co banks 1924 blue water cicr descriptions adult banks 1924 as NEWNLW COUNIYCOUNTY lii111CORDSRICORDS CSU sierra co ephorusepeorus morrisonimorrisonnmornsonmornsommorrisoni banks see also traver 1935 palomas cicr PERC sandoval co jemez R larva seemann 1927 as iron petpetulantpetulansulans see- pecos san miguel co R mann see also key by alienallenailen and chao 1978b REMARKS this relatively common species PREVIOUS RECORDS alienallenailen and chao 1978b is known flomfrom throughout the western USA colfax co cimarron R san miguel co pecos mcdunnough 1927 tiavertravergiaver 1935 edmunds R taos co red R rio chiquito 1954 peters and 1961 and edmunds alienallenailen NEW COUNTY RECORDS NMHU san juan chao 1978b nixe criddlcnddleiei has been taken co san juan R from all diadladrainageinage systems within new major REMARKS this species is known from much mexico of the west from alberta mcdunnough nixe simplicisimplicioidesoides 1934 to baja california in northern mexico mcdunnough 1924 lugoortizlugo ortiz and mccafferty 1994 with respect to the states adjacent to new mexico it is descriptionsSCRIP IONS adult mcdunnough 1924a dl lons known only from arizona see lugoortizlugo ortiz and as heptageniaheptagemaHeptagema simplicioidessimplicioides mcdunnough agenia mccafferty 1995a although mccafferty et al bednarikbednarlk and edmunds 1980 as H laivalarva 1993 did record mor from col- simplicioidessimplicioides not R morrisonimorrisonnmomsonmomsomrisoni orado they noted that materials commonly PREVIOUSPRLVIOUS RECORDSblobeoRLC ORDS peters and edmunds identified as R undulataundungulataulata banks in colorado 1961 as heptageniaheptagemaHeptageniaagema simplicisimplicioidesoides san juan in and elsewhere probably referable to this co san juan R were new mexico it has been taken in NEWNLW COUNIYCOUNTY RECORDSRLCORDS BYU grant co species in in the of the gila R turkey cr sandoval co jemez R northern part the state in canadian pecos grande san NMHU rio arriba co rio chama PERC river river rio and juan catron co gila R lincoln co bonito cr river drainage systems moiamolamora co coyote cr san miguel co pecos rhithrogena plana R alienallenailen and chao 1978 RLREMARKSMARKS this species is known from the westeinwestern USA traver 1935 edmunds 1954 descriptions adult unknown larva peters and edmunds 1961 alienallenailen and chao alienallenailen and chao 1978b 1978b19781 bednarik and edmunds 1980 and NEW RECORDS PERC grant co duck albeltaalberta mcdunnough 1924a previously it cr at cliff US 180 5 V 1981 WPWE mccafferty was known flomfrom new mexico only from the AXA V provonsha and D bloodgood larva gila san juan river diadladiainagedrainageinage system in the north- R at US 180 nr cliff 5 V 1981 WPWE mccaf- west coinercolnercorner of the state we report it from all ferty AXA V provonsha and D bloodgood larva major drainagediadlainage systems in new mexico taos co S fork of red R carson national forest 27 VII 1937 C tarzwell larvae rhithrogena hagenihagemhagenl REMARKS this species was previously 1885 eaton known only from arizona alienalienallenailen and chao descriptions adult eaton 1885 larva 1978b where it was described from the white not described but keyed by day 1963 river in navajo county we report it for the PREVIOUSPRLVIOUS RECORDS alienallenailen and chao first time from new mexico from the gila river 1978b taos co red R and rio grande drainage systems mccafferty rlREMARKSMARKS this western USA species is et al 1993 indicated that R plana may even- also known from colorado dodds 1923 and tually prove to be a variant of R morrisonimornsonmornsommorrisoni 199719971 NEW MEXICO MAYFLIES 299

rhithrogena lobustarobusta rhithrogena vitta dodds 1923 alienallenailen and chao 1978 descriptions adult dodds 1923 larva descriptions adult unknown larva dodds 1923 see also key by alienallenailen and chao alienallenailen and chao 1978b 1978b NEW RECORDS BYU grant co gila R NEW RECORDS NMHU santa fe co rio above mouth turkey cr 1455 in 27 V 1985 B en medio 2990 in 13 V 1992 GXG Z jacobi lar- jensen larvae PERC grant co E fork of vae PERC grant co I1 mi S cliff tributary gila R at rd 15 gila national forest 6 V to gila R 14 VII 1967 R and D koss larvae 1981 WP mccafferty AV provonsha and D taos co creek in hondo canyon nr taos ski bloodgood larvae valley 22111196722 iliillIII111 1967 RWR W baumann larvae REMARKS this species was previously REMARKS the new new mexico state known only from arizona where it was de- records of the distinctive R lobustarobusta represent scribed from the white river in apache and the southernmost limits of the known range of navajo counties middle colorado drainage this boreal western species in new mexico it system by alienallenailen and chao 1978b in new has been taken in the gila river and upper mexico we found it in the southwestern por- grande drainage co with rio systems it occurs tion of the state in the gila river drainage sys- R plana grant county where the in species tem also a part of the greater colorado river are distinct from each other cohabitation by drainage system rhithrogenarhtthrogena species is often common leonard and leonard 1962 mccafferty et al 1997 isonychiidae rhithrogenarhzthrogena undulataundungulataulata Isonychia interintermediaintermedialmedia banks 1924 eaton 1885 descriptions adult banks 1924 as epe descriptions adult eaton 1885 as chi orus undulatusundulates banks larva not published rotonrotonetesetes intermedius eaton larva kondra- but keyed by flowers and hilsenhoff 1975 tieff and voshell 1984 and alienallenailen and chao 1978b PREVIOUS RECORDRECORDSS kondratieff and PREVIOUS RECORDS banks 1924 as epe voshell 1984 catron co san francisco R orus undulatesundulatus sandoval co jemez R peters dona ana co rio grande grant co gila R san and edmunds 1961 san juan co juan NEW COUNTY RECORDS NMHU socorro R alienallenailen and chao 1978b colfax co cimar- co grande pecos rio ron R san miguel co R taos co REMARKS Isonychia interintermediaintermedialmedia is known red R from several localities in arizona and chi- NEW COUNTY RECORDS NMHU rio ambaarriba huahua kondratieff and voshell 1984 lugo- co PERC grant co gila R rio chama ortiz and mccafferty 1995a in addition to arriba co brazos rio rio those localities in new mexico noted above REMARKS rhithrogena undundulataungulataulata is a west- in new mexico it has been taken from the ern and central north american but species it gila grande and san was originally described from new mexico river rio francisco drainage systems the southern part of banks 1924 with respect to states adjaceadjacentnt river in the we do to new mexico the has been reportereportedd state not expect it to be found in species either rocky great plains from arizonaanzona ailenalienalienallen and chao 1978b and col-Col01 the mountain or orado peters and edmunds 1961 but it is provinces not known from mexico oklahoma or texas Isonychia sicca mccafferty et al 1993 indicated that this walsh 1862 species was confined to the western slope of the colorado plateau and lugoortizlugo ortiz and descriptions adult walsh 1862 as mccafferty 1995a indicated that in arizona it baetisbabtis sicca walsh see also kondratieff and was restricted to the colorado river drainage voshell 1984 larva alienallenailen and cohen 1977 system however it is not confined to the san as I1 sicca manca eaton see also provonsha juan river drainage system middle colorado and mccafferty 1982 river drainage system in new mexico but is NEW RECORDS BYU sandoval co rio relatively widespread being found in all mainmaln grande coronado state park bernalillo 21 drainage systems vilivillVIII 1985 baumann and whiting larvae 300 GREAT BASIN naturalist volume 57

RLREMARKSMARKS Isonychia sicca is a relatively REMARKS tricorythodes condolusconcondylusdylus is a widespread species inm north america kon- southwestern species known from arizona dratieffdratieff and voshell 1984 being especially chihuahua new mexico and sonora ailenalienalienallen common in the midwest but extending to 1967 kilgore and alienallenailen 1973 lugoortizlugo ortiz and central america with respect to areas adja- mccafferty 1994 in new mexico it is known cent to new mexico it is common in texas only from the gila river drainage system lugoortizlugo ortiz and mccafferty 1995b but has also been reported from chihuahua lugo- tricorythodes corpulentcorpulentusus ortiz and mccafferty 1996a and oklahoma kilgore and alienallenailen 1973 mccafferty et al 1997 its discovery in new descriptions adult unknown larva kil- where is thus far mexico it is known only from gore and alienallenailen 1973 the rio grande drainage system is somewhat PREVIOUS RECORDRECORDSS kilgore and alienallenailen expected of I1 sicca and I1 larvae sicca campestriscampecampestnsstris 1973 catron co gila R mcdunnough of which the latter occurs in in REMARKS this species is known only from alberta saskatchewan wyoming northeast the gunnison colorado and utah and northwest colorado woodbury and river in ward stanford 1990 and gila new argyle 1963 cannot be distinguished confi- river in mexico mccafferty al 1993 dently kondratieff and voshell 1984 theref- et pointed out that T corpulentusus was only tentatively valid we ore although there is the possibility that lar- corpulent have this vae we have seen belong to I1 campestriscampestnscampestris the not seen specimens of species sandoval county record would appear more tricorythodes dimorphusdimdimorphousorphus consistent geographically with the range of I1 alienallenailen 1967 sicca also the I1 sicca identification is more consistent with larvae taken from the rio descriptions adult kilgore and alienallenailen grande yanoviak and mccafferty 1995 1973 larva alienallenailen 1967 showed that I1 sicca is typically found in very PREVIOUS RECORDS alienallenailen 1967 taos co long stream regions 200 km whereas I1 red R kilgore and alienallenailen 1973 catron co campecampestriscampestnsstris is known only from stream regions N glenwood grant co sapillocapilloSapillo cr sierra 100 200 kmkra in length co rio grande REMARKS this southwestern species is leptohyphidae known from arizona ailenalienalienallen 1967 kilgore and leptohyphes apache alienallenailen 1973 gray 1981 lugoortizlugo ortiz and mccaf- alienallenailen 1967 ferty 1995a chihuahua and new mexico lugo and mccafferty 1994 new descriptions adult unknown lugoortizortiz in larva mexico it has been taken from the gila and alienallenailen 1967 rio grande drainage systems PREVIOUSPRLVIOUS RECORDS alienallenailen 1967 rio arriba co taylor cr alienallenailen 1978 catron co tricorythodes explicatusexplicates san francisco R grant co gila R eaton 1892 REMARKS this southwestern species is descriptions known fromflom arizona and southern utah allenailenalienalienallen adult eaton 1892 as tri 1967 1978 lugoortizlugo ortiz and mccafferty 1995a corythus explicatesexplicatus eaton larva alienallenailen and and chihuahua lugoortizlugo ortiz and mccafferty murvosh 1987c 1994 in addition to new mexico in new PREVIOUS RECORDS alienallenailen and murvosh mexico L apache is known only from the inter- 1983 dona ana co trib rio grande montane plateau in the western part of the NEW COUNTY RECORDS NMHU colfax state including the gila san francisco and co cieneguilla cr six mile cr eddy co san juan river drainage systems penasco R lincoln co rio ruidosoRuidoso rio arriba co rio chama sierra co rio grande tricorythodes condylusdylus tncorythodes condoluscon PERC san juan co san juan R alienallenailen 1967 REMARKS this species is known from descriptions adult alienallenailen 1967 larva arizona and throughout northern mexico see alienallenailen 1967 lugoortizlugo ortiz and mccafferty 1995a mccaf- PREVIOUS RECORDS kilgore and alienallenailen ferty and lugoortizlugo ortiz 1996 in new mexico 1973 catron co gila R grant co sapillocapilloSapillo T explicatesexplicatus is relatively common and is now cr known from the canadian river pecos river 199711997 NEW MEXICO MAYFLIES 301

rio grande and san juan river drainage sys- drainage system in arizona colorado and tems lugoortizlugo ortiz and mccafferty 1995a in- utah it could eventually also be found in that dicated some possibility that T explicatusexplicates and drainage system in new mexico T minutusminutes traver were synonymous leptophlebia bradleyi tricorythodes minutesminutus needham1932needham 1932 traver 1935 descriptions adult needham 1932 descriptions adult traver 1935 larva larva berner 1975 kilgore and alienallenailen 1973 NEW RECORDS CSU eddy co sitting bull PREVIOUS RECORDS kilgore and alienallenailen falls 161199316 1 1993 S fitzgerald and S hoffman 1973 catron co san francisco R chaves larvae male adults co rio penasco colfax co bonilponil cr grant REMARKS this species is known from the co gila R lincoln co rio ruidosoRuidoso mora southeastern USA and texas henry and kon- co mora R rio arriba co rio chama san- dratieffdratieff 1982 here we report it from new doval co jemez R san juan co san juan mexico near texas in the lower pecos river R sierra co rio grande san miguel co drainage system the new mexico record rep- sapello cr santa fe co rio grande taos resents the westernmost limits of its presently co taos cr known range REMARKS tricorythodes minutesminutus is apparently widespread and ubiquitous in much of neochoroterpes kossl canada and the USA it has not been found in alienalienallenailen 1974 mexico or texas it was reported as common descriptions adult henry 1993 larva throughout much of colorado by mccafferty alienallenailen 1974 as choroterpes kossibossi alienallenailen see et al 1993 in new mexico kilgore and also henry 1993 alienallenailen 1973 have reported it from all major PREVIOUS RECORDS henry 1993 catron drainage systems although we have not seen co san francisco R this any material of species from the many col- NEW COUNTY RECORDS PERC grant co lections we have examined lugoortizlugo ortiz and cherry cr little cr mccafferty 1995a indicated that T minutesminutus REMARKS in addition to new mexico N could easily T be confused with explicatesexplicatus kossl has been known from arizona allenailenalienalienallen see above 1974 from where it was originally described leptophlebiidae and texas henry 1993 previous and new records from new mexico are confined to the choroterpes inornatainorincornatanata gila river drainage system eaton 1892 neochoroterpes nanita descriptions adult eaton 1892 larva traver 1934 kilgore and alienallenailen 1973 PREVIOUS RECORDRECORDSS kilgore and alienallenailen descriptions adult traver 1934 as 1973 catron co san francisco R colfax choroterpes nanita traver see also henry co cimarron R grant co Sasapillocapillopillo cr san 1993 larva alienallenailen 1974 as C nanita see miguel co windsor cr also henry 1993 NEW COUNTY RECORDS NMHU sandoval NEW RECORDS NMHU san miguel co co rio cebollabebollaCebolla PERC mora co coyote cr canadian R at sanchez 21111198821 iiiliiIH111 1988 LR REMARKS in addition to new mexico the smolka larvae southwestern species C inornatainorincornatanata is known from REMARKS this species was previously arizona and northern mexico eaton 1892 known only from texas traver 1934 in new including chihuahua ailenalienalienallen and murvosh mexico it has been taken in the canadian river 1987b colorado ward and berner 1980 and drainage system which continues east into more recently in texas lugoortizlugo ortiz and mccaf- texas ferty 1995b and utah mccafferty and mac- neochoroterpes oklahoma donald 1994 in new mexico C inornatainorincornatanata is 1934 now known from all major drainage systems traver except the san juan river drainage system descriptions adult traver 1934 as since it is found in the greater colorado river choroterpes oklahoma traver see also henry 302 GREAT BASIN naturalist volume 57

1993 larva alienallenailen 1974 as C mexicanusmexic anus NEW RECORDS NMHU mora co jack s alienallenailen see also henlyhenry 1993 cicr 10650 ft 18 VI 1991 M hatch larvae rio PREVIOUS RECORDS henry 1993 chaves arriba co san pedro park south entrance co rio penasco eddy co sitting bull falls 13 VI 1981 S carygary male and female adults NEW COUNIYCOUNTY RECORDS CSU colfax co REMARKS mcdunnough 1924a first de- bermejovermejovermejorVermejoRmejo R NMHU otero co dog canyon scribed paraleptophlebia heteroneaheteronea from alberta PERC mora co coyote cicr the species was misidentified from colorado REMARKS this species which has been as leptophlebia palkpespalpaipallipespalmipeslipes hagen now P memo known more commonly by the synonyms chor naltsnalis see below by dodds 1923 and was oterpes inexicanusmexicanusmexicannsanus alienallenailen or neochoroterpes similarly misidentified from utah somewhat mexicansmexicanameximeyicana ailenalienalienallen see mccafferty et al 1993 later by needham and christenson 1927 and henry 1993 is known from colorado however it was correctly reported from those parts of mexico including chihuahua okla- states by ward and bemerberner 1980 and edmunds homa and texas see henry 1993 mccafferty 1954 respectively our new records of this et al 1993 lugoortizlugo ortiz and mccafferty 1996a species in new mexico are based on both the in new mexico this species is now known more identifiable adults as well as larvae in fromfi om the eastern part of the state in both the new mexico it has been taken in the northern canadian and pecos rivelriver drainage systems part of the state in the canadian river and rio grande drainage systems paraleptophlebia debiksdebilisdabilis walker 1853 paraleptophlebia memmemonalismemomemorialismemorialistnalisorialis eaton 1884 descriptions adult walker 1853 as baetisbabtis debilisdebihsdabilis waikelwalkerwaiker see also key of halperharper and descriptions adult eaton 1884 as lep harper 1986 larva ide 1930 as leptophlebia tophlebiatophlebia memmemomemonalismemorialismemorialistnalisnaltsorialis eaton see also key of dedebihsdabilisdebilisbilis harper and harper 1986 larva seemann NEW RECORDS BYU eddy co castle cr 1927 as leptophlebia tufituftrufiruftvenosarufivenosadenosavenosa eaton 1884 black river village 141198714 1 1987 baumann sar- see also kilgore and alienallenailen 1973 gent and kondratieff larvae PERC grant PREVIOUS RECORDS peters and edmunds co N of silver city cherry cr 2422.44 mi N of 1961 as paraleptophlebia palpallipespalmipeslipes san juan cherry creek campgrounds 7 vilVII 1969 RW co san juan R kilgore and alienallenailen 1973 koss WP mccafferty and AXA V provonsha lar- catron co whitewater R lincoln co cr at vae sandoval co ca 32 km W of los alamos pine lodge santa fe national forest las conchas picnic NEW COUNTY RECORDS BYU sandoval co site E forkfolkmolkmork jemez R 1 IX 1969 R and D jemez R PERC colfax co cimarron R koss san miguel co I1 mi N of pecos pecos san miguel co pecos R rio arriba co rio R just below lisboa springs fish hatchery brazos taos co pueblo cr 31 VIII 1969 R and D koss larvae REMARKS this common western species REMARKS this transcontinental species is is known from all USA states adjacent to new known from colorado ward and stanford mexico except oklahoma and texas edmunds 1990 and utah edmunds 1954 but no other 1954 argyle and edmunds 1962 kilgore and states adjoining new mexico its presence in alienallenailen 1973 paraleptophlebia memmemomemorialismemonalismemorialistnaltsnalisorialis has new mexico therefore appears to represent often been recorded under its junior synonym the southernmostsoutheinmost range limits in western north leptophlebia palpallipespalmipeslipes hagen which proved to americaamenea in new mexico P debihsdebilisdabilis is presently be an unavailable name because it was a known flomfrom all major drainage systems with homonym of L palpallipespalmipeslipes walker 1875 para the exception of the san francisco and san leptophlebia memmemomemonalismemorialismemorialistnaltsnalisorialis has been taken in juan river drainagedi abageamage systems mexico only in baja california lugoortizlugo ortiz and mccafferty 1994 in new mexico it is paraleptophlebia heteroneaheteronea now known from all major drainage systems mcdunnough 1924 thraulodes brunbrunneusbrunneousneus descriptions adult mcdunnough 1924a koss 1966 as leptophlebia heteroneaheteronea mcdunnough see also key of harper and harper 1986 larva descriptions adult koss 1966 larva not published kilgore and alienallenailen 1973 199711997 NEW MEXICO MAYFLIES 303

PREVIOUS RECORDS koss 1966 grant co record cited above we have seen this species N silver city in catron county from the west fork of the NEW COUNTY RECORDS PERC catron co gila river and white water creek in new gila R mexico the species is known only from the REMARKS thraulodes brunbrunneusbrunneousneus is known gila river drainage system new from arizona and mexico and through- Traverella albertana out much of mexico including chihuahua mcdunnough 1931 alienallenailen and murvosh 1987b in the USA it has been taken only from the gila river drainage descriptions adult mcdunnough 1931a system as thraulusThraulus albertanusalbertanus mcdunnough see also traver and edmunds 1967 larva edmunds thraulodes gonzgonzalesigonzalesmalesi 1948 traver and edmunds 1967 PREVIOUS RECORDS kilgore and alienallenailen san description adult traver and edmunds 1973 as Traverella castanea catron co francisco R grant co gila R 1967 larva traver and edmunds 1967 NEW COUNTY RECORDS NMHU santa fe NEW RECORDS BYU grant co sapillocapillo co rio grande cr jet with gila R 1555 m 26 V 1985 B REMARKS Traverella castanea kilgore and jensen larva NMHU grant co E fork gila alienallenailen was shown to be a junior synonym of T 1950 X PE stew- R nr grapevine m 12 1993 albeitalbertanaana by lugoortizlugo ortiz and mccafferty 1995a art larvae PERC grant co sapillocapillo cr rd this western species therefore is known from 15 gila national forest 7 V 1981 WPWE mccaf- alberta and saskatchewan south to chihuahua ferty AXAV provonsha and D bloodgood larvae see alienallenailen 1973 alienallenailen and murvosh 1987b REMARKS thraulodes salinusgalinus kilgore and lugoortizlugo ortiz and mccafferty 1995a 1996a alienallenailen was recently shown by lugoortizlugo ortiz and although it is known from the surrounding mccafferty 1995a to be a junior synonym of areas of arizona chihuahua and colorado it T gonzgonzalesigonzalesmalesi therefore in addition to the new is not known from texas in new mexico it new mexico distribution the species is known has been taken only in the gila river upper from arizona kilgore and alienallenailen 1973 mexico rio grande and san francisco river drainage including chihuahua alienallenailen and brusca 1978 systems and texas traver and edmunds 1967 in new oligoneuriidaeoligoneurndae mexico it has been taken only in the gila river drainage system homoeoneunahomoeoneuria allem pescador and peters 1980 thraulodes speciosusspeciosus traver 1934 descriptions adult unknown larva pescador and peters 1980 descriptions adult traver 1934 larva PREVIOUS RECORDS pescador and peters mayo 1969 1980 dona ana co rio grande PREVIOUS RECORDS and durfee kondra REMARKS this species was originally de- tieff1995tieff 1995 as T arizonicusarizonicus catron co taylor scribed from new mexico by pescador and cr peters 1980 and those authors also reported NEW COUNTY RECORDS BYU grant co it from chihuahua and utah it has since been turkey cr found in colorado by durfee and kondratieff REMARKS thraulodes arizonicusarizonicus kilgore 1994 in new mexico it is known only from and alienallenailen was recently shown to be a junior the lower rio grande synonym of T speciosusspeciosus by lugoortizlugo ortiz and mccafferty 19951995aa this species previously lachlanialachlamaLachlanialamaiamaiania dencyannaedency annae has been known only from arizona traver koss 1970 1934 and northern mexico alienallenailen and murvosh descriptions adult koss and edmunds 1987b lugoortizlugo ortiz and mccafferty 1996a 1970 as L dencyannadencyanna larva koss and ed- including chihuahua alienallenailen and brusca 1978 munds 1970 as L dencyannadencyanna may have seen material from new mexico be- PREVIOUS RECORDS koss and edmunds cause they plotted the species in new mexico 1970 grant co gila R on a range map but presented no substantiat- REMARKS this is the only species of north ing data for it in addition to the new county american maymayfliesflies known exclusively from 304 GREAT BASIN naturalist volume 57 new mexico and therefore endemic to the state from mexico in adjacent chihuahua lugo- and the gila river drainage system it may ortiz and mccafferty 1994 and as far south as eventually be found inm the gila river drainage hidalgo lugoortizlugo ortiz and mccafferty 1996a system area of eastern arizona we consider in new mexico it has been taken in all major this species rare and endangered see species drainage systems except the canadian river of special concern below the ending of the drainage system specific epithet of this species has been emenderemended to conform to rules of nomenclature because it faunistics was based on a modern female name dency anne latinizedlatiniplatinizedzed by the original author and thus regional affinities should have further been given the appropri- we have categorized new mexico maymayfliesflies ate genitive ending by their general regional geographic affinities and present this data in table 1 western lachlanialachlamaLachlanialamaiamaiania saskatchewanensis in species are those that have a considerable north south ide 1941 distribution pattern in western north amer- descriptions adult ide 1941 see also icaleaiea only those new mexico maymayfliesflies found edmunds 1951 as L nowellipowelli edmunds larva north of colorado and utah and not in texas edmunds 1951 as L nowellipowelli unless restricted to west texas are consid- NEW RECORDS PERC san juan co san ered inm the western species category two sub juan R blanco state hwybwy 17 bridge 10 VIII categories of such western maymayfliesflies are infor- 1961 DWD W argyle larva san juan co animas mative with respect to faunistics these sub R hwybwy 17 bridge farmington 25 X 1961 categories segregate those species in which DWD W argyle larva new mexico is the southern limit of their REMARKS mccafferty 1996 recently ranges and those that occur further south showed S nowellipowellipowelh edmunds from utah ed- than new mexico and arizona southwestern munds 1951 and mexico lugoortizlugo ortiz and species are the and2nd mainmaln distributional cate- mccafferty 1994 to be a junior synonym oflofkoflL gory for new mexico maymayfliesflies these species saskatchewanensis thus L saskatchewanen occur no further north than colorado or utah sis occurs from saskatchewan to guerrero in and no further east than oklahoma or texas mexico it is known from the adjacent state of widespread species are the ard3rd main distribu- colorado mccafferty et al 1993 but is not tional category these are new mexico species reported from arizona chihuahua or texas that are western but also occur inm at least 1 new has taken only the in mexico it been in other large region of the continent iei e central san juan river drainage system northeastern or southeastern mexican species siphlonundaesiphlonuridae constitute a ath4th category new mexico species placed in this category are distributed mainly siphlonurus occidentoccidentalisoccidentalistalis eaton 1885 through mexico and occur no further north description adult eaton 1885 as than arizona new mexico or texas southern siphlurusstphlurus occidentoccidentalisoccidentahsoccidentalistalisails eaton larva clemens species constitute a ath5th category new mexico 1915 species placed in this category are distributed PREVIOUS RECORD traver 1935 santa in the southern USA further east than texas fe co santa fe R peters and edmunds 1961 and range no further north than new mexico san juan co san juan R durfee and kon inm the west eastern species constitute a ath6th dratieff1995dratieffdradragiefftieff 1995 catron co taylor cr category these species occur in the central NEW COUNTY RECORDS BYU grant co and northeastern USA in addition to new gila R turkey cr NMHU otero co agua mexico chiquita taos co rio hondo PERC grant table 1 shows that the largest component of co buck cr cherry cr little cr sapillocapillo cr the new mexico mayfly fauna is made up of san miguel co pecos R taos co rio grande species generally well represented in the west REMARKS mccafferty et al 1993 indi- most of these western species are restricted to cated that S occidentoccidentalisoccidentahsoccidentalistalisallsails was the most common the USA and canada with new mexico or species of siphlonurus in western north amer- new mexico and arizona representing their icaiealea it is known from all USA states adjoining southern range limit moreover many of these new mexico except texas and was reported latter species ege g drunella grandis ephorusepeorus 199719971 NEW MEXICO MAYFLIES 305

TABLE 1 major geographic affinities of the new mexico mayfly species see text for detailed definitions of the distributional categoriescategones western species southern limits at NM southern limits south of NM ameletus sparsparsatussatus ephemerellaEphetephemerellamerellamerelia infrequentinfrequens acentrella insignificantinsignificansinsigmficans pametropusametropusAmettopusropus albrighalbnghtialbrightiti heptageniaHeptagenia elegantulaelegantula babtisbaetis magnus tattenellatAtAttenella marmargaritaganta heptageniaHeptagenia solitanosolitanasolitariasolisolzsolitatiaariatanagriatanu caeniscanniscaems bajaensis tbaetisbaetisbabtis bicaudatus leucrocuta petersipelersipeterstpegerst callibaetis dictuspictus cmygmulat0nygmula par nixe criddlcriddleiei camelobaetidius varreniwarreni tdrunellatDrunellaruneliadrunella coloradensis nixe simplicioidessimplicioides lachlaniaLachlania saskatchewanensis tdrunellatDrunellaruneliadrunella doddsi 1 paraleptophlebiatparaleptophlebia heteroneaheteronea paraleptophlebiamemorialisparaleptophlebia memomemonalisnaltsnalis tDtdrunellarunelladrunella grandisgi andis pi ocloeonprocloeon conturbatumconturba tum rhithrogena mossonimomsonimorrisonimotmorrisoni tEpetepeorusephorusEpeepeorusofusorus albertae i rhithrogenatrhithrogena robusto serratella michenerimicheneri tEpetepeorusephorusepeorusorus deceptiousdeceptivus tserratellaserratella tibialis siphlonurussiphlorturus occidentoccidentalisoccidentalistalis tEpetepeorusephorusepeorusorus longimanuslongilongimanousmanus ttimpanogatimpanogaTimpanoga hecuba Traverella albertanaallger tana ephemerellaEphe merella inermisinermia southwestern species ameletus doddsianusdoddsianus ephorusepeorus margaritamarmayganta neochoroterpes oklahoma ameletusfalsusameletus falsusbalsus ephemerellaEphemerella altana rhithrogena hahhagemhaggeni baetisbabtis adonis ephemerellamollitiaephemerellaEphemetellamerella molmoimollitiaittialitia rhithrogena plana babtisbaetis caecoelestiscaelestislestis homoeoneuriahomoeoneuna allenialbeni r rhithrogena vitta babtisbaetis notesnotos IsJsonychia interintennediaintermediaintermedialmedia thraulodes gonzalesigonzgonzalesmalesi baetodesBaetodes edmundedmundsiedmundsosi f labiobaetis apache thcorythodestncorythodes condolusconcondylusdylus camelobaetidius musseri lachlaniaLachlania dencyannaedency annae tricorythodestncorythodes corpulentcorpulentusus choroterpes inorinornataincornatanata neochoroterpes bossikossi tncorythodestricorythodes dimorphusdimdimorphousorphus t0oeodesciocloeodesCloeodes macrolamellus neochoroterpes nanita tricorythodestncorythodes explicatesexplicatus widespread species acentrella turbidasurbida callibaetisfluctuanscallibaetis fluctuantfluctuans Isonychia sicca baetisflavistrigababtisbaetis flaviflavistrigastriga ephemera simsimulantsimulansulans paraleptophlebia debilisdehilisdabilis babtisbaetis tncaudatustricaudatus fallceonFallceon quillenquilqulileriihrl rhithrogena undulataundungulataulata caeniscanniscaems latipennislatipenmsiatilatipennis hethexageniahexagemahexagyniaHexageniaagema limbatalimbaga tncorythodestricorythodesminutusminutesminutus callibaefisferrugineuscallibaetis ferruginous mexican species baetodesBae todes deficiensdefiniens thraulodes brunbrunneusbrunneousneus thraulodes speciosusspeciosus callibaetis montanus southern species leptophlebia bradleyi eastern species labiobaetis propinquuspropinquous

1 speciesispccies restricted to mountmountainun streamsitieitic ims I1 foundpound only inin thetlletile greatergre ttcitaci coloradocoloichloi idoudo riverkiver drainagedi midge system longilongimanuslongimanousmanus are restricted to mountain streams restricted to the greater colorado river drain- and some have their southern range limit in age system the rocky mountain province of northern thirteen species in new mexico are con- new mexico eg serratella tibialis none of sidered widespread some of these are rela- the western species whose ranges extend into tively ubiquitous ege g babtisbaetis tncaudatustricaudatus cal mexico are restricted exclusively to mountain libietishbaetislibaetislibaetis ferrugiferruginousneus tncorythodestricorythodes minutesminutusminnmmutusminufustus streams ephemera simsimulantsimulansulans has its southwestern range the more geographically restricted south- limit represented in new mexico on the other western species in new mexico make up almost hand caeniscannis latipennislatipennis and fallceonFallceon quilleriquilquillerzlerilerz as large a component as the western species in are widely distributed in much of the USA but new mexico none of these species are re- also occur through mexico and central amer- strictstricteded to mountain streams except cloeodesCloeodes leaicaiea acentrella turbidasurbida is widespread but gener- macrolamellus but that species is not found at ally found in cool well oxygenated water and high altitudes the southwestern species lab thus in new mexico is found in mountain iobaetisiobaetis apache and lachlaniaLachlania dencyannaedency annae are streams 306 GREAT BASIN naturalist volume 57

four species represent an essentially mexican be made with a comparative index that can fauna in new mexico of these all but takes into account the degree of similarity and callibaetismontanuscallzbaetis montanus aiealeare restricted in distribu- difference in the faunalfaunas of any 2 regions we tion in new mexico to the gila river drainage have used Sorensorensenserssetss coefficient of similarity system although C montanus has invaded SC to compare new mexico more thor- other drainage systems in the state besides the oughly with each of its broadly adjoining gila it is restricted to lower lying areas san states SC 2a2a b c where a num- juan river and southernmost area of the rio ber of species in common between new mex- grande drainage system baetodesBaetodes definiensdeficiens is icoleoieo and a comparative state b the number notable because its presence in sapillocapillo creek of species in new mexico and c the num- gila river drainage system is its only known ber of species in the comparative state the occurrence in the USA results of such calculations taking into consid- two species show atypical distribution pat- eration any recent taxonomic revisions such as terns by their presence in new mexico lep zloty 1996 are as follows for new mexico tophlebiatophlebmtophlebialehla bradleyi is a southern species that and arizona there are 52 species in common reaches its westernmost limits in extreme south- and SC 04100.4141 for new mexico and chihua- eastern new mexico labiobaetis propinquuspropinquous hua 18 species in common and SC 0350.350 35 for is a mainly central USA species that has its new mexico and colorado 48 species in com- southwestern range limit in northeastern and mon and SC 0350.350 35 and for new mexico and south central new mexico we regard its dis- texas 22 species in common and SC 0200.20090ogo0 20 tributiontribution in new mexico as anomalous since it to a large extent arizona mirrors the aquatic does not fit any predictable pattern however environment of new mexico the shared may- the species may prove to be more widespread fly fauna of the 2 states is mainly southwestern in north america than now thought and there- but also contains many of the western moun- fore fall into the widespread category tain species table 1 this accounts for the we have documented the occurrence of a high coefficient of similarity for these 2 states total of 12 families 37 genera and 81 species of eighteen of the 27 species thus far known maymayfliesflies in the state of new mexico by com- from chihuahua occur in new mexico and parison with broadly adjoining states 9 families this is the mainmaln reason for considerable simi-simisiml 28 genera and 66 species have been recorded larity between these states with respect to re- for arizonaanzona see lugoortizlugo ortiz and mccafferty lationshipslationships with colorado just as with arizona 1995a zloty 1996 5 families 14 genera and over half of the new mexico species are found 27 species have been recorded for chihuahua in colorado also however less than half of see lugoortizlugo ortiz and mccafferty 1995d 1996a the colorado species are found in new mexico mccafferty and lugoortizlugo ortiz 1996 14 families this is because of a considerable number of 44 genera and 100 species for colorado see more boreal transcontinental species that occur mccafferty et al 1993 durfee and kondra- in colorado mccafferty et al 1993 but not tieff 1994 and 11 families 34 genera and 92 new mexico the large number of shared species roifoifotforror texas see lugoortizlugo ortiz and mccaf- mountain species all those mountain restricted ferty 1995b the low number of taxa for the species shown under the western southern mexican state of chihuahua is due to the more limits new mexico categories of table 1 is restricted availability of freshwater habitats in offset by the fact that many of the southwest- that primarily desert area and also the fact that ern species in new mexico do not reach col- collecting efforts there have not been on a par orado thus giving a basically intermediate with those inm the other states mentioned on coefficient of similarity between the 2 states the othelother handband the relatively high mayfly taxa the low coefficient of similarity between new number in colorado is due to the diverse mexico and texas is explained by the fact that topography and array of freshwater habitats only one of the basically eastern and south- available in that state as well as the fact that easternn species of east and central texas see colorado has been one of the better worked lugoortizlugo ortiz and mccafferty 1995b reaches new north american regions with respect to sam- mexico and in the same respect western moun- pling of aquatic macromaeromacromvertebratesmacroinvertebratesinvertebrates tain species of new mexico do not reach texas A more informative comparison of the new the vast majority of the 22 species shared by mexico mayfly fauna with that of other regions new mexico and texas are southwestern or 199719971 NEW MEXICO MAYFLIES 307 mexican species but some are widespread fig 2 by summing all intra drainage coeffi- acentrella insignificantinsigmficansinsignificans represents a rare case cients for each of the drainage systems we can of a species distributed mainly in the north- see that the rio grande system fauna is most west but which has also penetrated west texas representative of the entire new mexico fauna evidently via the canadian and pecos river whereas the san juan and canadian system drainage systems faunalfaunas are least representative of the state eighteen new mexico mayfly are drainage system relationships species known exclusively from western drainages in in table 2 we have sorted the new mexico the state san juan zunizum san francisco gila mayfly species by major drainage system and 23 species are known exclusively from species richness is greatest in the gila system eastern drainages essentially all drainages east 49 species followed closely by the rio grande of the continental divide dry cimarron cana- 46 species then pecos 39 species canadian dian rio grande pecos see table 2 fig 1 29 species and finally san juan 25 species the genera to which these exclusive drainage the gila although not large compared with system species belong can be considered with some other diadladrainagesmages does have a unique com- respect to their recent biogeographic origins plex of both western mountain species and edmunds et al 1976 mccafferty et al 1992 species that could be considered lowland twelve of the 18 species confined to the west- species many of which are southwestern or ern drainage systems are species that are mem- mexican the species richness of the rio bers of genera that have recent neotropical grande and pecos systems is in laigelarge part due biogeographic origins baetodesBaetodes camelobaecarnelobae to the western mountain species present inm fidiustidius leptohyphes tncorythodestricorythodes thraulodes headwaters and mountain tributatributariesries and the LachlochlanLachlanlachlanialachlanidlaniaianiaid on the other hand nearly all species range of habitats represented over their rela- 22 of 23 that aie found exclusively in eastern tively long courses including large low lying drainages in new mexico belong to genera rivers the canadian and san juan systems that have boreal nearctic biogeographic oriorlori- are somewhat restricted both in area and in gins Ameletus baetisbabtis labiobaetis caeniscanniscaems habitats both primarily with low lying courses tenellaattinellaattenellaAt drunella ephemerellaEphemerella serratella with considerable silting particularly the san timpanogaTimpatzmpanoganoga ephemera hexageniahexagemahexagyniaHex ageniaagema cinygmula juan A number of species collected from the ephorusEpeepeorusorus rhithrogenarhtthrogena Isonychia neochoro san juan river prior to 1961 may no longer be terpescerpes leptophlebia paraleptophlebia homolohomoeo present there see species of special concern neunaneuriaurla this comparison suggests that the below greater colorado drainage system has been a if intrastate comparisons of the drainage major northern dispersal corridor for maymayfliesflies system mayfly faunalfaunas are made by calculating during its 353 5 million year existence it also sorensenisorenseiisSorensorensenssens coefficients see formulation above suggests that the greatelgreater rio grande drainage it becomes apparent that the greatest similar- system has not been amenable to northern dis- ity of any 2 systems is that shown by the gila persal of austral taxa but has been primarily and rio crandeglandegrande systems fig 2 which also colonized fromblombrom the north however mountain happen to be the richest systems the pecos species found exclusively in the eastern drain- system is also relatively similar to both the age systems there are 9 of these evidently gila and rio grande fig 2 the least simisimlsimi- have been limited to the northern part of the larity is shown between the gila and canadian state by topography the 38 species of new systems probably due to the distance between mexico maymayfliesflies that occur in both eastern and them compounded by the fact that one is a western drainage systems in the state belong western drainage and the other an eastern to a mix of nearctic and neotropical genera drainage and by their overall dissimilarity in and demonstrate that none of the drainage habitat availability the canadian system has systems in new mexico should be considered only 14 species in common with the san juan one way coineoineolncorridorsdors system and another 14 in common with the in species of special concern gila system factoring in the faunal size of the systems in question however shows that the recent studies of the maymayfliesflies of the south- canadian and san juan systems have a greater west indicate that only I1 endemic species occurs similarity than the canadian and gila systems in new mexico it is lachlanialachlamaLachlanialamaiamaiania dencyannaedency annae a 308 GREAT BASIN naturalist volume 57

TABLETAB 1 2 drainagediadlaDidindge system distributions of new mexico mayfly species san juan including zuni acentrella insignificantinsigmficansinsignificans ephemerellaEphemerella mollitiamollitia paraleptophlebia memmemomemorialismemonalismemorialistnaltsnalisorialis ametropusametiAmet topusopusropus albrighalhnghtialbrightiti heptageniaheptagemaHeptageniaagema elegantulaelegantula procloeon conturbatumconturbatum baetisflavistrigabaetisbabtis flavistnga heptageniaheptagemaneptHeptageniaagema solitanasolitanosolitariasolisolitariaafiaaflatana rhithrogena morrisonimonsommormomsommoyrisoni babtisbaetis tricaudatustncaudatus labiobaetis apache rhithrogena undundulataungulataulata callibaetisferrugineuscallibaetis ferruginous lachlaniaLachlania saskatchewanensis serratella chenenmimichenermichenenmichenerimicheneri callibaetiscallihaetisCallihaetis montanus leptohyphes apache siphlonurus occidentoccidentalisoccidentalistalis callibaetis dictuspictus nixe criddlcnddleicriddleiei tncorythodestricorythodes explicatesexplicatus camelobaetidius varreniwarreni nixe simplicioidessimplicioides tncorythodestricorythodes minutesminutus ephemerellaepheinerellaEphemerella inermisinerinisinermiaincinerinisninis gila including san francisco acentrella insignifteansinsigmficans drunella doddsi paraleptophlebia dabilisdebilis Aineaineletusameletusletuslefus doddsianusdodds ianus ephorus deceptiousdeceptivus paraleptophlebia memdemoimemoimemorialismemorialistorialistaltslalistaits baetisbabtis bicaudatushicaudatus ephorusepeorus longimanuslongilongimanousmanus rhithrogena plana baetisbabtis caecoelestiscaelestislestis ephorusepeorus marmargaritamurganta rhithrogena lobustarobusta babtisbaetis magnus drunella grandis rhithrogena undulataundungulataulata babtisbaetis notos ephemerellaEphemerella altana rhithrogena vitta baetisbabtis tricaudatustriuiudatus ephemerellaEphemerella inermisinermia serratella mimichcheneneneri Baebaetodesbtietodestodes definiensdeficiens fallceonFallceon quilleriquilleri siphlonurus occidentoccidentalisoccidentalistalis baetodesBae todes edmundedmundsiedinundsiedmundsosi heptageniaheptagemaHeptageniaagema solitanasolitariasolitanosolisolosolitariaarlatana thraulodes brunbrunneusbrunneousneus canniscaeniscaems bajaensisimjaensis Isonychia interintermediaintermedialmedia thraulodes gonzgonzalesigonzalesmalesi callibaetifcallibaetisferrugineusfenpen ugineuh lachlaniaLachlania dencyannaedency annae thraulodes speciosusspeciosus callibaetiscullibaetis inonmontanustanus leptohyphes apache Traverella albertana callibaetiscalhbaetis dictuspictus leucrocuta pelersipetersi tncorythodestricorythodes condolusconcondylusdylus cennelobaetidiuscamelobaetidius musseri neochoroterpesneochoi oterpes bossikossi tricorythodestncorythodes corpulentcorpulentusus camelobaetidiuscamelolwetidius warrent nixe criddlcnddleicriddleiei tncorythodestricorythodes dimdimorphusdimorphousorphus choroterpes inorinornataincornatanata nixe simplicioidessimplicioides tncorythodestricorythodes minutesminutus cloeodesCloeodes macrolamellusmacrolamellw canadian including dry cimarron acentrella infimficansinsignficans drunella grandisgt andis nixe simplicioidessimplicioides baetisbabtis bicaudatus ephemerellaEphemerella altana paraleptophlebia dabilisdebilis baetisbabtis manusinagnus ephemerellaEphemerella inermisinermia paraleptophlebia heteroneaheteronea baetisbabtis notos heptageniaheptagemaHeptageniaagema elegantulaelegantula paraleptophlebia memmemomemorialismemonalismemorialistnalisnelisorialis baetisbabtis tncaudatustricaudatus heptageniaheptagemaHept ageniaagema solitanasolitariasolitanosolisoltsoitsolitariaarlatana rhithrogena morriconimorrisommorrisonimormoyrisonirisom baetodesBaetodes edmundedinundsiedmundsiedmundsosi labiobaetis propinquuspropinquouspropmquus rhithrogena undulataundungulataulata caeniscannis bqjaensisiwjaensis leucrocuta pelersipetersi serratella mimichcheneneneri camelobaetidius warrent neochoroterpes nanita tricorythodestncorythodes explicatesexplicatus choroterpes inorinomatanata neochoroterpes oklahoma tricorythodestncorythodes minutesminutus drunella doddsi nixe criddlcnddleiei

notably distinct and unique mayfly with a age system can therefore be viewed as a re restrictive habitat koss and edmunds 1970 fugiumfulgium in new mexico for certain southwest- most other species once thought to be ern species in addition lachlaniaLachlania is a highly endemic to new mexico have eventually been unusual group of maymayfliesflies with rapid flight found in arizona often in the same drainage habits and associated wing morphology unique system middle or lower colorado river drain- among the ephemeroptera see edmunds et age system lachlanialachlamaLachlanialamaiamaiania dencyannaedency annae deserves al 1976 and a boltingmolting process unique among special attention for other reasons beside its all insects see edmunds and mccafferty 1988 being endemic it is limited in distribution to given the highly restricted range and remark- the gila river drainage system camelobaetid able nature of this mayfly species we recom- tuslusiuszus musserirnussen cloeodesCloeodes macrolamellus rhithro mend that L dencyannaedency annae be given special con- gena vitta tncorythodestricorythodes condolusconcondylusdylus T corpu sideration as a rare and endangered species lentusclentus neochoroterpes bossikossi thraulodes brun both at the state and federal levels because of neus T gonzgonzalesigonzalesmalesi and T speciospeciosusspeczosussus are addi- the degradation of the cilacliagilaglia river especially in tional new mexico species that are found only arizona see ege g mcnamee 1994 and impend- in the gila river drainage system this draindram ing degradation in new mexico we suggest 199711997 NEW MEXICO MAYFLIES 309

TABLE 2 continued rio grande acentrella insignificantinsigmficansinsignificans ephorusepeorus deceptiousdeceptivus paraleptophlebia heteroneaheteronea acentrella turbidasurbida ephorusepeorus longilongimanuslongimanousmanus paraleptophlebia memmcmmcmomemomemonalismemorialismemorialistnaltsnalisorialis ameletus doddsianusdodds ianus ephorus matmarmargaritaganta rhithrogena hagenihagemhagenl ameletusfalsusameletus falsusbalsus ephemera simsimulantsimulansulans rhithrogena mormornsonimorrisonimorrisonnmornrisonisonisont Attenella dargaimargaimargaritaita Epheephemerellamerella altana rhithrogena plana babtisbaetis bicaudatus ephemerellaEphemerella inennisinermisinermia rhithrogena lobustarobusta babtisbaetis magnus ephemerellaEphemerella infrequentinfrequens rhithrogena undulataundungulataulata baetisbabtis tncaudatustricaudatus fallceonFallceon quillenquilqulilerilefi serratella chenenmimichenermichenenmichenerimicheneri callibaetiscallibaetisferrugineusferruginous heptageniaheptagemaHeptageniaagema solitanasolitanosolitariasolisoltsoitsolitariaafiaaflatana serratella tibialis callibaetiscalhbaetis montanus homoeoneunahomoeoneuria allem siphlonurus occidentoccioccidentalsoccidentaleoccidentalisdentalsalisallsails callibaetis dictuspictus Isonychia interintermediaintermedialmedia timpanogaTimpanoga hecuba choroterpes inorinornataincornatanata Isonychia sicca Traverella albertana drunella coloradensis nixe criddlcriddleiei tricorythodestncorythodes dimorphusdimdimorphousorphus drunella doddsi nixe simplicisimplicioidesoides tricorythodestncorythodes explicatesexplicatus drunella grandis paraleptophlebia dabilisdebilis tricorythodestncorythodes minutesminutus ephorusepeorus albertae pecos acentrella insignificansinsigmficansinsignificant callibaetis dictuspictus leptophlebia bradleyi acentrella turbidasurbida choroterpes inorinornataincornatanata neochoroterpes oklaokiaoklahomahoina ameletusfalsusameletus falsusbalsus cinygmula par nixe criddlcriddleiei ameletus sparsparsatussatus drunella coloradensis nixe simplicisimplicioidesoides Attenella margaritamarganta drunella doddsi paraleptophlebia dabilisdebilis babtisbaetis adonisadorns drunella grandis faraleptophlebiaparaleptophlebia memmemomemorialismemonalismemorialistnalisnaltsorialis babtisbaetis bicaudatus ephorusepeorus deceptiousdeceptivus rhithrogena mormornsonimorrisonimorrisonnmornmonnrisonisoni babtisbaetis caecoelestiscaelestislestis ephorus longimanuslongilongimanousmanus rhithrogena undundulataungulataulata babtisbaetis magnus ephemerellaEphemerella altana serratella chenenmichenerimichmi eneri babtisbaetis tricaudatustncaudatus ephemerellaEphemerella inermisinermia siphlonurus occidentoccidentalisoccidentalistalis caeniscanniscaems latipennislatipenmslatiiatipennis fallceonFallceon quillenquilqulileri timpanogaTimpanoga hecuba callibaetiscallibaetisferrugineusferruginous hexagemahexageniahexagyniaHexageniaagema bilbtlblibilineatabtlineatabilineateineata tncorythodestricorythodes explicatesexplicatus callibaetisfluctuanscallibaetis fluctuantfluctuans labiobaetis propinquuspropinquous tncorythodestricorythodes minutesminutus

that the other gila river drainage system mayfly county lower pecos river drainage system it species mentioned above minimally be con- may have only recently invaded the state and sidered at risk we expect that it may become more widespread we consider the following mayfly species if it is relatively vagile as is typical of other rare or in some cases possibly recently extir- species in the genus mccafferty et al 1992 pated in new mexico Ametametropusropus albrighalbrightiti is a procloeon conturbatumconturbatum is known in new mex- sand dwelling mayfly with very specific habitat ico only from adult males taken at the san requirements see clifford and barton 1979 juan river leptophlebia bradleyi is known only that has been taken only in the san juan river from a single sample from eddy county which in new mexico and the green river system in apparently is the westernmost record of this the region where colorado wyoming and utah southern USA species neochoroterpes nanita is meet allenailenalienalienallen and edmunds 1976 although known from only I1 sample from the canadian such sand dwelling species can be difficult to river in san miguel county otherwise it is sample mccafferty 1991 A albrighalbrightiti has not known only from central and east central been taken in new mexico since 1935 sub- texas henry 1993 lugoortizlugo ortiz and mccaf- lette et al 1990 discussed the changes to the ferty 1995b rhithrogena hagenihagemhagenl is known san juan river related to a recent host of large only from an isolated fringe population in the scale irrigation facilities in the san juan basin red river inm taos county upper rio glandegrande and construction of the navajo dam calli drainage system although also rare in arizona baetisfluctuansbaetisbabtis fluctuantfluctuans is a common species elsewhere it is relatively common in colorado but in new mexico it has been taken only from hexageniahexagemahexagyniaHexageniaagema bilbihbibbilineatabihneatabilineateneataineata is a large common and a single sample from castle creek in eddy well known species in eastern and central areas 310 GREAT BASIN naturalist volume 57

G C R P singular new mexico record was questioned by alienallenailen and edmunds 1965 the fact that the species is also known in california from S 15.2911529 13.331 1427 152915.2915291 133313.3313331 1631116131161.3116.311 142714.271427114.271 the san gabriel mountains as is B adonisadorns above would possibly add credence to the new mexico report if it was taken from new G 13125131.25 3013930391301.3930.391 223622.362236122.361 mexico it may now be extirpated there due to alteration of habitats in the san juan river C 173117.311731117.311 153115.311531115.311 that have occurred inm the last half of this century it is possible that leucrocuta pelersipetersi presently occurs only in new mexico outside of R 253725.37 new mexico the species has been known only from the green river of northeastern utah fisfigpig 2 matrix of new mexico species shared by and sorensensolSoi ensen s coefficients of similarity in brackets between and southwestern wyoming however the majolmajor drainagediadlainage sysystemssterns in new mexico S san juan species may now be extirpated in these latter zuni G gila san flanrianfranciscoFian elseocisco C canadian dry states because the construction of flaming R glandegrande pecos cimarron rio clandecrande P gorge dam has had some detrimental effects on maymayfliesflies of that area edmunds personal of north america in new mexico however communication and L pelersipetersi has not been its distribution is disjunct the population is found there subsequent to the collections of unusual see species accounts above and the the type material 1945 1962 alienallenailen 1966 the single habitat where it is found is a one of a species apparently is common in the gila river drainage new also kind area in the state it consists of a 1 km system in mexico and it has long 2 m deep pool with fine calcareous sed- been taken in a sample from the canadian iments it is located just upstream from harkey river drainage system in another part of the crossing on the black river eddy county state we doubt there is another habitat with this type of burrowing substrate required by hexa acknowledgments1 S genia maymayfliesflies in new mexico the road cross- provo ing mentioned above acts as a partial dam we thank dick baumann utah across the limestone basin and any alteration george edmunds salt lake city utah and of the crossing or roaroadworkdwork would seriously boris kondratieff fort collins colorado for affect the depositional nature and sediment the loan and donation of collections of new trapping feature of the extensive pool mexico maymayfliesflies we also thank arwin provon- homoeoneuria allenialbeni is known somewhat sha west lafayette indiana for the artwork north and south of new mexico in new mex- to all those who have collected maymayfliesflies in new ico it has been taken only once from the lower mexico beginning in the 1960s and whose rio grande in dona ana county however names appear in the new records sections of we expect that this sand dwelling species is this paper we offer our sincere gratitude for difficult to collect in larger rivers likewise making this study possible we also thank SLS L lachlaniaLachlania saskatchewanensis known only from joseph and ARA R alexander who collected new the san juan river drainage system in new county records but whose names did not appear mexico is also difficult to sample and in reality in the text the research was supported in part may not be rare by a grant from the american philosophical baetisbabtis adonis ephemerellaEphemerella molmoimollitiatnollitialitia and leu society additional financial support for some crocuta pelersipetersi are apparently rare north amer- collections since 1991 was provided in part by ican species baetisbabtis adonis is known only from grant F 59 R through the federal aid in sport the san gabriel mountains of southern cali- fish restoration program administered by the fornia and the upper pecos river in the foot- new mexico department of game and fish and hills of the sangre de cristo mountains in san the USU S fish and wildlife service for develop- miguel county new mexico ephemerellaEphemerella ment of an index of biotic integrity IBI for molmoitnollitiamollitialitia is also known only from california and surface waters of new mexico the study has new mexico although it has not been reported been assigned purdue agricultural research from the san juan river since 1935 and this program journal number 15347 199711997 NEW MEXICO MAYFLIES 311

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maymayfliesflies 01or ephemeropteiaephemeroptera of illi- 449 456 noisnols illinois natural history survey bulletin 26 ALLEN RKR K AND SDS D COHEN 1977 Maymayfliesflies ephemerop- 1 216 tera of mexico and central america new species CLEMENS WA 1913 new species and new life historieshistones descriptions and records canadian entomologist of ephemeridae or maymayfliesflies canadian entomologist 109399109log 399 414 4524645 246 262262329329 341 ALLEN RKR K AND GEG F EDMUNDS JRTR 1959 A revision of 1915 Maymayfliesflies of the siphlonurus group cana- the genusgenns ephemerellaEphe merella ephemeroptera echemeepheme dian entomologist 4724547 245 260 rellidaerallidaerellidae I1 the subsubgenusgenusgenns timpanogaTimpanoga canadian CLIFFORDCEIFFORD HFH F AND DRD R BARTON 1979 observations on entomologist 915191 51 58 the biology of ametropusAmetropus neavelneavei ephemeropteraa 1961 A revision of the genus ephemerellaEphemerella ephem- ametropodidae from a large aveinveiriverfiver in noinolnorthernthern alberta eroptera ephemerellidae liiiliIII111 the subsubgenusgemis atten canada canadian entomologist ililii111855111 855 858 uatellacatella journal of the kansas entomological society COHEN SDS D AND RKR K ALLEN 1972 new species ofbaeokbaeof bae 3416134 igi161 173 todes fromflom mexico and centralgentral america pan pacific 1962 A revision of the genus ephemerellaEphemerella ephem- entomologist 4812348 123 135 eroptera ephemerellidae V the subgenus dru- DAY WC 1963 ephemeroptera pages 79 105 in RLR L nella in north america miscellaneous publications usinger editor aquatic insects of california uni- of the entomological society ofamerica 31473 147 179 versity of california press berkeleybelBei keley 1963 A revision of the genus ephemerellaEphemerella ephemepbemepremephem- DODDS GSG S 1923 Maymayfliesflies from colorado descriptions eroptera ephemerellidae VI the subgenus ser- of certain species and notes on otheisothersnotheis transactions ratella in north america annals of the entomologi- of the american entomological society 699369 93 116 cal society of americaofamerica 5658356 583 600 DURFEE R AND BCB C kondratieff 1993 description of 1965 A revision of the genus ephenwrellaephemerellaEpheEphenwrellamerella ephem- adults of baetisbabtis magnus ephemeroptera baetidaeBaetidae eroptera ephemerellidae VIII the subgenus entomological news 104227104 227 232 ephemerellaEphemerella in northn01 th america miscellaneous pub- 1994 new additions to the inventory of coloradocoloichloi ado licationslications of the entomological society of americaamenea maymayfliesflies ephemeroptera entomological news 105 42434 243 282 222 227 1976 A revision of the genus Ametametropusropus in noinorthth 1995 description of adults ofbaetisbabtisbaetisnotosnotos ephem america ephemeroptera ametropodidae journal eiopteiacropteracroptera Baetbaetidaebaetidacidae entomological news 10671106log 71 74 of the kansas entomological society 4962549 625 635 1997 description of adults of labiobaetis apache ALLEN RKR K AND CMC M MURVOSH 1983 taxonomy and ephemeroptera Baetbaetidaebactidaeidae with additions and col-cor- zoogeography of the maymayfliesflies ephemeroptera lectionsrectionsreetrectionslons to the inventory of Colorado maymayfliesflies ento- insecta ofbajaof baja califoiniacalifornia annals of the entomolog- momologicallogical news 10897losios108 97 101 ical society of america 7642576 425 433 earonEATON AE 1871 A monogiaphmonograph of the ephemeridae 1987a new Baetbaetidaeidae from the southwestern transactions of the entomological society of london united states and northern mexico insecta ephem 19119igi 1 164 312 GREAT BASIN naturalist volume 57

1883 88 A revisional monograph of lecentrecent ephem- 1941 Maymayfliesflies of two tropical genera lachlanialachlamaLachlanialamaiamaiania eridae 01or maymayfliesflies transactions of the linnaean and Campcampsurussurus from canada with descriptions cana- society of london zoological series 313 1 352 dian entomologist 7315373 153 156 1892 Biologia centialicentralicentioli americana insecta neur- JOHNSON SCS 1978 larvae of ephemerellaEphemerella inermismerimsmerrms and opteraa ephemeridae bernardbeibel nainal d quaritchquaritschQuaquaiqual itchritch london E infrequentinfrequens ephemeroptera ephemerellidae pan EDMUNDS GFG F JR 1948 A new genus of maymayfliesflies from pacific entomologist 541954 19 25 westelwestern n northnor th america leptophlebiinaeleptophlebnnae proceed- KELTNER J AND WPWE mccafferty 1986 functional ings of the biological society of washington 61 morphology of burrowing in the maymayfliesflies hexageniahexagemahexagyniaHex ageniaagema 141 148 limbatalimbaga and pentageniarentpentagyniabentPentpentagemaageniaagemaogema vitvittigeratigera zoological journal 1950 new records of the mayfly genus baetodesBaetodes of the linnaean society 6713967 139 162 with notes on the genus entomological news 61 KILGORE JIJ I1 AND RKR ALLEN 1973 Maymayfliesflies of the 203 205 southwest new species descriptions and records 1951 new species of utah maymayfliesflies I1 aligooligo ephemeroptera annals of the entomological soci- neumdacnetiriidae ephemeroptera proceedings of the ety ofamericaof america 6632166 321 332 entomological society of washington 5332753 327 331 kondratieffkondratiefe BC AND JRJ R VOSIIELLVOSHELL JR 1984 the north pi 1954 the maymayfliesflies of utah oceoceedingsproceedingsedings of the and central american species of Isonychia ephemer- utah academy of sciences aitsaltsarts and letters 316431 64 66 optera oligoneurndaeoligoneuriidae transactions of the ameri- EDMUNDS GF JR ANDRKAND RK ALLEN 1964 the rocky can entomological society iloiio110129110 129 244 epeorus mountain species of ephorus iron eaton ephe- KOSS RWR W 1966 A new species of thraulodes from new meropmeropteratera heptageniidaeheptagenndae journal of the kansas mexico ephemeroptera leptophlebndaeleptophlebiidae michi- entomological 37 275 288 society 37275 gan entomologist igi1911 91 94 EDMUNDS GFGE JR AND WPWE mccafferty 1988 the 1972 baetodesBaetodes new species and new records for mayfly subimago annual review of entomology 33 north america ephemeroptera baetidaeBaet idae entomol- 509 529 ogical news 839383 93 102 EDMUNDS GEG F JR AND GGG G MUSSERMUSSLR 1960 mayfly the koss RWR W AND GEG F EDMUNDS JR 1970 A new species fauna of green the Fl gorge gi een rivelriver inm flamingarning basin of lachlanialachlamaLachlanialamaiamaiania from new mexico with notes on the anthropol- wyoming and utah university of utah genus proceedings of the entomological society of ogical papers 4811148 111 123 ililii washington 725572 55 65 EDMUNDS GFG F JR SLS L JENSENJLNSLN AND L BERNER 1976 LEONARD JW AND FAEA LEONARD 1962 Maymayfliesflies of rhe maymayfliesflies of north and central america univer- the michigan trout streams cranbrook institute of sci- sity of minnesota presspi ess minneapolis ence bloomfield hills MI eislniioodwW 1979 precipitation pagepagelomjl10 in JL williams eislniiood LOWEN RGR G AND J F FLANNAGAN 1991 four manitoba and PE mcallisterMcAllistelisterlster editorsediedltoistols new mexico in maps JE in of centroptilumofcentroptilum eaton ephemeroptera bae technology application center university of new species tidae with remarks the genus pages 189 205 mexico albuquerque otidae on in J alba tercedor and A sanchez ortega editors FLOWERS RW 1987 adult stage of three central the overview and of ephemeroptera and ple- american baetodesBaetodes ephemeroptera baetidaeBaetidae with strategies coptera sandhill crane press gainesville FL notes on the genus aquatic insects 9 1 10 LUGOORTIZLUGO ORTIZ CRR AND WPWE mccafferty 1993 genera FLOWERSFLOWLRS RW AND WL hilsenhoff 1975 heptageniheptagem idae ephemeroptera of wisconsin great lakes of Baetbaetidaebactidaeidae ephemeroptera from mexico entomol- news 197 entomologist 82018 201 218 ogical 104193104 193 1994 new records of ephemeroptera from mex- GRAY LJL J 1981 species composition and life histories of ieoicoleo entomological news 105 17 26 aquatic insects in a lowland sonoran desert stream ico 10517losios american midland naturalist 106229 242 1995a annotated inventory ofthe maymayfliesflies ephem- news 106 harperHARPLRHABPER F AND PP HARPERHARPLR 1986 an annotated key to eroptera of arizona entomological the adult males of the northwestern nearcticNeaictic species 131 140 of paraleptophlebia lestage ephemeroptera lep 1995b the maymayfliesflies ephemeroptera of texas pages tophleliidactophlebiidae with the description of a new species and their biogeographic affinities 151 169 in canadian journal of zoology 64146064 1460 1468 L corkum and J ciborowski editors current direc- IIHARRISARRis LG 1984 new mexico water lightsrights new mex- tions in research on ephemeroptera canadian schol- press icoleoieo watelwater resources research institute las cruces ars toronto HENRYULNRY BC 1993 A revisionlevision ofneochoroterpes ephemerephemera 1995 taxonomy of the north and central amer- olteiaopteiaopteraaptera leptophlebndaeleptophlebiidae new status transactions of ican species of camelobaetidius ephemeroptera the american entomological society 119317119 317 333 baetidaeBaetidae entomological news 106178106log 178 192 HENRYililiiIII111 NRY BC AND BC kondraiilffkondratieffkondratiefe 1982 new state 1995d new species stage descriptions and rec- records of the mayfly leptophlebia bradleyi need- ords of baetodesBaetodes ephemeroptera Baetbaetidaeidae from ham entomological news 9312593 125 126 mexico and central america entomological news idlIDIIDE FP 1930 contribution to the biology of ontario may 10681 86 flies with descriptions of new species canadian ento- 1996a new central american and mexican rec- mornmologistlogist 6220462 204 213 ords of ephemeroptera species entomological news 1935 post embryonic development of ephemer- 107303107 303 310 optera maymayfliesflies external characters only canadian 1996b contribution to the taxonomy of callibaetis journal of research 1243312 433 478 ephemeroptera baetidaeBaetidae in southwestern north 1937 descriptions of eastern north americanamerlean america and middle america aquatic insects 18 species of baetinebactinebaccine maymayfliesflies with particular reference 1 9 to the nymphal stage canadian entomologist 69 LUGOORTIZLUGO ORTIZ CRR WPWE mccafferty AND RDR D WALTZ 219 231235 243 1994 review of the Panpanamericanpanamencanamericanamerlean genus fallceonFallceon 199719971 NEW MEXICO MAYFLIES 313

ephemeroptera baetidaebaetidacBaetidae journal of the new york mccafferty WPWE RK HETIIHETH AND RD WALTZ 1997 entomological society 102460102 460 475 the ephemeroptera of spring creekgreek oklahoma with MAYO VK 1969 nymph of thraulodes speciosusspeciosus traver remarks on notable records entomological news with notes on a symbiotic chironomid pan pacific losios108193108 193 200 entomologist 4510345 103 112 mccafferty WPWE MJ WIGLE AND RD WALTZ 1994 mccafferty WP 1968 the mayfly genus hexageniahexagemahexagyniaHexnexageniaagema in systematics and biology of acentrella turbidasurbida mexico proceedings of the entomological society of mcdunnough ephemeroptera raetidaebaetidaeBaetidae pan washington 7035870 358 359 pacific entomologist 7030170 301 308 1975 the burrowing maymayfliesflies ephemeroptera mcdunnough J 1921 two new canadian maymayfliesflies ephemeroidea of the united states transactions of ephemeridae canadian entomologist 5311753 117 120 the american entomological society 101447lolloi101 447 504 1924a new canadian ephemeridae with notes 1990 biogeographic affinities of the ephemer- II11 canadian entomologist 5690 98 113 122 optera of the black hills south dakota entomolog- 128 133 ical news loi101193101 193 199 1924b new north american ephemeridae cana- 1991 comparison of old and new world acan dian entomologist 5622156 221 226 thametropus ephemeroptera acanthametropodi 1926 new canadian ephemeridae with notes IV dae and other psammophilous maymayfliesflies entomolog- canadian entomologist 5829658 296 303 ical news 102205102 205 214 1927 A new heptageniaHeptagenia from the yellowstone 1994 distributional and classificatory supple- rexionlegionregion ephemeroptera canadian entomologist ment to the burrowing maymayfliesflies ephemeroptera 59261 ephemeroidea of the united states entomological 1929 notes on north american ephemeroptera news 1051losios105 1 13 with descriptions of new species II11 canadian ento- 1996 ephemeroptera of amer- the species north momologistlogist 6116961 169 180 ica and index to their complete nomenclature trans- 1931a new species of north american ephemer- actions of the entomological society of america optera canadian entomologist 638263 82 93 1221122 1 54 1931b the eastern north american species of mccafferty WPWE AND R DAVIS 1992 new and addi- JRJ ephemerellaEphemerella and their nymphs ephemeroptera tional records of small minnow maymayfliesflies ephemer- minnow canadian entomologist 6318763 187 197 201 219 optera Baetbaetidaeidae from texas entomological news 1934 new of north 103199103 199 209 species american ephemeroptera IV canadian entomologist 6615466 154 164 mccafferty WPWE AND CRC R LUGOORTIZLUGO ORTIZ 1992 registrosRegistros 181 188 nuevoscuevos y notas sobrebobre los ephemeroptera de nica- 1935 notes on western species of ephemer- ragua revistadevista nicaraguensenicaragfiense de entomologfaentomologiaentomologic 19119igi 1 7 optera canadian entomologist 679567 95 104 1996 ephemeroptera pages 133 145 in J E in JE 1938 new of llorente bousquetsbousqueto ANA N garcia aldrete and E species north american ephemeroptera with critical notes canadian entomologist gonzalez soriano editors biodiversidad taxonomiataxonornfataxonomictaxonomia 702370 23 34 y biogeografia de artroartr6podosartropodospodos de mexico hacia una MCNAMEE G 1994 gila the life and smtesissintesissintenissintesis de su conocimiento universidad nacional death of an ameri- new york aut6nomaautonomaAutautonomyonoma de mexico mexico DFD F can rivernver orion MORIHARA DKD K AND WPWE mccafferty 1979a baitisbaetisbcietis mccafferty WPWE AND JFJE MACDONALD 1994 new the larvae of th idae records of ephemeroptera in utah with notes on noinorth america ephemeroptera Baetbaetidae biogeography entomological news 105217losios105 217 221 transactions of the american entomological society 105 139 mccafferty WPWE AND AV PROVONSHA 1993 new losios105139 221 species subspecies and stage descriptions of texas 1979b systematics of the propinquuspropinquous group of baetidaebaetidacBaetidae ephemeroptera proceedings of the ento- baetisbabtis species ephemeroptera baetidaeBaetidae annals of momologicallogical society of washington 955995 59 69 the entomological society of americaofamerica 7213072 130 135 mccafferty WPWE AND RD WALTZ 1986 babtisbaetis mag- NEEDHAM JGJ G 1905 ephemeroptera pages 17 62 in JGG nus new species formal new name for baetisbabtis sp B needham K J morton and 0 A jojohannsenharmsen editors of morihara and mccafferty ephemeroptera baetibaebi Maymayfliesflies and midges of new york new york museum dae proceedings of the entomological society of bulletin 86186 1 352 washington 8860488 604 1927 the rocky mountain species of the mayfly 1990 revisionary synopsis oftheodtheof the baetidaeBaetidae ephe- genus ephemerellaEphemerella annals oftheodtheof the entomological soci- meropteramerop tera of north and middle america transac- ety of america 2010720 107 117 tions of the american entomological society 116 1932 three new americanamerlean maymayfliesflies ephem 769 799 eroparop canadian entomologist 6427364 273 276 1995 labiobaetis ephemeroptera baetidaeBaetidae new NEEDHAM JGJ G AND ROR 0 christenson 1927 economic status related new genus and new north american insects in some streams of northernofnorthern utah bulletin of species entomological news 10619106log 19 28 the utah agricultural experiment station 2016201 6 16 mccafferty WPWE R DURFEE AND BC kondratieff NEWELL R L 1970 checklist of some aquatic insects from 1993 colorado maymayfliesflies ephemeroptera an anno- montana proceedings of the montana academy of tated inventory southwestern naturalist 3825238 252 274 science 304530 45 56 mccaffertymccaffMcCArr erryERTY WPWE RWR W FLOWERS AND RDR D WALTZ 1992 PESCADOR MLM L AND WL PETERS 1980 A revision of the biogeography of Mesomesoamericanmesoamencanamerican maymayfliesflies pages the genus homoeoneuriahomoeoneuna ephemeroptera aligooligo 173 193 in SPS P darwin and ALA L weldon editors neuneuriidaeneurudaerudae transactions of the american entomol- biogeography ofmesoamencaof M esoamerica proceedings of a sym- ogical society 106357106log 357 393 posium tulane studies in zoology and botany sup- PETERS WL AND GFG F EDMUNDS JR 1961 the maymayfliesflies plementalplemental publication 1 ephemeroptera of the navajo reservoir basin new 314 GREAT BASIN naturalist volume 57

mexico and Coloiacoloradodo university of utah anthropol- WALLEY GSG S 1930 review of the ephemerellaEphemerella nymphs of ogical papelpaper 55 upper coiocoloradoColotadoiddo seriesserlessenes 51075 107 111 western north america ephemeroptera canadian PROVONSHA AXAV 1990 A revisionlevision of the genus caeniscanniscaems in entomologist 621262 12 20 noinorthth america ephemeroptera caenidaeCaenmaenidaeidae trans- WALSH B D 1862 list of the pseudoneuroptera of illinois actions of the americanamerlean entomological society 116 contained in the cabinet of the writer with descrip- 801 884 tions of over forty new species and notes on their pkovonsiiap11ovonsha AXAV AND WP mccalmccafferty1 ertyLRTY 1982 new structural affinities proceedings of the academy of species and previously undescribed larvae ofofnoithnorth natural sciences of philadelphia 13 1436114 361 402 american ephemeroptera journal of the kansas 1863 observations on certain NAN A neuroptera entomological society 552355 23 33 by H hagen MDM D of koenigsberg prussia trans- ralliRAIIIRAHEL ejFJ AND CSC S KOLAR 1990 tradeoffstrade offs in the related from the original french MS and published sponse of maymayfliesflies to low oxygen and fish predation by permission of the author with notes and descrip- Occooecologiaoccologialogia 843984 39 44 tions of about twenty new NAN A species of pseudo- SAY T1 1924 in narrativeNanative on an expedition to the source neuroptera entomological society of philadelphia of the st peters rivelriver lake winnipeg lake of the proceedings 21672 167 272 woods etc performed in the year 1823 by order of WALTZ RD AND WP mccafferty 1987 revision of the hmlim JC calhoun secleSeciesecretarytary of waiwalwarwan undertinderlinder the the genus cloeodesCloeodes traver ephemeroptera baetibaebi command of stephen H long majolmajor USTE by dae annals of the entomological society of amer- WWMM keating philadelphia 22682 268 378 leaicaiea 8019180 igligi191 207 SEEMANNSILMANNTMTM 1927 Dragondiagonfliesdragonfliesflies stonestonefhesstoneflicsfliesflics and maymayfliesflies WARD JV AND L BERNERBLRNLR 1980 abundance and altitu- of sousouthernthein california journal of entomology and dinal distribution of ephemeroptera in a rocky zoology 194019 40 51 mountain stream pages 169 186 in JEJ F flannagan SNEADSNI AD RER E 1979 landformsLandforms page 6 in JLL williams and and KEK E marshall editors advances in ephemer- PE mcallister editorsediedltoistols new mexico in maps tech- optera biology plenum new york nology applications centelcentergentelgenter university of new mex- WARD JV AND JA STANFORD 1990 ephemeropteraephcmeroptera of icoieoleo albuquerquealbuqueiAlbuquei que the gunnison river colorado USA pages 215 220 splethSPIETH ILTII11 T 1941 taxonomic studies on the ephemer- min ICI1 C campbell editor Maymayfliesflies and stonestonefhesstonefliesflies life optera II11 the genus hexageniahexagemahexagyniaHexnexageniaagema americanamerlean midland historieshistones and biology kluwer dordrechtDordrecht the Natnaturalistmalist 2623326 233 280 netherlands suiulriljemdsublelsubiel M JE MD hxrciihaiciiandmannAND M SUBLEITESUBLETTE 1990 the WOODBURY AMA M AND DWD W ARGYLE 1963 ecology and fishes of new mexico university of new mexico limnology of green river pages 7 48 inm studies of pipressess albuquerque biota in dinosaur national monument utah and colo- TRAVERTKAVLHJRJR 1934 new north american species of may rado university of utah division of biological sci- flies ephemeridaeephemeiEphe meimel idae journal of the elisha mitchell ences miscellaneous paper 1 salt lake city scientific society 5018950 189 254 YANOVIAK SPS P AND WP mccafferty 1995 stream size 1935 part II11 systematic pages 237 739 in JGG and the distribution of selected ephemeroptera needham JR traver and YC hsu editorsediedltoismols the pages 225 236 in L corkum and J ciborowski edi- biology of maymayfliesflies with a systematic account of tors current directions in research on ephemer- northnoi th americanamerlean species comstock publishing co optera canadian scholars press toronto ontario ithaca new yorkyoimoimol k ZLOTY J 1996 A revision of the nearctic ameletus may THAVLTRAVER it JRJ R AND GEG F EDMUNDS JHJR 1967 A revision of flies based on adult males with descriptions of seven the genus thraulodes ephemeropteraephemeiEphemei apteraoptera leptophlebi new species ephemeroptera ameletidae canadian idae miscellaneous publications oftheodtheof the entomologi- entomologist 128293128 293 346 cal society ofofamencaofAamericamenca 53495 349 395 1968 A revisioni evisiondevision of the Baetbaetidaeidae with spatulate received 8 april 1997 clawed nymphs ephemeroptera pacific insects 10 accepted 27 may 1997 629 677 walWAIWALKERKL n FE 1853 catalogue of the specimens of neur- opterousoptopte iouslouserous insects in the collection of the british museum paitpaltpart 34773 477 585 great basin naturalist 574 0 1997 appp 315 326

HOLOCENE vegetation AND HISTORIC GRAZING IMPACTS AT CAPITOL REEF NATIONAL PARK reconstructed USING PACKRAT MIDDENS

kenneth L colel norman Henderson 2 and david S shafertshafer3

ABSTRACT mid to late holocene vegetation change from a remote high desert site was reconstructed using plant macromicrofossilsmacrofossilsfossils and pollen from 9 packratpankrat biddensmiddens ranging from 0 to 5400 yr in age presettlement biddensmiddens consistently contained abundant macromicrofossilsmacrofossilsfossils of plant species palatable to large herbherbivoresivores that are now absent or reduced such as winterwinterfatfat Ceratceratoidesoides lanata and ricegrassncricriencegrassegrass stipa hymenoideshymenoides microfossilsmacrofossilsMacro fossils and pollen ofofpmyonpinyon pine pinus edadulisedulisulis sagebrush artemisia sppapp and roundleafroundleaf buffabuffaloberryloberry shepherdia rotundifoliarotundifolidrotundifoliafoila were also recently reduced to their lowest levels for the 5400 yr record conversely species typical of overgrazedovergrazed range such as snakeweed Gutiergutierreziagutierrezbarezia sarothraesarothrae viscid rabbitbrushrabbitbrush chrysothamnus visidiflorusvistdiflorus and russian thistle salsola sp were not recorded prior to the historic introduction of grazing animals pollen of utah juniper juniperus osteosperma also increased during the last 200 yr these records demonstrate that the most severe vegetation changes of the last 5400 yr occurred during the past 200 yr the nature and timing of these changes suggest that they were primarily caused by 19th century open land sheep and cattle ranching the reduction ofofpmyonpinyon and sagebrush concurrentconcu nent with otherothel grazing impacts suggests that effects of cattle grazing at modern stocking levels may be a pooipoor analog for the effects of intense sheep grazing during drought

key words holocene vegetation history grazing impacts pankratpackrat biddensmiddensmid dens fossil pollen presettlement vegetation

STUDY AREA torreyanatorreyana viscid rabbitbrushrabbitbrush chrysothamnus visidiflorus and central pricklypricklypearpricldypearpear opuntia this purpose of this study was to produce a polyacanthapoly acantha table 1 low areas with thicker holocene vegetation history of capitol reef soil support a sparse growth of grasses rice national park reconstructing past changes in grass stipa hymenoideshymenoides sand dropseed sporo- vegetation and relating those changes to their bolus cryptcryptandrusandrus and blue grama bouteloua most probable causes packratpankrat midden chronol- gracilis plant taxonomy follows welsh et al ogies were developed from several sites cole 1987 1992 but only the most complete series from mean annual precipitation at the site is close a single site collected from the hartnet draw to the 18 cm yr 1 recorded at fruita 15 km to site is reported here seven fossil and 2 modemmodern the southwest at 1670 m elevation heil et al biddensmiddens were collected from hartnet draw in 1993 precipitation is bi seasonal with winter northern capitol reef national park 3815n and late summer peaks temperature extremes IHWWI1 I1 120w fig 1 this site at 1920 m elevation are great at this andaridarld continental site a mean in wayne county utah was chosen because january minimum temperature at fruita of of its remote location free from most anthro- 8cac and a mean july maximum of33cof 33c pogenic disturbances other than grazing and GRAZINGCRAZING HISTORY historical records men- the abundant fossil packratpankrat biddensmiddensmiddens tion no disturbances to this area other than the the site is underlain by the salt creek mem- introduction of exotic herbherbivoresivores during the ber of the morrison formation which forms mid 19th century native large herbherbivoresivores that many overhangs protecting the fossil packratpankrat may have been present in the study area dur- biddensmiddens fig 2aaa today the most abundant ing the last 5000 yr include bighorn sheep plant species are utah juniper juniperus osteoosoteo ovis canadensis mule deer odocoileus hemi- spennaspsperspermdspermaaspennaennamd bigelow sagebrush artemisia biglovbigloviiii onus pronghorn antilocapra americanaanzericanaanzericana and big sagebrush A tritrlfridentata snakeweed cutiguticutiguti possibly bison bison bison and elk cervus errezia sarothraesarothrae torrey ephedra ephedra elaphusclaphus van gelder 1928 mead et al 1991

geological survey green lusius MNCPSU 115 hall department of foiestforest resourceresources university of minnesota st paul MN 55108 presentpiesentaddiessaddress Coloiacoloradoclo plateau field station USGS BRD box 5614 northernnolNoi thein arizona university flagstaff AZ 86011 rational2national park selvleeservice glen canyon national recreation aleaarea box 1507 page AZ 86040 sqffice doffice3office of environmental restoration US department of energy richland operations office box 550 MS s7sa 54 richland WA 99352

315 316 GREAT BASIN naturalist volume 57

capitol reef national park

hartnet dra UTAH

wayne county garfield county

hailshallshalis creek

fig 1 map showing site location

native north american equilsequids horses and prior to the passage of the taylor grazing act in asses became extinct before 10000 yr BYB P 1934 large numbers of livestock were brought from wayne emery 1984 sevier and counties to winwin- mead and meltzer eurasian horses ter on these lands many of the animals remained and asses equus sppapp were introduced to on the range yearlong resulting in the progressive new mexico by spanish colonists as early as destruction of soils and vegetation reports from ADA D 1598 underhillunderbill 1971 by the late 1600s stocklenstockmenstockmen in the area indicate that many trespass feral horses were reported in parts of the horses used the area until about 1955 prior to in 1946 there were at least 163 cattle and 20 horses is likely west but it is not that they existed in yearlong in this area hartnet allotment file 1966 the study area prior to the 1800s in the late 1800s introduced herbivore popu- currently the area is grazed under an allot- lations increased dramatically in southern utah ment issued by capitol reef national park with the widespread increase in open land graz- reconstructing PAST vegetation fossil ing livestock grazing within and near capitol packratpankrat biddensmiddens are valuable sources of pale reef national park has been documented since oecologicaloncological information in andaridanidannd regions of the at least the 1870s frye 1995 the earliest southwestern united states betancourt et al detailed herbivore population estimates from 1990 cole 1990 plant fossils in packratpankrat mid the capitol reef area are from summer graz- dens often identifiable to the species level ing permits issued for powell now part of grew close to the midden most likely within dixie national forest frye 1995 in 1909 50 m because plant identification and loca- the forest service issued permits for 67000 tion can be precisely known this method has sheep and 11000 cattle the animals grazing extremely high spatial and taxonomic resolution these high summer pastures presumably spent compared to other methods of reconstructing the winter in the lower adjacent areas of capi- past vegetation tol reef national park studies comparing trees and shrubs at mid- A bureau of land management survey de- den sites with plant specimens from modern scribed past use at the hartnet draw site biddensmiddens typically report similarities exceeding 19971 HOLOCENE vegetation AT CAPITAL REEFreep 317

473

2zaz

04 1 WM RK

k ai0i tat1 pl g V V ae aaa4

fig 2 A photograph of hartnet draw study site showing shelter where biddensmiddens s 1 4 were collected B recon- strucstruestructedted image of study site with presettlement vegetation as visualized from fossil data 318 GREAT BASIN naturalist volume 57

TABLETABLL 1 plant macromicrofossilmacrofossilfossil concentrations from hartnet draw biddensmiddens in 1090logo concentrationconcentrationkgkg of washed matrix modemmodern percent cover classes are estimated from a relev6relevedeleve ofof3030 m radius coverage classes 3 3 cover 2 1 3 cover I1 1 cover R rare located outside of relev6relevedeleve midden 3 2 8 la ab7b 9 5 6 midden age yr BPB P 0 330 630 1020 1275 2570 3615 5450 modern relev6relevedeleve cover class logology concentrationconcentrationkgkg of washedofwashed matrix trekstreetTRLLS SHRUBS AND succulentsSU CULENTS artemisia sec tridenttridentataetndentataeatae 3 09 18 28 28 18 13 20 atriplex sppapp 2 28 33 30 43 31 32 28 41 Bricbnckelliabrickelliakellia scabiascabra 2 13 chrysothamnus sp 2 16 dowaniacowania mexicansmexicanamexicana 2 20 19 19 26 31 25 29 12 ephedra sppapp 2 17 05 31 13 23 Ceratceratoidesoides lanata 20 17 36 24 25 23 30 Gutiergutierreziagutierrezbarezia sarothsarothraerae 3 20 13 heterothecaHetero theca sp 1 09 18 hymenopappushyrnenopappus 1 19 juniperusJumperus osteosperma 3 34 39 38 43 43 43 40 41 machaeranthera grindgrindeliodeseliodes 1 04 opuntia polyacanthapolyacantha 2 31 37 31 43 42 35 39 39 pediocactusechinocereuspediocactuslechinocereus 1 11 14 21 24 21 finuspinus eduiseduhsedulis R 20 28 47 26 41 30 20 salsola sp 1 13 sarcobatus sp R 10 Shephershepherdiadw rotundifoliarotundifolia 1 25 18 30 yucca angustissimaangu&tissima R 10 19 herbsHLRBS AMAND GRASSESGRASSLS cf amaranthus sp 23 17 19 16 16 astragalus mollissimusmollissimus 1 09 aristida purpurpurpureapurpureanea 2 bouteloua gracigraciasgracihsgracifisfishis 3 17 14 22 chryptantha sppapp 3 08 05 13 12 descuraniadescurama pinnatepinnata 3 eriogonum sp 10 18 etloerloerioneuronEnoErioenoneuronneuron pulchellumpulc hellum 1 euphorbia sp 1 17 17 hilaria sp 13 hymenoxys acauhsacaulis 1 lappula occidentoccidentalisoccidentalistalis 1 04 09 08 18 lepidium densiflorum 1 18 05 13 23 peanut shell 09 phacelia 13 plantago 09 14 sphaeralcea coccineacoccinea 1 13 12 22 25 sporobolissporobohssporobolus sppapp 3 09 13 stipa hymenoideshymenoides 3 15 17 16 13 28 19 17 stipa sp 13 12 streptanthella longirolongirostnslongirostrisstrisstoisstols 1

80 using a Sorensorensenserss index of similarity cole cation of diverse forbs and grasses within mid- 1985 cole and webb 1985 spauldingSpauldmg et al den assemblages is very challenging 1990 frase and sera 1993 especially when the quantitative correlation between species small macromicrofossilsmacrofossilsfossils 2 mm are identified using abundance and midden specimens is complex a lox microscope similarity with forbs and spaulding et al 1990 plant species produc- grasses has been reported to be lower frase ing abundant readily identifiable plant parts and sera 1993 but inventories of current forbs eg atriplex leaves or pankratpackrat food items are usually incomplete due to seasonal and eg juniper or plants having deterrence val- yearly variability in the forb flora and identifiidentify ues in protecting pacpackratspankratskrats from predators eg 199719971 HOLOCENE vegetation AT CAPITAL REEF 319 cactus spines tend to occur in the highest gregregatedgated in water two unconsolidated mid numbers within biddensmiddensmiddens but perennial species dens hartnet draw s 3 and 4 were consid- that are abundant near biddensmiddens are most often ered modern because of the presence of green represented by high numbers of plant speci- leafy material cow feces and a peanut shell mens while less abundant species or those fur- pollen samples were taken from the wash ther from biddensmiddensmiddens are represented by fewer water after several days of soaking and the specimens As a result interpreting changes pollen was separated using standard methods from midden assemblages requires experience faegrifaegre and iversen 1975 microfossilsmacrofossilsMacro fossils of 1 with macromicrofossilmacrofossilfossil types leaves twigs flowers modern midden hartnet draw 4 could not seeds and abundances typically found for that be analyzed because the midden had been species this element of judgment is present burned chairingcharringcharring much of the plant debris in any retrospective study for example fossil the pollen content of this midden however pollen magnifies the presence of wind polli was not destroyed by the fire packratpankrat debris natedbated plants while insect pollinated species piles are often burned in rangeland shelters may not be represented at all photolithphytolithPhytolith stud- most likely to eliminate the rats that occupy a ies detect only those species producing identi- pleasant shelter fiable opal phytoliths historical writings record after sieving vegetable debris fecal pellets only those species of interest to the writers and rocks from the dissolved biddensmiddens with a repeat photography is useful only for those imm1 mm sieve we mixed dried and weighed species identifiable in photographs midden the resulting matrix producing 100 200 g of viewed records can be as representing some- washed midden matrix the dried matrix was thing similar to a plant releverelev6deleve mueller dom sorted by hand under a loxIOX dissecting micro- bois and ellenberg 1974 from the past this scope pankratpackrat fecal pellets and rocks were releverelev6deleve list also contains a detailed species but removed and weighed identifiable plant macro more complex information on past species fossils vertebrate bones and insect fossils were abundance comparable to coverage classes identified counted labeled and stored in plastic fossil pollen within biddensmiddens can also be vials analyzed king and van devender 1977 six to 13 g of packratpankrat fecal pellets were sub- thompson 1985 davis and anderson 1988 mitted to radiocarbon laboratories for dating emphasizing different types of vegetation and hartnet draw 5 was dated at 3615 70 yr representing a larger source area than the BPBY using a single pinuspintis needle after the initial plant microfossilsmacromacrofossilsfossils interpretation of fossil pollen pellet sample yielded an impossible result of abundances like macromicrofossilmacrofossilfossil abundances re- 142 modern carbon some type of sample quires caution and experience as some species with are better represented than others by consid- contamination artificial carbon isotopes sample ering both macromicrofossilmacrofossilfossil and pollen records we or mismislabelinglabeling is suspected as it is impossible can achieve a more comprehensive under- to contaminate an old sample with standing of past environments enough modern natural carbon isotopes to yield such a high number calendar year ranges for MATERIALS AND METHODS radiocarbon ages were calculated using stuiver and reimer s 1993 calibration program we collected 7 of the 8 biddensmiddens within a data on midden contents were quantified radius of about 200 in from a small shelter by number weight percent of identified spec- fig 2 the ath8th was found I1 km east of the imens and 1090logo of macromicrofossilmacrofossilfossil concentration shelter using a hammer and chisel we sepa- in midden matrix to compensate for variability rated approximately 1 kg of each midden from between biddensmiddensmiddens we adjusted midden matrix larger masses and returned the samples to the weights by subtracting the weight of rocks and laboratory samples were then dissected pro- pellets from the dried washed matrix weight ducing horizontally stratified subsamples typi- before calculating the concentration as sug- cally measuring about 15 x 20 cm and several gested in betancourt 1990 using 1090logo of centimeters thick weathering rinds and large macromicrofossilmacrofossilfossil concentration calculates a number rocks were removed from each subsample similar to the semiquantitativesemi quantitative abundance scale yielding 300 600 g of hardened midden mate- used by several other authors but it has the rial this sample was then weighed and disagdicag advantage of being quantitative 320 GREAT BASIN naturalist volume 57

TABITABLE L 2 ages of biddensmiddens based on radiocarboni adioadlocarbonearbon dates radiocarbon ages are calibrated to calendar years based on stuiver and reimer 1993 MC percent modemmodern carbon sample postdates atmospheric testing of nuclearofnuclear weapons calendar year range sample name radiocarbon date 813c8 13c lab ID no at one sigma material dated

hartnethaihal tnetanet drawdi AW 4 modernmodem debusdebris pile not dated hartnet draw 3 137 121 2 MC 222 A 5197 AD 1960 1986 neotoma pellets hartnetdiaw2hartnet draw 2 330 60 218 A 5204 AD 1495 1643 neotoma pellets iiaitnetdiaw8hartnet draw 8 630 100 212 GX 16259 AD 1280 1410 neotoma pellets hartnet draw la 1020 70 217 A 5203 AD 898 1152 neotoma pellets iiaitnetdiaw7bhartnet draw 71 1275 110 208 GX 15554 AD 640 890 neotoma pellets iiaitnetdiaw9hartnet draw 9 2570 135 218 GX 15553 889 434 BC neotoma pellets hartnet draw 5 3615 70 AA 6447 2128 1889 BC pinus adulisedulis needle hartnet draw 6 5450 90 218 A 5205 4363 4235 BC neotoma pellets

we used the program CONISS grimm settlement similarly globe mallow sphaeral- 1987 on the plant macromicrofossilmacrofossilfossil and pollen results cea sp needleneedlegrassgrass stipa sp blue grama to conduct a stratigraphically constrained clus- bouteloua gracilis dropseed Sporasporabolisbolis crapcryp ter analysis using a square root transformation tantandrusgandrusdrus and roundleafroundleaf buffabuffaloberryloberry shepher- and edwards and cavalli sforzas chord dis- dia rotundifoliarotundifolia are common in presettlement tance as a dissimilarity coefficient the square biddensmiddens but absent from the 1 modemmodern midden root transformation makes the skewed distri- in contrast hartnet 3 the modern mid- butions of abundant species more closely con- den is the only midden containing viscid rab form to normal distributions plant taxa occur- bitbrushbitbrush chrysothamnus visidiflorus grease- ring in only a single midden sample were de- wood sarcobatus vermiculatus and russian leted from the analysis to eliminate false posi- thistle salsola sp macromicrofossilsmacrofossilsfossils only the 2 tive correlations due to shared absences these most recent biddensmiddens 3 and 2 contain deletions cause any differences between the snakeweed Gutiergutierreziagutierrezbarezia sarothraesarothrae rabbit modern and fossil biddensmiddens to be understated brush snakeweed and russian thistle are fre- quent at the site today the absence of these RESULTS species from presettlement biddensmiddens indicates that these species were formerly absent or so MIDDEN AGE the 9 biddensmiddens ranged in infrequent as to not be represented age from modern to 5450 yr BYB P table 2 the cluster analysis right side fig 3 dem- hartnet draw 3 contained 137 modern onstratesonstrates the difference between the modern s carboncalbonearbon modern is defined as ADA D 1950 and macromicrofossilmacrofossilfossil assemblage and other assemblages thus postdates atmospheric testing of nuclear it is the primary branch in the dendrogram weapons seven biddensmiddens dated to presettle- even though the single occurrences of rabbit ment times brush greasewood and russian thistle in the PLANIPLANT microfossilsmacrofossilsMACRO FOSSILS all of the biddensmiddens modern midden were disregarded in the contain abundant macromicrofossilsmacrofossilsfossils of utah juniper analysis Jumjunipentsjumperusperasperus osteosperma saltbush atnplexatfiplexatriplex sppapp POLLEN SAMPLES results of the pollen cliff rose dowaniacowaniaCowania mexicansmexicanameximeyicana and prickly pear analysis fig 4 are similar to those from plant opuntia sp which are all plentiful at the site macromicrofossilsmacrofossilsfossils presettlement biddensmiddens contained today table 1 fig 3 the presettlement mid much more pine and sagebrush pollen than dens also contain pinyon pine pinus edadulisedulisulis the 2 modern samples 3 and 4 similarly Winterwmterfatwinterfatfatfht Ceratceratoidesoides lanata sagebrush arte- percentages of grass and buffaloberrybuffaloberry pollen misia sp and riericricegrassncncegrassegrass stipa hymehymenhymenoideshymenozdesozdesnoides were generally higher in presettlement mid which are rare or absent from the single mod- dens in contrast only the 2 modern biddensmiddens ern midden analyzed for macromicrofossilsmacrofossilsfossils hartnet contained pollen of the exotic russian thistle 3 winterfatWinter fat was not observed during the and high amounts of juniper pollen like the fieldwork and pinyon pine was rare in the macromicrofossilmacrofossilfossil cluster analysis the pollen cluster area the rarity or absence of sagebrush and analysis right side of fig 4 showed that mod- ricriericegrassncncegrassegrass from the modern midden suggests ern samples are very different from all preset- that they are less common now than prior to tlement biddensmiddensmiddens HOLOCENEz vegetationz AT CAPITAL REEF

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discussion 1666 16701684 168517281685 1728 17291822 1823 1879 1880 and 1899 1900 figure 2 shows the site as it is now and a less is known about climatic variability in reconstructed image of how it may have looked this region over the previous 5000 yr prior to prior to settlement the reconstructed image this tree ring record but it seems unlikely shows greater coverage of grasses winterwinterfatfat that any climatic event of the last 200 yr was sagebrush and pinyon inferred from the macro sufficient to cause a change with no precedent fossils and pollen found in the packratpankrat mid during the previous 5000 drought may have dens although vegetation probably fluctuated precipitated some of the dramatic vegetation continuously throughout the late holocene changes of the last 200 yr but it did not set the this midden record suggests that previous stage for them this would require an event changes were minor compared to those of the unprecedented during the previous 5000 yr last 200 yr sites similar in appearance to the FIRE HISTORY some changes recorded in reconstructed image are now present on un- the biddensmiddens could have been caused by changes grazed terraces that are inaccessible to large in fire regime the increase in juniper could herbherbivoresivores along halls creek 90 km to the have resulted from a decrease in fire frequency south heil et al 1993 the presettlement caused by elimination of grassy fuels by graz- plant community was undoubtedly more like ing but this does little to explain the shift from the pinyon juniper grass community described palatable to palatablenonnonpalatable species or the reduc- by heil et al 1993 than the juniper shrub tions of pinyon sagebrush and buffaloberrybuffaloberry community present at the site today just at the time that fire frequency decreased it is clear from macromicrofossilmacrofossilfossil and pollen analy- fires of unprecedented severity could have ses reinforced by the 2 cluster analyses figs been set during the settlement era but this 3 4 that the modern midden plant contents hypothesis has no data to support it studies of are dramatically different from the presettle- tree fire scars or sedimentary charcoal would ment biddensmiddensmiddens furthermore the presettlement have to be conducted to test this possibility biddensmiddens are more similar to each other than to GRAZING IMPACTS impacts from introduced either of the modern biddensmiddensmiddens this suggests herbherbivoresivores especially large sheep herds in the that the magnitude of change in vegetation late 19th and early 20th centuries are the most during the last 200 yr was far greater than dur- likely cause of recent radical vegetation changes ing the previous 5000 yr hypothetical causes the introduction of sheep goats cattle and of this vegetation change should account for horses was without precedent during the pre- both the timing of the change and the specific vious 5000 yr overall vegetation has shifted taxa that increased or decreased from palatable toward less palatable forage DROUGHT HISTORY reductions in winter specifically palatable grasses winterwinterfatfat and fat pinyon pine sagebrush and ricricegrassegrass and buffaloberrybuffaloberry decreased while less palatable increases in juniper rabbitbrushrabbitbrush and snake- species rabbitbrushrabbitbrush snakeweed and grease- weed might be attributed to droughts during wood increased rabbitbrushRabbitbrush and greasewood the 19th or 20th centuries however an analy- are poor forage while snakeweed is typically sis of past drought frequency for southeastern an invader or increaser on overovergrazedgrazed range utah fig 5 using 400 yr of tree ring data benson and darrow 1981 heil et al 1993 compiled by fritts 1991 suggests that droughts cronquist et al 1994 of the 19th century were not unusually severe other studies conducted on grazing at capi- when compared to the 17th century severe tol reef support this conclusion heil et al droughts defined here as years with 165 1993 in a survey of the vegetation of capitol nunmm of annual precipitation reconstructed for reef national park suggest some of the most moab utah occurred 9 times in the 17th cen- preferred plant species for glazersgragrazerszers eg cera tury 4 times in the 18th century and 7 times toidesboides lanata and stipa comatacamata may have been in the 19th century the 5 driest years recon- locally extirpated by grazing this packratpankrat mid- structedstructed from the tree ring record were AD den record demonstrates that for the hartnet 1667 132 mm 1684 142 mm 1668 143 mm draw site this was the case 1879 147 mm and 1861 150 mm excep- additional research at capitol reef national tiotionallynally dry successive years were 1624 1626 park indicates grazing has caused and may 324 GREAT BASIN naturalist volume 57

reconstructed precipitation for moab utah 12 V 300

11 280

26060 E 10 E 240 c 0 0 9 220 t CL 8 200 CL

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160 6 140

5 i 1600 1650 1700 1750 1800 1850 1900 1950 calendar years

fig 5 reconstruction of the last 400 yr of precipitation at moab utah from tree ring data generated by a program distributed by fritts 1991

still be causing changes to the natural habitat sequent reductions in fire frequency but stud- plant photolithphytolithphytol ith analysis in buried soil horizons ies discriminating between pinyon and juniper shows a reduction of palatable grass species do not portray identical histories for both over the last several hundred years fisher et al species pine and sagebrush both declined 1995 an analysis of riparian areas indicates while juniper dramatically increased during that dramatic changes had occurred prior to the settlement period at peck s lake arizona the taylor grazing act of 1934 forage plants davis 1987 A study of permanent plots in a were heavily used and in many instances cover presently ungrazed part of pine valley utah was entirely removed recent grazing has per- demonstrated a significant decrease in juniper petuapetuatedted this removal or reduction of species and significant increase in pinyon between 1933 and inhibited potential recovery barth and and 1989 yorks et al 1994 A study of tree mccullough 1988 in a lightly grazed area age structure on a presently ungrazed site in palatable shrubs and grasses have increased southwestern utah s needle range found that significantly during the 19th century many surviving juniper dramatic declines in pinyon sagebrush and and few pinyon were established by 1915 the buffaloberrybuffaloberry may also have been caused by the situation had reversed with far more surviv- grazing history but effects on these species ing pinyon becoming established in this cen- are less well understood these declines are tury tausch and west 1988 these results also present in 14 additional biddensmiddens from sites demonstrate that pinyon and juniper respond elsewhere in capitol reef national park such differently to changing regimes of grazing fire as along hall s creek murray 1989 cole 1992 or climate the observation that heavy grazing pinyon juniper woodlands have reportedly causes an expansion of pinyon juniper wood- increased during the historic period this is land west et al 1975 does not equate with especially evident when comparative photo- the expansion of both species in all habitats graphic techniques are used west et al 1975 these results suggest that pinyon may be tausch et al 1981 this increase in pinyon recovering now at some sites from a late 19th juniper woodlands is thought to be caused by centuryearlycentury early 20th century decline caused by reduced competition from grasses and forbs grazing impacts this recovery has not yet which were eliminated by grazing and by con occurred at hartnet draw this hypothesis has 199719971 HOLOCENE vegetation AT CAPITAL REEF 325 support despite the lack of recent observations spence debra maddox debra daugherty and of pine removal by grazing cattle knowledge betsy jernigan helped sort and count plant of the effects of cattle grazing at present stock- macromicrofossilsmacrofossilsfossils lyn murray completed the analy- ing levels forms an inadequate basis for judg- sis of some pollen samples robyn flakne ing the effects of an overoverstockedstocked sheep range assisted with copy editing and walter loope during the droughts of the late 19th century and an anonymous reviewer contributed sug- although cattle will consume some pine when gestionsgestions on the manuscript this project was it is available pfister and adams 1993 sheep funded by the national park service and the readily consume pine needles and strip pine national biological service bark even in the absence of drought conditions anderson et al 1985 sheep accomplished literature CITED the near complete elimination of the bishop pine forest pinus muricatemuricatamuricata on santa cruz ANDERSON GW H HAWKE AND RW MOORE 1985 pine island california where they were not fenced needle consumption and bark stripping by sheep annual pastures in young stands of widely 1980 grazing in out hobbs spaced pinus radiata and P pinaster Agroagroforestryforestry sagebrush populations may have a similar systems 3373 37 45 history despite observation of increases in BARTH RCR AND EJE J mccullough 1988 livestock graz- sagebrush caused by the removal of their grass ing impacts on riparian areas within capitol reef national park unpublished competitors young et al 1978 although sage- report to capitol reef national park torrey UT brush may be increasing on land presently BENSONBCNSON L AND RA DARROW 1981 trees and shrubs of grazed by cattle this is not an appropriate the southwestern deserts university ofarizonaof arizona press analog for intense 19th century sheep graz- tucson ing sagebrush is consumed by sheep during betancourt JLJ L 1990 late quaternary biogeography of the coloradodo plateau pages 292 droughts the late 19th Coloia 259 in JL betan- during century sheep court TR van devender and PS martin editors severely reduced populations of california fossil packratpankrat biddensmiddensmiddens the last 4000040 000ooo years ofbioticof biotic sagebrush artemisia californicacalifornica on santa rosa change university ofarizonaof arizona press tucson island california after first consuming the betancourt JL TR vandevender AND PS MARTIN grass cole and 1994 EDITORS 1990 fossil packratpankrat biddensmiddensmiddens the last 40000 liu years of biotic change university of arizona press MAGNITUDE OF CHANGE recent vegetation tucson changes recorded at capitol reef national park COLE KLL 1985 past rates of change species richness are unique when compared to natural changes and a model of vegetational inertia in the grand of the last 5000 yr these results echo those of canyon arizona american naturalist 125289125 289 303 1990 davis et al 1977 who found the vegetation reconstruction of past desert vegetation along the colorado river using packratpankrat biddensmiddens change caused by domestic livestock reflected palaeogeography palaeoclimatology and palaeoecol in fossil pollen at wildcat lake washington ogyagy 7634976 349 366 to be greater than any other event of the last 1992 A survey of the fossil pankratpackrat biddensmiddens and 1000 yr reconstruction of the prepregrazingpregrazmggrazing vegetation of capitol reef national park unpublished report to the it is also possible that both climate and graz- national park serviceselvleeSeivice capitol reef national park ing combined to produce the dramatic vegeta- torrey UT tion shifts of the last 200 yr there is little doubt COLE KLL AND G liu 1994 holocene paleoecology of that the most severe grazing damage occurs an estuary on santa rosa island california USA when high populations of herbherbivoresivores compete quaternary research 4132641 326 335 gollgoleCOLECOLL KLL AMAND RHR H WEBBWLBB 1985 holocene vegeta- for food during a severe drought drou ats late the droughts9hts tion changes in Greengreenwaterwater valley mojave desert of 1879 1880 and 1899 1900 probably exacer- california quaternary research 2322723 227 235 bated damage caused by high herbivore popu- CRONQUIST A AHA H HOLMGREM NHN H HOLMCREN AND lations more severe earlier droughts such as JLJ L REVEAL 1994 intermountain flora volume IV asterales new york garden new york those during the 17th century did not cause botanical davis OK 1987 palynologicalPalynological evidence for such changes historic because the large introduced juniperjkniper invasion in central arizona a late quater herbherbivoresivores were absent nary perspective pages 120 124 in the pinyon juniper ecosystem a symposium utah state univer- acknowledgments sity logan DAVIS OK AND RSR S ANDERSON 1988 pollen in pankratpackrat neotoma biddensmiddens pollen transport and the rela- we were assisted in the field by penny hoge tiontionshipship of pollen to vegetation palynology 11 dan huff rick harris bill romme and john 185 198 326 GREAT BASIN naturalist volume 57

DAVIS OK0 K DAD A KOLVA AND PJ MEHRINGER JR 1977 MURRAY LKL K 1989 report on fossil pollen from pankratpackrat pollen analysis of wildcat lake whitman county biddensmiddensrniddens from capitol reef national park unpub- washington the last 1000 years northwest science lished report to the national park service indiana 511351 13 30 dunes national lakeshoreLake shore porter IN FAEGRI K AND J IVERSEN 1975 textbook of pollen analy- PFISTER JAJ A AND DCD C ADAMS 1993 factors influencing sis hafner new york pine needle consumption by grazing cattle during FISHER RFR F CNC N BOURN AND WF FISHER 1995 opal winter journal of range management 4639446 394 398 phytoliths as an indicator of the floristics of prehis- SPAULDING WG JLJ L betancourtBETANCOUBT KLK L COLE AND L toric grasslands Geogeodermaderma 6824368 243 255 CROFT 1990 fossil packratpankrat biddensmiddens their nature FRASE BAB A AND WE SERA 1993 comparison between and methods of analysis pages 59 84 in JLL betan- plant species in bushy tailed woodratwoodral biddensmiddens and court TR van devender and PS martin editors in the habitat great basin naturalist 5337353 373 378 fossil packratpankrat biddensmiddensmiddens the last 40000 years ofbioticof biotic FRITTS HCH C 1991 climate reconstructions data and dis- change university ofarizonaofarizona press tucson play software program accompanying reconstruct- STUIVER M AND PJ REIMER 1993 A radiocarbon calibra- ing large scale climatic patterns from tree ring data tion program radiocarbon 3521535 215 230 a diagnostic analysis university of arizona press TAUSH RJR J AND NEN E WEST 1988 differential establish- tucson ment of pinyonpryonpmyon and juniper following fire american FRYE BJB J 1995 an administrative history of capitol reef midland naturalist ilg119174119 174 184 national park utah unpublished report to capitol TAUSCH RJR J NEN E WEST AND AAA A NABI 1981 tree age reef national park torrey UT and dominance patterns in great basin pinyon juniper GRIMM ECE C 1987 CONISS a fortran 77 program for woodlands journal of range management 3425934 259 264 stratigraphically constrained cluster analysis by the THOMPSON RSR S 1985 palynology and neotoma biddensmiddens method of incremental sum of square pergamon pages 89 112 in BLB L fine jacobs PL fall and OK0 K journals 131313 13 35 davis editors late quaternary vegetation and cli- HARTNET ALLOTMENT FILE 1966 wonderland resource mates in the american southwest american associ- area middle desert planning unit hartnet allot- ation of stratigraphic palynologists contributions ment plan hanksvilleHanksville office of the bureau of land series 16 management files UNDERHILLUNDERBILL RMR M 1971 red mans america university of HEIL KDK D JMJ M PROCTER R FLEMING AND WH ROMME chicago press chicago IL 1993 vascular flora and vegetation of capitol reef VAN GELDER RGR G 1928 mammals of the national parks national park NPSNFS technical report npsnaucare johns hopkins university press baltimore MD NRTR 9301 flagstaff AZ WEST NEN E KHK H REA AND RJR J TAUSCH 1975 basic syn HOBBS ERE R 1978 the effects of feral sheep grazing on ecological relationships in pinyon juniper woodlands the northern population of pinus suricatamuricatamutimuncatamurimurfcata on santa pages 41 53 in the pinyon juniper ecosystem a cruz island california pages 158 172 in DMD M power symposium utah state university logan editor the california islands proceedings of a mul- WELSH SLS L NDN D ATWOOD S GOODRICH AND LCL C HIG- tidisciplinaryti symposium santa barbara museum of GINS EDITORS 1987 A utah flora great basin natu- natural history santa barbara CA ralist memoirs no 9 KING JEJ E AND TR VAN DEVENDER 1977 pollen analysis YOUNG JAJ A RER E ECKERT AND RAR A EVANS 1978 histori- of fossil pankratpackrat biddensmiddens from the sonoran desert cal perspectives regarding the sagebrush ecosystem quaternary research 81918 igiigl191 204 pages 1 13 in the sagebrush ecosystem a symposium MEAD JIJ I1 AND D MELTZER 1984 north american qua- utah state university logan ternary extinctextinctionsions and the radiocarbon record pages YORKS TP NEN E WEST AND KMK M CAPELS 1994 changes 440 450 in PS martin and R klein editors quater- in pinyon juniper woodlands in western utah s pine nary extinctextinctionsions university ofarizonaofarizona press tucson valley between 1933 1989 journal of range manage- MEAD JIJ I1 SES E SHARPE AND LDL D AGENBROAD 1991 ment 4735947 359 364 holocene bison from arches national park southwestern utah great basin naturalist 5133651 336 342 received 8 october 1996 MMUELLERUELLERUELLERDOMDOMDOMBOISboisbols D AND H ELLENBERG 1974 aims accepted 29 may 1997 and methods of vegetation ecology john wiley and sons new york great basin naturalist 574 0 1997 appp 327 337 MULE DEER AND PRONGHORN USE OF wastewater PONDS IN A COLD DESERT

karen L cierninskil2ciemmski12 and lester D flakelalakelflake1

ABSTRACT pronghorn antilocapra ameriametiamertamericanacandcany and mule deer odocoileus hemionus were counted at wastewater ponds at the idaho national engineering and environmental laboratory INEEL in southeastern idaho 4 to 8 times per month from august 1989 through july 1991 mule deer used wastewater ponds n 15 from june through december and were most commonly observed august through november pronghorn frequented wastewater ponds from may through november and were most common from july through september the driest and warmest months ponds were also used heavily in november 1990 diel activity was studied from july through october mule deer use of ponds varied in relation to 8 diel time periods in august P 0020.020 02 and september P ooi0010.010 01 while pronghorn use varied by time period P 0010 01 in all 4 months mule deer were more active at ponds during nocturnal than diurnal counts from july through september P ooi0010.010 01 pronghorn diurnal activity exceeded nocturnal activity P oolooi0010.010 01 august through october mule deer and pronghorn use of ponds was not related to distance from site facilities groups of buildings used for research and other purposes pronghorn made greater use of individual ponds lacking additional nearby watering sites and both pronghorn and mule deer were attracted to ponds with grassforbgrass forb and shrub cover around the upland periphery

key words pronghorn mule deer desert idaho diel activity ponds wastewater

during warmer and drier periods of the year the INEEL studies thus far have shown that mule deer odocoileus hemionus and prong- radiation does not pose a hazard to animals that horn antilocapra americana in desert habi- directly use the ponds halford et al 1982 tats may seek free water beale and smith millard et al 1990 or to secondary consumers 1970 yoakum 1978 hervert and krausman markham and autenrieth 1976 arthur and 1986 diel temperature patterns during warm markham 1982 and dry conditions may influence watering and artificial systems such as INEEL wastewater activity patterns of mule deer eberhardt et al ponds are becoming increasingly common in 1984 hervert and krausman 1986 and poten- north america and many other regions of the tially pronghorn deblinger and alldredge world the degree to which such ponds are 1991 mule deer and pronghorn are found used by large mammals and their potential sympatricsympatricallyally on the idaho national engineer- influences either negative or positive have ing and environmental laboratory INEEL been largely overlooked our objectives were a department of energy research area located to evaluate wastewater ponds to 1 determine in the upper snake river plain in this semiarid monthly and diel patterns of pond use by mule environment ephemeral water sources are often deer and pronghorn and 2 determine char- available in the spring permanent watering acteristicsacteristics associated with use or nonuse of sites on the 231600 ha INEEL however are ponds by mule deer and pronghorn limited to a few game watering cisterns and sanitary industrial and radioactive wastewater STUDY AREA ponds concerns about mammal use of wastewater the INEEL is located in southeastern idaho ponds stem from possible effects to the mam- and has an average elevation of 1485 m fig mals halford and markham 1978 kuzo et al 1 temperatures range from 44c to 39c 1978 and possible transport of contaminants with july normally the warmest month average to hunters reynolds and rose 1978 hoskin- daily temperatures range from 11clic to 21c son and tester 1980 if game mammals leave average annual precipitation is 21 cm typically

ldepartmentidepartment of wildlife and fisheries sciences south dakota state university box 2140b brookingsBrookmgs SD 57007 send reprint requests to LDL D flake at this address present address national park service 13025 riley s lock road Poopoolesvillelesville MD 20837

327 328 GREAT BASIN naturalist volume 577

bitterootbitterrootBitteroot idaho range

birch creek

meelWEEL

rang technica leahilemhi support facility uttleattle lost river containment test A facility lost riverriven navalnavai range reactors facil Y N M argonne national test reactor west area idaho chemical laboratory MTRUA processing plantpiant ANIW ICPP to idaho fallsfalis

big lost river key 0 facilities rivers and sinks roads 0 4 8 12 V i M i 1471.47 i i 00y kilometers

fig 1 location of facilities at the idaho national engineering and environmental laboratory INEEL where waste watelwater ponds weiewelewere surveyed for mule deer and pronghorn august 1989 july 1991 precipitation levels are highest in may and anderson et al 1996 the primary native june and lowest in october relative humidity grasses include thick spiked wheatwheatgrassgrass ely- is commonly at its yearly minimum daily mus lanceolalanceolatustus bottlebrushbottlebrush squirreltail E average approximately 30 in july and maxi-maxi elymoelymoidesides indian ricricegrassegrass oryzopsis hymen mum 69 from december through february oides needleneedie anand thread grass stipa cocomacomatdcomatacamatacomandmatatd clawsonetalclawsonglawson etetalal 1989 and nevada bluegrass poa secunda anderson most plant communities on the INEEL are et al 1996 dominated by desert shrubs particularly wyo- wastewater ponds at which mule deer and ming big sagebrush artemisia dentatatntritridentatatridentate wyo pronghorn were studied contained sanitary minmingensisgensis anderson et al 1996 other com- waste industrial waste radioactive waste or a mon shrubs are green rabbitbrushrabbitbrush chryso- combination of waste types ponds were located thamnus viscidiflorusviscidiflorus gray rabbitbrushrabbitbrush C on the periphery of groups of buildings site nauseosus winterwinterfatwmterfatfat Krasheninnikkrashemnmkoviakrasheninnikoviaodiaovlaovia lanata facilities being used for research maintenance and spiny hopsagehousage craylagrayiagrabia spinosaspinosd assem- operations management and other purposes blages dominated by shadshadscalescale atriplex con fig 1 we eliminated 2 INEEL ponds from fertifertifoliafolia nuttall saltbush atnplexatriplex falcata analysis because they were surrounded by a 3 and winterwinterfatwmterfatfat similar to salt desert shrub com- m high chain link fence that excluded prong- munimunitiesties may occur on old lakebed sediment horn and mule deer fifteen ponds were readily 199719971 PRONGHORN AND MULE DEER USE OF PONDS 329 accessible to mule deer and 13 to pronghorn due to shortened daylight hours and de- most ponds were rectangular and ranged in creased activity around the frozen ponds we size from 0020.02 to 2212.21 ha one additional waste- reduced monthly surveys to 3 diurnal sunrise water source was a waste ditch that we included midday and sunset and 2 nocturnal dusk and only in the monthly use and diel activity stud- dawn counts per month in november and ies shorelines were gravel and subsoil plas december in january and february 1990 tic lined or cobble over plastic lined only 3 counts were conducted as in november and ponds contained emergent vegetation and I1 december except only I1 nocturnal count either other had abundant submerged vegetation dusk or dawn was conducted ponds were not ponds were partially to completely ice covered surveyed in january and february 1991 due to from late november through early march lack of target species observations during ponds are described in further detail by these months in 1990 diurnal and nocturnal cieminski 1993 counts were rotated from november through birch creek and little lost river at one february to assure sampling of all ponds dur- time terminated in playasolayas on and at the edge ing all time periods used in those months of the INEEL respectively fig 1 most flow we attempted to initiate monthly diurnal is now diverted for upstream irrigation of crops and nocturnal counts about 30 d after initia- big lost river flows onto the INEEL only in tion of counts in the preceding month ponds years following heavy snowfall the last of which were scheduled to be counted only once per prior to our study was 1987 the ponds may day because observer presence at the ponds in small part compensate for wetland habitat could influence later pond counts monthly lost since surface water no longer flows regu- counts were conducted on consecutive days larly onto the INEEL unless interrupted by inclement weather monthly and diel use analysis METHODS observations were used as an indication of counts of mule deer and pronghorn pond use for example 2 prongpronghornhom seen once we conducted monthly diurnal and nocturnal or I1 pronghorn seen twice would both equal counts on each pond august 1989 through july 2 observations for monthly use for the entire 1991 the period from august 1989 through july year we made data comparable between months 1990 was defined as year I1 when year was by summing the number of observations in a used in analyses august 1990 through july 1991 month for year I1 and year 2 at all ponds com- was defined as year 2 mule deer and prong- bined and dividing by the number of counts horn were considered to be at the pond site if conducted during that month for both years they were within 100 ra of a pond we used a combined for example the total number of spotlight for nocturnal counts which lasted pronghorn observed during diurnal periods at about 15 min and were not conducted during ponds in july of year I1 plus year 2 would be inclement weather divided by the following denominator 5 count five diurnal sunrise through sunset and 3 periods per pond X number of ponds surveyed nocturnal dusk through dawn time periods X 2 yr monthly data were then presented as the each 2 h long were established during which average number of prongpronghornhom or mule deer ob- counts were conducted from july through served per diurnal or nocturnal count october time periods were sunrise centered for analysis ofofrielofdieldiel use patterns we summed 050.5os hb after sunrise midmorningmid morning midday mid- the number of target species observations in afternoon sunset centered 050.5os h before sun- each year month and time period 8 diel time set dusk centered 151.5isls h after sunset midnight periods over all study ponds using log linear centered around 2400 military time and dawn analysis under categorical data modeling pro- centered 151.5 h before sunrise diurnal and cedures CATMOD SAS institute inc 1989a nocturnal counts from march through october we examined differences P 0050.05oos in diel use were conducted in a manner that assured sam- patterns diel activity was analyzed for differ- pling of all ponds in all time periods we ences due to diel time period and month for attempted to visit a pond no more than once the period from july through october for year per 24 h period 1 and year 2 in a second analysis we compared 330 GREAT BASIN naturalist volume 57 diurnal use of ponds with nocturnal use includ- ular pond and its characteristics could fall into ing month and time diurnal or nocturnal as a different category each month depending explanatory variables in the logistic regression on target species observation data we assumed in this analysis we also used the period from that the target species were selecting ponds july through october the sum of observations based on pond characteristics and location in over the 5 diurnal time periods was compared relation to facilities and other ponds logistic with an adjusted sum of observations for the 3 regression SAS institute inc 1989b was used nocturnal time periods nocturnal counts were to identify possible pond characteristics asso- adjusted to make them comparable with diurnal ciated with use of ponds by target species counts as follows adjusted sum mean obser- logistic regression models were developed vations per nocturnal count x 5 with a stepwise procedure at an alpha level of oos0.05 pond characteristics 005

the number of ponds or ditches within a I1 RESULTS AND discussion km circle of a surveyed pond from the pond use center was obtained from maps and aerial pho- monthly tos as was the distance to site facilities shore- mule deer first appeared at study ponds in line distance meters of shorelinepondshoreline pond was june after a january through may absence fig determined by superimposing sketches of water 2 in the intermountain region june is the surface area and shoreline interface on blue- beginning of mule deer fawning season which prints of ponds or drawings made from direct runs through mid july robinette and olsen measurements shoreline distance was remeacremea 1944 juveniles were first seen at the ponds in sured whenever fluctuating water levels ap- july but were not commonly seen until mid peared to influence this measurement during august pond use by juveniles increased through summer 1991 we determined percent cover of the summer and fall to a peak diurnal plus shrubs and grassesforbsgrasses forbs combined around nocturnal in november observations of adults the ponds by running six 20 m line intercepts steadily increased through the summer and early evenly spaced and perpendicular to the shore- fall then remained constant through novem- line at each pond plant coverage around ponds ber mule deer were not reported by age in appeared to have changed little from 1989 december because we had increasing diffi- through 1991 the method used was that de- culty distinguishing between adults and juve- scribed by canfield 1941 except only I1 inter- niles under survey conditions cept line placed I1 m off the ground was used many mule deer are year round residents for both shrubs and grassesforbsgrasses forbs because of on the INEEL but apparently are not depen- restricted access we visually estimated vegeta- dent on ponds in the spring swank 1958 and tion coverage inside the fences around radio- hervert and krausman 1986 reported move- active ponds n 2 percentage of shoreline ments to water by mule deer in arizona asso- from the water to I1 m onto the shore lacking ciated with increased temperatures these vegetation or with vegetation 15 cm tall was movements may be associated with changing estimated and defined as bare shoreline the needs for water as metabolic rates increase vertical distance from the water surface to the and at higher ambient temperatures 38c top of the surrounding berm was defined as as evaporative cooling increases hervert and pond relief we obtained weather data from a krausman 1986 during these hot dry periods national oceanic and atmospheric adminis- mule deer nonnallynormally remain in their home range tration weather station located in the south even if regular excursions are necessary to central portion of the INEEL seek water hervert and krausman 1986 boro- mule deer and pronghorn occurrences at ski and mossman 1996 in our study the peak ponds were compared to pond characteristics in mule deer observations in september lagged from july through november each month 2 months behind the temperature peak in july ponds were grouped by target species into july follows the 2 wettest months of the year those with no pronghorn or mule deer obser- on the INEEL july through november is the vations and those where these target species driest series of months clawson et al 1989 were observed months were then combined during our study precipitation in july was for the analysis thus in the analysis a barticpartic lower than in any other month INEEL 199711997 PRONGHORN AND MULE DEER USE OF PONDS 331

mule deer adult

count periods 55 0.303 01 03 F 1 01 diurnal 01 01 M nocturnal KX a diurnalanddiurna nocturnal KX 01 020.2 land 5zaz KX 0 rxarx KX rxarx 10 01 arx KX 01 rx KX 01 01oi0.1 KX 01 01 0 jan feb mar aarapr may jun juljui aug sep oct nov decdoc nulemule deer juvenile N 030.3

020.2 k 0 1 I1 c0 0 faf7 F F jan feb mar aarapr may jun juijul aug sep oct nov decdoc C 0 mule deer allaliail ages 060.6og

05 050.5 10 KX 0 10 040.4 KX 030.3 020.2 ii 01oi0.1 0 jan feb mar aarapr may jun jul aug sep oct nov decdocdeodoo month

fig 2 monthly use ofwastewaterof wastewater ponds in southeastern idaho by mule deer monthly use was calculated as the mean number of mule deer observations seen per visit to a pond count august 1989 july 1991 dotted line represents mean observations per visit for all 8 diel time periods national oceanic and atmospheric adminis- ber 1989 excluding november adult prong- tration unpublished data this precipitation horn observations peaked in july through sep- pattern and availability of forbs around pond tember and then declined in october fig 3 edges may influence pond visitation patterns juvenile pronghorn were first seen at study by mule deer on the INEEL ponds in june use by juvenile pronghorn then beale and smith 1970 reported that highest remained low through october with the same water consumption by pronghorn in utah var- november peak due to large herds in 1990 as ied from july to september depending upon in adults forage succulence moisture content which during colder months resident pronghorn was dependent on precipitation pronghorn on or near the INEEL are joined by pronghorn pond visitation in our study fig 3 peaked in moving to lower altitudes hoskinson and november however the november peak in tester 1980 with the exception of november adult and juvenile numbers was caused by from herds observed in 1990 pronghorn use several observations of large herds in novem- of INEEL ponds during 1989 and 1990 gener- ber 1990 no pronghorn were seen in novem ally declined after august or september lack 332 GREAT BASIN naturalist volume 57

pronghorn adult count periods 6 M diurnal 46 D og 22lalialesi nocturnal 060.6 diurnal and nocturnal 050.5 040.4 030.3

0 2.2

01oi0.1 0 jan feb mar aarapr may jun jul aug sep oct nov dec pronghorn juvenile &0 3c5ca 030.3

020.2 ssigligilc11

01oi0.1 i KX 0 1 1 pri I1 0 n01 r CT nfri Fn 10 jan feb mar aarapr mamay juni jul aug sep oct nov dec

C pronghorn all ages 0 2292.292929.29 L 1371.3737 0 060.6og 050.5 040.4 0.303oa 03 DO 020.2 01oi0.1 0 RM jan feb mar aarapr may jun jul aug sep oct nov dec month

fig 3 monthly use of wastewater ponds in southeastern idaho by pronghorn monthly use was calculated as the mean number of pronghorn observations seen per visit to a pond count august 1989 july 1991 dotted line represents mean observations per visit for all 8 diel time periods of heat stress and availability of snow may attracted to free water during summer but did have reduced or eliminated the need for drink- not move from an area when the water source ing water during the cooler months we can- was no longer available deblinger and all- not explain the later seasonal peak in pond ob- dredge 1991 these authors note that moisture servationsservations of mule deer compared with prong- content of vegetation was high throughout the horn summers of their study and that these results although ponds were iceieelee free by mid may not apply to an unusually hot dry sum- march we observed no pronghorn at ponds mer until may may and during june pronghorn use dependence on INEEL ponds was also low diel because air temperatures were low plant observations from the 8 diel time periods moisture content was high beale and smith were used in analysis of diel pond use by tar- 1970 and temporary rainram pools were plenti- get species from july through october of year ful pronghorn in wyoming s red desert were 1 and year 2 figs 4 5 both time period and 199719971 PRONGHORN AND MULE DEER USE OF PONDS 333 month influenced P oolooi0010.01 numbers of mule on numbers of pronghorn observed at ponds deer observed at ponds fig 4 there was a in all 4 months they were observed at ponds time period X month interaction P ooi0010.01 in all time periods except dawn fig 5 when months were analyzed individually pronghorn used ponds at different rates P time period had a significant influence on ooiool0010.01 during the diurnal and nocturnal por- mule deer observations only in august P tions of the diel cycle there was also a month 0020.02 and september P ooi0010.01 effect P ooi0010.01 and a time X month interaction log linear analysis indicated that mule P ooi0010.01 when months were analyzed sepa- deer use of ponds was greater P ooi0010.01 dur- rately pronghorn use of ponds was greater P ing the nocturnal dusk to dawn than diurnal ooi0010.01 during daylight hours from august sunrise to sunset portion of the diel cycle through october change in use due to time fig 4 the analysis also indicated that months diurnal and nocturnal was not statistically P ooi0010.01 influenced our counts and that significant P 0050.05oos in july greater diurnal there was a month X time nocturnal and diur- use of ponds from august through october at nal interaction P ooi0010.01 when we exam- the INEEL is similar to observations in wyo- ined months individually mule deer were ming by amstrup 1978 nocturnal activity is more likely P 0050.05oos to be at ponds during consistent with observations that daytime activ- nocturnal periods in all months except octo- ities are also engaged in at night albeit generally ber in october of year 1 we observed mule at a lower frequency buechner 1950 kitchen deer only during daylight hours fig 4 1974 amstrup 1978 due to the small sample size we did not amstrup 1978 observed crepuscular daily separate antlered mule deer from adult females peaks of pronghorn activity july through the percentages of antlered adults were simi- november reynolds 1984 observed similar lar between diurnal 16716716.7 antlered and noc- patterns in summer on the INEEL taylor turnal 17317317.3 antlered surveys from july 1972 recorded a midday peak june through through october august in wyoming in pronghorn activity in among black tailed deer 0 h columbianuscolumbianus addition to crepuscular peaks peaks in activ- miller 1970 found differences in diurnal activity from september through november were ity due to time in all months morning and at 0600 0900 and 1300 h taylor 1972 our ob- twilight peaks occurred june through septem- servationsservations related to occurrence of pronghorn ber early morning and midday peaks in octo- at ponds and not to general increases in activ- ber and november and high use midmorningmid morning ity still pronghorn at ponds were actively feed- till twilight in december we observed con- ing and watering and would have been rated siderablesiderable midday or midafternoonmid afternoon occurrence as active by other authors pronghorn use of of mule deer at ponds in september and octo- wastewater ponds was irregular for diel time ber but no strong sunrise or sunset peaks in periods from july through october in our study activity during the diurnal cycle fig 4 inter- fig 5 there was no strong trend of increased esestingly within the nocturnal period deer were use of ponds in crepuscular hours as active at ponds during midnight as during use in relation to dusk and dawn and krausman 1986 hervert pond characteristics suggested that desert mule deer does during the warmer drier months in arizona may have fifteen ponds were accessible to mule deer remained less active during the diurnal period on the INEEL and 13 to pronghorn the 2 to avoid water loss and to conserve energy in ponds used by mule deer but not by pronghorn general diurnal activity in midsummer in our lacked an open gate and were surrounded by idaho study was greater than that observed by mesh wire plus barbed wire on the top that hervert and krausman 1986 probably a result mule deer readily jumped in addition to waste- of milder daytime temperatures water ponds mule deer and pronghorn fre- numbers of pronghorn observed at ponds quently used a wastewater ditch that extended during the 8 diel time periods were related to for over 1000 m across the shrub desert dur- time period P ooi0010.01 and month P ooiool0010.01 ing the period from july through november there was a time period x month interaction P mule deer were observed at 12 of 15 available ooiool0010.01 analysis by individual months indi- study ponds pronghorn werewcreobservedobserved at 10 cated that time period had an effect P 0010.01 of 13 ponds accessible to them 334 GREAT BASIN naturalist volume 57

july 25

year I1 aug 1989 july 1990 20 0 5nan YOWyomyem 2 aug 1990 july iggi1991 15

0 august 25 C 0 20

10 aA 0 september 25 c1ca 20

10 0 5 L 0 october E 20

z 15

E 0 2mam 2 cn 2 time period

fig 4 diel use stacked bar graph nonoverlapping patterns of wastewater ponds in southeastern idaho by mule deerdoerdeen diedlediel use was calculated as the sum of mule deer seen at all ponds each pond counted once during each time period within a month and year

differences in habitat variables between we suspect that greater amounts of vegeta- observation categories mule deer observed or tion in the upland periphery next to the pond not observed were found only for percent bare percent shrubs was nearly significant attracted shoreline and percent grassforbgrass forb cover within mule deer to ponds either because of forage 20 in of the pond log linear analysis table 1 value or increased concealment ponds sur- buildings and other physical facilities had no rounded by bare soil or subsoil and gravel on apparent influence on mule deer use of ponds the uplands looked much less natural and were logistic regression using these 2 variables apparently less attractive to mule deer identified percent grassforbgrass forb within 20 in of for pronghorn log linear analysis using sin- the ponds as the only significant variable sepa- gle variables indicated that all habitat variables rating observation categories P ooi0010.01 con- other than distance to facilities were signifi- cordant pairs 619gig61961.9 indicated that this is not cantly different P 0050.05 between pond obser- a particularly strong logistic regression model vation categories table 1 when these variables 199719971 PRONGHORN AND MULE DEER USE OF PONDS 335

july 40

year i aug 30 year 2 aug

0 aimnim enr august 40 C 2.2

10 0 C september 0 40 IM 30 C 0 20

10 0 0 october E Z 30 20

ELMzum IWO

M C 0 2C

co CD time period

fig 5 diel use stacked bar graph nonoverlapping patterns of wastewater ponds in southeastern idaho by pronghorn diel use was calculated as the sum of pronghorn seen at all ponds each pond counted once during each time periodwithinperiodpernod within a monthandyearmonth and year

were evaluated as a group in logistic regres- serving pronghorn both percent shrub cover sion only number of ponds within I1 kmkin entered and percent grassshrubgrass shrub cover were associated the equation FP ooi0010.01 as with mule deer with increased use of ponds by pronghorn concordant pairs 63063063.0 indicated that the we commonly observed pronghorn drink- model separating observation categories was ing water from some usually larger ponds at not strong smaller ponds pronghorn were flushed by increased isolation of ponds fewer ponds observers arrival and we rarely saw them actu- within I1 km was apparently associated with ally drinking pronghorn were also observed concentration ofpronghornofpronghom use at a single pond drinking from sources such as parking lot run- other variables such as shoreline distance and off catchmentscatchments guzzlepuzzlersguzzlersrs road construction percent shrub cover could also be used to de- ponds and leaks in piping to construction or velop a logistic regression model larger ponds maintenance work areas these temporary water were associated with increased chances of ob sources quickly discovered by pronghorn were 336 GREAT BASIN naturalist volume 57

TABLE 1 wastewater pond characteristics median mean and association logistic regression by single variables a1 with occurrence1occurrenceoccurrenceaoccurrences of target speciesspecie 1 at ponds from july through october

pond observation shoreline bare pond ponds distance to cover cover category distance shoreline relief 1 km facilities grass shrubs m m m forbs MULE DEERDFER not observed 195 100 181.8 3 82 51si5.1 141.4 nn100loo100 29112911291.1 815 lgig191.9 28 937 12112112.1 75757.5 observed 392 80 17itlt1.7 3 52 17817.8 919.1gi n 30 3619 64 19 31313.1 72672 6 23323323.3 ilslis11511.5 Ppchisqchiehi sqaq 007 ooi0010.01 099 0230.23 0140.14 ooi0010.01 0070.07 PRONGHORN not observed 1562156.2 100 15 3 59 3 0 84 n 2489 852 18 30303.0 8002 13313313.3 3943943.94 observed 434 945 181.8lsis 1 82 18 999.9gg n 26 3634363 4 69 22222.2 1961961.96 91691691.6 218 ilg11911.9 P chi sqaq ooi0010 01 0030.03 001 ooiool0010.01 0420.42 0030.030 03 ooi0010 01 each month on thetiietile from july through INEEL domhom october wastewater pondponds accessible to mule deer and prongpronghornhornhoin were grouped into those with target species and those without eight months ailaugustAli gultgust octoberOctoleimielmial 1989 july october 1990 july 1991 were combined foiforhblhorbor the entire study for analysis ip is dicenteicentent bare shoreline is the percentage ofaof a pond shoreline where vegetation isis absentisabsent 01or 15 cm inin height within I1 m oftheodtheof the land water interface pond reliefisicliefisreliefis thetiietile distance froinnom the watelwaterwatey to the top ofoftlieoatliethe surrounding bembegbenn ponds 1 pond within I km include any ponds or ditches including the observation pond that aieare available and have water for target species use

probably selected over ponds if they were nearer mediate pronghorn use vegetation around the pronghorn s center of activity however these ponds included willow leaved dock R none of these aforementioned water sources salicifoliussalicifolius at all 5 well vegetated ponds pros- were permanent and some lasted little more trate knotweed at I1 of the highest use ponds than a day and poverty sumpsumpweedweed at another of the high- beale and smith 1970 observed that prong- est use ponds horn did not drink available water when mois- mule deer and pronghorn readily use waste- ture content of forbs was 75 when mois- water ponds at the INEEL perhaps as much ture content in forage plants was insufficient for the surrounding succulent vegetation as the pronghorn regularly drank water reynolds drinking water the presence of drinking water 1984 found open water was within only I1 of may be important to both species during the the 5 home ranges of pronghorn bands studied warmest and driest months of the year partic- at the INEEL pronghorn probably drink water ularly during unusually hot and dry years if available and otherwise depend on moisture where ponds are determined to be safe for from vegetation consumed einarson 1948 wildlife use designs that include grass forb it was not unusual to see pronghorn feed- and shrub cover around wetlands would likely ing in the vicinity of a pond for a few minutes improve use by mule deer and pronghorn after drinking probably an attraction to herba- ceous vegetation around some of the INEEL acknowledgments ponds several authors have found that pronghorn use was greater in areas of higher soil we thank OD markham TD reynolds moisture good and crawford 1978 or that and RG mitchell for assistance in developing pronghorn selection of forage was influenced our project and for manuscript review thanks by succulence beale and smith 1970 forbs to JA jenks for critique of the manuscript and common at some study ponds that can be to PDED evenson and JA jenks for suggestions important in pronghorn diets were prostrate on design and statistical analysis we acknowl- knotweed polygonum aviculaavicularenavicularere dock rumex edge PE saffel and S alienallenailen for assistance in col- sppapp and poverty sumpsumpweedweed iva axillarisaxillaris lecting and recording field data our research ferrel and leach 1950 bruns 1977 good and was funded by the idaho operations office crawford 1978 of the 5 INEEL ponds with US department of energy and is a contribu- well vegetated shorelines 3 had the highest tion of the environmental science and research pronghorn use of all ponds and I1 had inter foundation idaho falls idaho 199711997 PRONGHORN AND MULE DEER USE OF PONDS 337

literature CITED tive waste disposal area journal of wildlife management 4690546 905 914 ANDERSON JEJ E KXK T RUPPEL JMJ M GLENNON KEK E HOLTE HERVERT JJJ J AND PR KRAUSMAN 1986 desert mule deer AND RCR C ROPE 1996 plant communities ethnoecol use of water developments in arizona journal of agyogy and flora of the INEL ESRF 005 environmen- wildlife management 5067050 670 676 tal science and research foundation idaho falls HOSKINSON RLR L AND JRJ R TESTER 1980 migration behav- ID lilililllppIII111 appp lorior of pronghorn in southeastern idaho journal of AMSTRUP SCS C 1978 activities and habitat use patterns of wildlife management 4413244 132 144 prongpronghornshorns on montana wyoming coal lands pro- KITCHEN DWD W 1974 social behavior and ecology of the ceedingsce of the biennial antelope states workshop pronghorn wildlife monographs 38 82708 270 304 kuhkumkuzo GBG B L FRALEY JR AND 0ODD MARKHAM 1978 in ARTHUR WJ AND OD0 D MARKHAM 1982 radloRadioradionuclideradionuchdenuclide transuranic concentrations in selected biotic and export and elimination by coyotes at two radioactive abiotic components of the idaho national engineer- pages 15 waste disposal areas in southeastern idaho health ing laboratory test reactor area ponds 27 physics 4349343 493 500 inOT 0ODD markham editor ecological studies on the BEALE DMD M AND ADA D SMITH 1970 forage use water idaho national engineering laboratory site 1978 consumption and productivity of pronghorn ante- proglprogressess report US department of energy technical lope in western utah journal of wildlife manage- report IDO 12087 ment 3457034 sto570 582 MARKHAM 0ODD AND RER E autenrieth 1976 radionsradionu clides BOROSKI BBB B AND ASA S MOSSMAN 1996 distribution of in pronghorn antelope tissues on and near the pages mule deer in relation to water sources in northern INEL site 3 12 in 0ODD markham editor california journal of wildlife management 60 idaho national engineering laboratory site radio 770 776 ecology ecology programs USU S department of energy BRUNS EHE H 1977 winter behavior of pronghorn in rela- technical report IDO 12080 tion to habitat journal of wildlife management MILLARD JBJ B FWEW WHICKER AND OD0 D MARKHAM 1990 4156041 560 571 radionuclideRadio nuclide uptake and growth of barn swallows BUECHNER HKH K 1950 life history ecology and range use nesting by radioactive leaching ponds health physics 58 of the pronghorn antelope in trans pecos texas 58429429 439 american midland naturalist 4325743 257 354 MILLER FLEL 1970 distribution patterns of black tailed CANFIELD RHR H 1941 application of the line interception deer odocoileus hemionus columbianuscolumbianus in relation method in sampling range vegetation journal of to environment journal of mammalogy 5124851 248 260 forestry 3938839 388 394 REYNOLDS TDT D 1984 daily summer movements activity CIEMINSKI KLK L 1993 wildlife use ofwastewaterof wastewater ponds at patterns and home range of pronghorn northwest the idaho national engineering laboratory unpub- science 5830058 300 311 lished master s thesis south dakota state univer- REYNOLDS TD AND FL ROSE 1978 pronghorn antelope sity brookingsBrookm gs 311 appp use of the INEL national environmental research park pages CLAWSON KLK L GEG E START AND NRN R RICKS 1989 cli- 219 223 inodenodin OD markham editor eco- matographymatography of the idaho national engineering lab- logical studies on the idaho national engineering oratory and2nd edition USU S department of commerce laboratory site 1978 progress leportreport US depart- national oceanic and atmospheric administration ment of energy technical report IDO 12087 idaho falls ID DOE ID 1211815512118 155 appp ROBINETTENETTE WL AND OA OLSEN 1944 studies of the DEBLINGER RDR D AND AWA W ALLDREDGE 1991 influence productivity of mule deer in central utah transac- of of free water on pronghorn distribution in a sage tions the north american wildlife and natural brushsteppebrush steppe grassland wildlife society bulletin 19 resources conference 91569 156 161 321 326 SAS INSTITUTE INC 1989a SASSTAT user s guide version 6 1 carygary EBERHARDT LEL E EEE E HANSON AND LLL L CADWELL 1984 ath4th edition volume cars NC 889 appp movement and activity pattern of mule deer in the 1989b SASSTAT user s guide version 6 ath4th edi- sagebrush steppe region journal of mammalogy tion volume 2 carygary NC 1686 appp 6540465 404 409 SWANK WG 1958 the mule deer in arizona chaparralchapanalanai game EINARSON ASA S 1948 the pronghorn and its management arizona and fish department wildlife bul- wildlife management institute washington DC letin 3 109 appp 235 appp TAYLOR E 1972 food habits and feeding behavior of pronghorn FERREL CMC M AND HRH R LEACH 1950 food habits of the antelope in the red desert of wyoming pronghorn antelope of california california fish proceedings of the biennial antelope states work- and game 362136 21 26 shop 52115 211 219 YOAKUM D 1978 pages 103 121 GOOD JRJ R AND JAJ A CRAWFORD 1978 factors influencing JDJ pronghorn in JLJ L pronghorn use of playasolayas in south central oregon schmidt and DLD L gilbert editors big game of north proceedings of the biennial antelope states work- america ecology and management wildlife manage- shop 8182 204 ment institute and stackpole books Harnsharrisburgburg PA HALFORD DK AND OD MARKHAM 1978 radiation dosimetry of small mammals inhabiting a liquid received 21 august 1996 radioactive waste disposal area ecology 59104759 1047 1054 accepted 23 july 1997 HALFORD DKD K OD0 D MARKHAM AND RLR L DICKSON 1982 radiation doses to waterfowl using a liquid radioac cleatcreatgleatgreat basin natmahst574naturalist 574 C 1997 appp 338 342

observations ON THE reproduction SOURCES OF MORTALITY AND DIET OF THE kendallwarmKENDALL WARM SPRINGS DACE

andrew D gryskalgryska1 and wayne A huberti

al3srractalisails 1 hachaoRAC 1 the life history of the endangereded kendall warm springs dace rhinichthysrhimchthys esculusosculus therthermalismalis is largely unknown ouioulour study of its reproduction sources of mortality and diet indicated that kendall warm springs dace seem to be reproductively active throughout the yeaiyear but the rate of reproduction appears to decrease during the winter males become sexually matuiematulemature at 34 mm total length and females at 40 mm total length we observed 2 sources of mortality 1 emigration from the warm spring stream over a waterfall into the green river and 2 predation on larvae by dragonfly libellula saturatesaturatasaturata nymphs stomachs of kendall warm springs dace contained small 151 5 mm total length benthic invertebrates primarilyprimal ilyliy dipterdipteransans and mollusks

key words kendall warm springs dace rhinichthysrhimchthys esculusosculus therthermalismalis reproduction mortality diet

the kendall warm springs dace rhinichthysbhimchthys through a braided network of channels among esculusosculus thermalstherthermahsthennalisthedthentheonalisnailsmahs is endemic to a single spring large vegetation mats largely chara sppapp creek because of its restricted habitat it has been listed as an endangered species and a METHODS recovery plan has been developed that requires a more thorough description of its life history during june july and august 1995 and USU S fish and wildlife service 1982 hubbs january may june and july 1996 we captured and khune 1937 and bilms 1978 made lim- fish in traps in kendall warm springs creek ited life history observations of the fish but traps were constructed of 3 mm square mesh most aspects of its life history are poorly hardware cloth measuring 555.5ss X 11 X 31 cm understood including reproduction sources of with a fusiform shape when viewed from the mortality and diet side the traps had a V shaped mouth with a vertical slot entrance traps were set in the STUDY AREA main channel secondary channels and vegetated mats over the length of the spring creek kendall warm springs creek originates from during day and night see gryska 1996 for a several thermal springs 295c29 5cac in a small detailed description of traps and sampling pro- limestone bluff 50 km north northwest of pine totocolscols gender and spawning condition were dale wyoming hubbs and khune 1937 binns determined from external morphology and by 1978 breckenridge and hinckley 1978 the manually extruding semen or ova sexually creek has a discharge of 202.090go2 0 252.595gs2 5 msm3sns and mature males were also identified by the pres- courses for 340 ra before cascading over a 4 ence of minute nuptial tuberclestubtubercledercles on the fore- m high waterfall into the green river this head dorsal surface and fins and a longer waterfall prevents green river fish from enter- than immature males whitish anal papilla ing the spring creek the creek has a mean we did not observe the purple coloration re- depth of 10610.610log 6 cm mean wetted width of 868.68 6 m ported in spawning males described by hubbs mean water velocity of 25 cmsemsams at the bottom and khune 1937 females were identified by and a substrate composed mostly of a mix the absence of nuptial tuberclestubtubercledercles a larger than of fine oi01010.10 1 323.23 2 cm and coarse 33333.33 3 767.67 6 cm immature females clear anal papilla females limestone pebbles gryska 1996 water flows were considered to be in spawning condition

lWyoningwyoming coopnatiroopuative fish and wildlife reseaithResreveaithrehyeh unit university of wyoming laramie WY 82071316682071 3166 the unit is jointly supportedsuppoisuppon ted by the university olof01 wyoming wyoming caniecanleganiegame and fish department USU S geological stsueysteysurveyey and wildlife management institute

338 199719971 KENDALL WARM SPRINGS DACE 339 if they had a distended abdomen and eggs could portion of total stream width 0078007800.078078 sampled be extruded by gentle pressure to the abdomen with the 2 nets gender and total length TL mm of captured trapping mortalitiesmortali ties and fish caught in drift fish judged to be in spawning condition were nets immediately upstream from the cascade recorded were preserved in 70 ethanol and their observations of the presence of larval fish stomach contents analyzed the abdominal 6 15 mm TQTL were made june through august cavity of fish 20 cm TL was injected with a 1994 and january and may through july 1995 70 ethanol solution to preserve stomach con- during each sampling month we made visual tents in the lab we removed the stomach of observations with the aid of a powerful light each specimen and pooled the contents into 4 between 2200 and 2400 h during at least a 5 d classes 1 fish 40 mm TL captured during period by walking along each bank of the january 1995 2 fish 40 mm TL captured stream we searched shoreline habitat that during january 1995 3 fish 40 mm TL cap- was 1 5 cm deep without measurable current tured may june or july 1995 and 4 fish 40 and vegetated by chara sppapp mm TL captured may june or july 1995 this we observed a dragonfly libellula saturatesaturatasaturata enabled a comparison of food items between nymph in association with larval fish in shallow winter and summer and between 2 length 5 cm deep slow moving near shore habi- classes but it did not allow for assessment of tat to assess the density of dragonfly nymphs variation among individual fish invertebrates and larval fish we randomly selected 10 sam- in each identifiable taxon were enumerated by pling sites 202.0 m long and 060.6og m away from the the bureau of land management aquatic bank where larval fish were observed thirty ecosystem laboratory logan utah quadratsquadquadransrats 0020020.02 x 0020.02 m were identified within each site each site was sampled 5 nights RESULTS 7 14 june 1995 between 2200 and 2400 h using a powerful light we counted dragonfly A total of 22942 juvenile and adult fish nymphs and larval fish and observed the pred- were sampled with traps during the study atory behavior of dragonfly nymphs the light during each sampling month january may did not appear to affect the behavior of either june july and august several fish were dragonfly nymphs or larval fish dragonfly examined for the presence of semen or ova nymphs and larval fish were counted in 3 ran- measurements of the lengths of individual fish domly selected quadratsquadquadransrats within each site with and without semen or ova could not be the number of fish emigrating downstream made because of the potential mortality asso- out of kendall warm springs creek was esti- ciated with handling the minimum length at mated using two 04 m diameter 500 micron which semen or ova could be extruded differed mesh openings drift nets set at 2 points at the by gender for males the minimum length at upstream edge of the cascade due to greater which semen could be extruded was 34 mm channel slope and current velocity 1 ms at TL whereas the shortest length at which ripe this location fish collected there were being eggs could be extruded from females was 40 carried over the waterfall when captured nets mm TL semen and ova could be extruded were set horizontal to each other on a transect from males and females during all sampling across the creek so that each net sampled a months unique portion of the flow nets were set daily larval fish were also observed during all for 24 h periods between 22 july and 22 august sampling months mean densities in june 1994 5 and 10 january 1995 and 25 may and were 2822.88 larvaelarvae0larvaeoo40 04 m2ma quadrat and larval 23 july 1995 each drift net occupied 393.9 of fish were observed in 67 of the sampled n the total stream width 10410410.4 m at the falls 150 quadratsquadquadransrats estimates of densities of lar- traps were set and retrieved between 1000 val fish were not made using the sampling and 1100 h fish were measured to the nearest protocol outside of july 1995 however visual millimeter TL and identified as larvae 16 observations along the shoreline indicated that mm TQTL or post larvae 16 mm TQTL total densities were similar from may through number of fish emigrating daily over the falls august but substantially lower in january was calculated by dividing the mean number A total of 453 fish 329 larvae were cap- of fish captured per 24 h net set by the pro tured in drift nets at the upstream edge of the 340 GREAT BASIN naturalist volume 57

waterfall from may through august average gryska 1996 captured significantly fewer juve- capture peipelper 24 h sampling period was 3963.963 96 sts nile and adult fish in traps during winter than 0350.350 35 fish including 2902.902 90 sifsafs5f 0340.340 34 larvae during summer and mean length of fish cap- in each of the 2 nets we estimated that 100 tured in january was significantly greater than 95 CIC I1 85 120 fish emigrated from the in summer we submit 2 potential reasons for stream each day may august 75 of which the decline 1 an overall reduction in primary were larval fish catch rates were substantially productivity due to shorter days and reduced lower in january averaging 0810.810 81 fish includ- intensity of sunlight and 2 cooler water tem- ing 0090.090 09 larvae in each net per 24 h sampling peraturesperatures in shallow near shore larval fish habi- period tat during the winter gryska 1996 measured dragonfly nymphs were 10 25 mm long and water temperatures as low as 10c in near larval fish were 16 mm TL estimated den- shore habitats in january 1995 kaya 1991 sity of dragonfly nymphs in larval fish habitat was unable to stimulate spawning by R 0 was 22522.522925 5 nymphsm2 safs3fs 2252.259252 25 n 138 therthennalisthermalisthennalisnallsnailsmalismallsmails in laboratory aquaria by varying pho- dragonfly nymphs were frequently observed toperiodto water temperature or current our either grasping or attempting to grasp larval observations suggest that photoperiod andor fish but no records of capture rates were water temperature influence reproductive rates obtained minimum length at sexual maturity was 34 many fish collected in traps had died and mm TL for males and 40 mm TL for females were in a state of decomposition thus render- john 1963 found female speckled dace in ing them unsuitable for stomach content arizona to mature at age 2 and 45 mm standard analysis some fish had empty stomachs two length SQSL kendall warm springs dace of 45 fish 40 mm TL sampled in january had food mm SL were 54 mm TL they may mature at a items in their stomachs and only 3 physellaphydellaPhy sella 4 smaller size compared to other speckled dace Hydrahidracannahydracarinacarina and I1 chironomidae were found subspecies our longest captured kendall warm table 1 two fish 40 mm TL sampled in springs dace was 63 mm TL see gryska 1996 january contained chironomidae optioservus for length frequency data phydellaphysellaPhysella and hyallelaHy allela table 1 during sum- kendall warm springs dace regularly drifted mer 32 fish 40 mm TL had orthocladnnaeorthocladiinae out of the creek into the green river during and planorbidae as the most common taxa all sampling months january may july and table 1 and 38 fish 40 mm TL hadbad pre- august and 75 of the drifting fish were lar- dominantly riffle beetles elmidaeElmelmidgeidae chirono- vae because larval fish are relatively poor midae PhphysellaphyseliaphydeliaPhyponpoyyseliasellashila planorbidae hyallelaHy allela and swimmers they are easily displaced down- simulium in their stomachs table 1 prey of stream by the current larger fish found in the kendall warm springs dace were all 151.5lsis1 5 mm nets were often partly decomposed and we in length suspect they may have been dead upon entrance we probably underestimated the mean discussion number of fish captured per 24 h period because larval fish were difficult to see and were our observations of kendall warm springs often torn apart either by water flowing through dace and the work by bmnsbirmsbarms 1978 indicate the nets or by dragonfly nymphs captured in that breeding occurs year round males and the nets females were observed in spawning condition we estimate that 75 larval fish per day drifted and larval fish were observed during each from the creek into the green river from may sampling month january may june july and through august a total of about 9200 fish august indicating that the kendall warm average larval fish densities were 282.8 lar springs dace is reproductively active through- vae004 m2ma quadrat and larval fish habitat out the year occupied 17 329 m2ma of the creek gryska we found evidence that reproduction de- 1996 given the density of larval fish and the creases during winter very few larval fish were area of larval fish habitat we estimate the pop- seen along the shoreline and the number of ulation to be about 24000 larval fish in june drifting larvae was substantially less in january but the actual population was probably higher 0090090.090 09 larvae24 h net set than in may through because it is unlikely that all larval fish were august 8902902.902 90 larvae24 h net set additionally observed in each sampled quadrat given these 199719971 KENDALL WARM SPRINGS DACE 341

TABLE 1 numbers of organisms found in the stomachs of 2 length classes mm of kendall warm springs dace collected during winter january and summer may july 1995 number offish stomachs pooled to form the sample is in parentheses

winterwnlteriter summersumimerrmer 40 40 40 40 taxon life stage 2 2 38 32

amphipoda hyallelaHy allela adult 6 3 hydracannahydracarinaHydracarinacarlnaearlnacanna adult 4 1 elmidaeElmelmidgeidae optioservus larval 22 chironomidae pupal 23 1 6 orthocladiinae larval 1 1 87 simuliidae simulium larval 3 stratiomyidaestratiomyiidae larval 2 hemiptera adult I1 odonata libellulidae larval I1 trichoptera pupal 1 hydropsychidaeHydropsychidae cheumatopsyche larval 1 hydroptilidaehydrophilidae larval I1 physidae physellaPhyphydellasella adult 19 3 4 1 planorbidae adult 1 4 9

estimates it appears that the drift of larval fish phytic organisms consuming small lsis 151.5 mm from the stream may represent a substantial total length benthic invertebrates particularly segment of the larval fish population how- dipterdipteransans riffle beetles elmidaeElmelmidgeidae mollusks ever because reproduction appears to occur amphipodsamphipoda hyallelaHyallela and water mites hydra throughout the year our estimates of larval carina this is similar to other speckled dace fish abundance should be considered minimal that suck and scrape invertebrates from the estimates substrate by using a subterminal mouth spe- we estimate that 25 juvenile and adult fish cifically adapted for benthic foraging baltz et per day drifted from the creek into the green al 1982 van eimeren 1988 although feed- river from may through august a total of about ing behavior has not been recorded for speck- 3000 fish we have no estimates of density of led dace other species of rhinichthys have juvenile and adult fish but a total of 22942 been observed feeding between and under fish were captured with 867 trap sets during rocks and on the upper and downstream sur- summer 1994 traps were set at least 4 m apart faces ofofrocksrocks gibbons and gee 1972 diets of over the length of kendall warm springs creek other speckled dace subspecies have included so only a portion of the population in the creek diptera ephemeroptera gastropoda coleop- was sampled at one time it seems unlikely tera water mites and algae baker 1967 li and that drift of juveniles and adults from the moyle 1976 van eimeren 1988 it appears creek into the green river causes a substan- that kendall warm springs dace feed on items tial loss from the population also utilized by other rhinichthys species dragonfly nymphs preying on larvae and gibbons and gee 1972 reed and moulton small juveniles 20 mm TQTL contribute to 1973 brazo et al 1978 sample sizes were mortality among kendall warm springs dace small because no fish could be sacrificed only but the extent of the mortality is unknown in incidental mortalitiesmortalities in traps and fish cap- addition to dragonfly nymphs a number of tured in drift nets at the waterfall into the other potential predators were observed along green river were available for stomach analy- kendall warm springs creek during our study sis diet data we obtained are limited because dippers cinclus mexicanamexicansmexicana brewer s black- stomach contents were pooled within seasons birds eupharuseuphagusEuphagus cyanocephalus great blue and length classes 40 and 40 mm TQTL and herons ardea herodias and wandering garter substantial digestion and deterioration of stom- snakes thamnophis elegansdelegans vavagransmagransgrans ach contents were common because samples kendall warm springs dace diet had not largely comprised fish that had died in traps been reported previously our stomach analy- or were captured in drift 12 24 h prior to sis indicates they feed on epibenthic and epi being collected and preserved additionally it 342 GREAT BASIN naturalist volume 57

is not known where the fish captured in drift breckenridge RMR M AND BSB S HINCKLEY 1978 thermal nets resided previously in the creek conse- springs of wyoming wyoming geological survey bulletin 60 quentquentlyly fish stomach contents that we analyzed GIBBONS JRHJ R H AND JHJ H GEE 1972 ecological segrega- may not fully represent the diet of kendall tion between longnoselongnose and blacknoseblacknose dace genus warm springs dace rhinichthysrhimchthys in the mink river manitoba journal of the fisheries researchResealchseaich board of canada 29 acknowledgments 1245 1252 GRYSKA ADA D 1996 development of population monitoring protocols and description of several life history we thank R beiswenger M jennings and aspects of kendall warmwaiwal m springs dace rhinichthysrhimchthys K nelson for assistance with the study K esculusosculus thermalstherthermalisthermahstheomahsmabsmalismallsmails unpublished master s thesis uni- krueger T marwitz and C white for help in versity of wyoming laramie the field and M vinson for identification of HUBBS CLC L AND ERE R KHUNE 1937 A new fish of the genus apocope from a wyoming warm spring occa- aquatic macromaeromacroinvertebratesmacromvertebratesinvertebrates funding was pro- sional papers museum of zoology university of vided by the US fish and wildlife service with michigan 3431343 1 21 additional support by the US forest service JOHN KRK R 1963 the effect of torrential rains on the reproductive cycle of rhinichthysrhimchthys esculusosculus in the chiricahuachiracahuachiracabuaChiracahua mountains arizona copela 19632861963 286 291 CITED literature KAYA CMC M 1991 laboratory spawning and rearing of speckled dace progressive fish culturist 5325953 259 260 BAKER PH 1967 size and rel- distribution size composition HW AND PRPB MOYLE 1976 feeding ecology of the ative abundance of the lahontan speckled dace li pit sculpin coituscottusgottus ritensispipitensistensis in ash creek california rhinichthysrhimchthys esculusosculus robustrobrobustnsustusns rutter in lake tahoe bulletin of the southern california academy of sci- california game fish and 5316553 165 173 ences 7511175 lii111 118 BAUZ DM PB MOYLE AND KNIGHT 1982 com- baltz NJ REED RJR J AND JCJ C MOULTON 1973 age and growth of petipetitivetive interactions between benthic stream fishes blacknoseblacknose dace rhinichthys atraatratulustulus and lonlongnosegnose riffle sculpin cottus gulosusgulosus and speckled dace dace R cataractcataractalcataractaeae in massachusetts american rhinichthysrhimchthys osculusesculus canadian journal of fisheries midland naturalist 9020690 206 210 and aquatic 39 1502 1511 sciences 391502 USU S FISH AND WILDLIFE SERVICE 1982 kendall warm BINNS N A 1978 warm NA habitat structure of kendall springs dace recovery plan united states fish and springs with the reference to endangered kendall wildlife service region 6 denver CO warm dace springs rhinichthysrhimchthys esculusosculus thettherthennalisthermalisthennalisnallsnailsmalismaltsmaits VAN EIMEREN PA 1988 comparative food habits of gila wyoming game and fish department fisheries trout and speckled dace in a southwestern headwa- technical bulletin 4 ter stream unpublished master s thesis new mex- BRAZO D C CRC R LISTON AND R C ANDERSON 1978 DC RC life ico state university las cruces history of the longnoselon gnose dace rhinichthysrhimchthys cataractcataractalcataractaeae in the surge of in zone eastern lake michigan near received 3 january 1997 ludington michigan of the transactions american accepted 14 july 199199779977 fisheries society 107550lot107 550 556 great basin naturalist 574 0 1997 appp 343 347 TWO NEW SPECIES OF chloroperlidaeCHLORO PERLIDAE plecoptera FROM california RWR W BaubaumannlbaumanniBaumbaumannnmanniannlanni and RLR L Bottorff2

ABSTRACT Suwallia sierra and sweltsa pisteripinteri are described as new species in the family chloroperlidaeChloroperlidae from california USA illustrations of the male terminalia of both species and the female and egg ofsofjofsS sierra are presented in addition detailed figures of the epiepiproctproct of 2 similar species sweltsa tornesitownesi and sweltsa resimabesima are included diagnoses are provided comparing the new taxa with related species observations indicate that S sierra may belong to the same clade as the genus neaviperla

key words stonefliesstoneflies plecoptera suwalliaSuwallia sweltsaSweltsa chloroperlidaeChloroperlidae california

the stonstoneflyefly fauna of california was first star shaped plate basal plate with small median treated as an entity by jewett 1960 who in- dome and 15 25 stout setae on anterior 13 cluded 14 species in the family chloroperlidaeChloro perlidae anterior margin of basal plate uneven and and gave limited distributional data and a key turned dorsally fig 3 aedeagus tubular ter- stark et al 1986 listed 23 species for the state minatingminating in 2 apical lobes and 2 lateral lobes then surdick 1995 named 2 additional species subapical frontal surface with 2 curved sclero- from california to bring the total to 25 chloro tized patches laterally separated by 3 scaled perperlissperlidslids lobes proximal surface inflated and covered we discovered 2 additional undescribed with fine spinspinulesspindlesules and scales figs 1 2 cerci species of chloroperlidaeChloroperlidae from california and curved in basal segments 1 4 segment I1 elon- decided to name them so they could be added gated length 252.5 4 times width the posterior to the growing knowledge of california stone 14 13 slightly demarked from the anterior flies portion but lacking the free articulation of a walliasuwalliaSu sierra new species separate segment segments 2 4 length width and posterior portion of segment I1 with figs 1 5 13 stout spines on interior surface remaining seg- MALE macropterous body length 5 7 ments slender straight and lacking stout spines mm forewing length 6 7 mm general body fig 3 color medium brown dorsum of headbead with FEMALE macropterous body length 6 8 dark brown interinterocellarocellar area and Y line mouth mm forewing length 6 8 mm general body parts lightly sclerotized Pronopronotumturn with margins color and external morphology similar to male and rugosities dark brown disks light angles subgenital plate wide at base narrowing to rounded meso and metanotalmetanotametanota with dark U median truncate swollen lobe that protrudes at marks wings light with dark veins abdomen least 12 over sternum 9 plate originates near with dark brown median stripe on dorsum of middle of sternum 8 cercalcereal segment I1 less segments 1 8 dark brown lateral marks on seg- elongated length 2 times width than on male ments 1 2 tergum 9 with dense setal patches segments 1 4 not curved and lacking stout posterior margin slightly excavated medially spines on interior surface figs 4 5 not projecting posteriorly or heavily sclerotized NYMPH body length 6 8 mm general Hemihemitergaltergal lobes very short pointed not atten- body color golden brown lacinia with main uated into medially directed digitate processes terminal tooth and small subapical tooth dou- fig 3 epiproctEpiproct tip a small membraneous knob ble row of marginal setae 20 24 equal spaced densely covered with fine curved setae basal setae on dorsal margin 12 stout 4 medium 8 anchor bar and paraparagenitalgenital plates fused to form fine 16 18 setae on ventral margin 10 stout

1 departmentidepartment of zoology and monte L bean life science museum brigham young university prosoprovo UT 84602 2boxaboxox 10504 south lake tahoe CA 96158

343 344 GREAT BASIN naturalist volume 57

7-7

figs 1 5 Susawalhasuwalhasuwalliawallia sierra 1 male terminalia and aedeagus dorsal 2 male terminaliatermmaliamallamaila and aedeagus lateral 3 male teimterminaliamaha dorsaldorsai 4 female termtermmaliaterminaliamaliamallamaila ventral 5 female subgenital plate lateral figs 6 8 sweltsa pistenpisteripinteripisterlsteri 6 male epiepiproctproct dorsal 7 male epiproctcpiproctepiproct lateral 8 male terminaliatermmaliamallamaila dorsal figs 9 10 sweltsa tornesitownesi 9 male epiepiproctproct dor sal 10 male epiepiproctproct lateral figs 11 12 sweltsa resimabesima 11 male epiepiproctproct dorsal 12 male epiepiproctproct lateral 199719971 NEW california chloroperlidaeCHLORO PERLIDAE 345

california nevada county sweetland creek hwybwy 49 323.2 km south of north san juan 18 mayay 9831983 RW baumann RC mower and ME whiting holotype and allotype deposited in the US national museum washington DC Paraparatypestypes were examined from the following 0 localities amador co big indian creek 6 km north of plymouth llII IV 1986 RL bottorff I1 female 25125 IV1V 1986 I1 male 1 female 29129 IV1V 1986 1 female 9 V 1986 I1 male little indian rbtret creek 3 km west of plymouth 18 IV 1986 RL bottorff 3 males 5 females I1 nymph 9 V 1986 2 females I1 V 1987 1 female 12 V 1987 2 males 3 females el dorado co indian creek 333.3 km northeast of michigan bar bridge 8 IV 1987 RL bottorff 15 males 10 females f 1 nymph 13 IV 1987 5 males 2 females unnamed north bank tributary to cosumnes x river 292.9 km upstream of michigan bar bridge 9 IV 1986 RL bottorff 16 males 13 females 14 nymphs unnamed creek tributary to north bank of north cosumnes river 6 km north of nashville I V 1987 RL bottorff I1 male 3 13 females I1 nymph 12 V 1987 15 males 19 females 25 nymphs mariposa co maxwell V fig 13 Suwallia sierra egg lateral view 20ox200x cali- river near coultervilleCoulterville 25 1974 DG den- fornia sacramento co small south bank tributary to ning 2 females USNM sacramento co cosumnes river upstream of michigan bar bridge 29 IV cosumnes river at michigan bar 616 IV1V 1983 1996 RL bottorff RL bottorff I1 female unnamed south bank tributary to cosumnes river 030.3 km upstream of the michigan bar bridge 21111198621 lilliiiliIII111 1986 RL near subapical tooth 6 fine near base prono- bottorff 25 males 16 females 2 nymphs tum with variable length setae around entire 29111198629 lilliiIII111 1986 17 males 21 females 3 nymphs margin angles rounded mesonotum margined llII IV 1986 25 males 20 females 5 nymphs laterally and anteriorly with stiff setae abdomen 29 IV 1986 24 males 23 females 25111198725 illlilliiIII111 1987 concolorous dorsum covered with fine setae 14 nymphs Paraparatypestypes are in the collections of tationsetationfetationse less dense ventrally membranes with the ML bean life science museum brigham chloride cells cerealcercalgereal segments with I1 long dor- young university provo utah and RL bot- sal and 1 ventral seta posteriorly setal length torff south lake tahoe california subequal to segment length ETYMOLOGY this species is named for the EGG length 500 gmim width 250 gmumu m sierra nevada of california general shape oval cross section circular color biological NOTES Suwallia sierra light golden brown collar and eclosion line emerged from late march to mid may and absent chorion covered with irregularly hex- occurred in low elevation small often inter- agonal follicle cell impressions with walls of mittent streams with bolshecapnia maculata varying thickness and occasionally broken fol- jewett cosumnoperla hypocrenahypo crena szczytko licle cell impression floors with 2 6 central and bottorff isoperla acalaacula jewett isoperla raised areas fig 13 aduncaaldunca jewett isoperla diwokmiwok bottorff and distribution this species is known only szczytko oemopteryx vanduzeevanduzeeaa claassen from lower elevations in the sierra nevada of and sweltsa californicacalifornica jewett nymphal gut california contents included chironomid larvae diatoms TYPES holotype male allotype female and and fine organic detritus well developed eggs 5 male and 10 female paraparatypestypes collected from were found in mature nymphs 346 GREAT BASIN naturalist volume 57

DIAGNOSIS because the epiepiproctproct tip is a S autumnsautumna occurs in high elevation perennial hairy membraneous knob the fused basal plate streams and emerges in august september star shaped the aedeagus spinuspinulatedspmulatedspiculatedlated and the sweltsa adult mandibles reduced S sierrasteffasterra is clearly pisteripipinteristeri new species within the tribe suwalliimsuwalliiniSuwal liim surdick 1985 figs 6 8 which includes the genera Su and neav wallia MALE macropterous body length 7 8 iperlasperla although S sierra shares important char- mm forewing length 8 9 mm general body acters with neaviperla such as cerci with color yellow brown dorsum of head from modified basal segments and an aedeagus with hind ocelli to front margin brown pronotum sclerotized lateral patches we have assigned with dark margin and dark lateral rugosities this species to Suwallia based on the adult with lateral disks light and corners rounded meso and metanotalmetanotametanota U marks and the shape of meso and metanotalmetanotametanota with dark U marks wings the male basal plate the most important fea- light brown and veins slightly darker abdomen ture absent in S sierra males that character- with dark brown median stripe that extends izes Suwallia is the lack of distinct medially to anterior margin of ath8th tergum ninth tergum directed digitate processes on the hemiteriahemitergahemiterga bearing small rounded sclerotized knob medi- in S sierra the hemiteriahemitergahemiterga have medially di- ally tenth tergum with large V shaped darkly rected points but these are not attenuated sclerotized area located under apex of epiproctepiproct into the typical digitate processes of Susawalhasuwalhasuwalliawallia hemiteriahemihemitergaterga sclerotized and bearing long hairs and are smaller than the digitate processes of fig 8 epiproctEpiproct large and well developed neaviperla Suwallia sierrasterra is not placed in lightly sclerotized dorsally tip with round neaviperla because of important character dif- sclerotized cap that extends ventrally dorsal ferencesferen ces including the lack of an anteriorly aspect nearly parallel sided but sometimes projecting process on the ath9th tergum of the slightly narrower at base or near apex lateral male lack of a bifurcated anterior edge on the aspect flat dorsally thin near base with mas- male basal plate less modified basal segments sive rectangular shaped apex encompassing of the male cerci dark U shaped pattern on apical 23 dorsal outline straight figs 6 7 the adult meso and metanotalmetanotametanota and lack of an FEMALE macropterous body length 858.5 mm forewing length 9 general egg collar apparently S sierrasterra is intermediate mm body color in appearance between walliasuwalliaSu and neav similar to male subgenital plate with broad iperlaforcipataiperlasperla forcipateforcipata neave base narrowly constricted posteriorly forming pointed is suwalliaSu slettasierrastettasterra males differ from all other tip the apex gently rounded but is wallia much than Suwallia by 1 the greatly reduced hemitergalhemitergal narrower in most sweltsa species processes 2 elongated and curved basal cer- distribution sweltsa pisteripinteri is presently known from the coast range in northern ealcal segments which possess stout on cali- spines fornia the interior surface 3 excavated instead of TYPES holotype male and allotype female posteriorly produced ath9th tergum and 4 lateral collected from california mendocino county sclerotized patches on the aedeagus female south caspar creek jackson state forest east Suwallia often cannot be clearly identified to of caspar 9 may 1991 RL bottorff the holo- although S females do species sierrasterra differ type male and allotype female are deposited at slightly by having a more swollen posterior the US national museum washington lobe DC on the subgenital plate and this lobe pro- Paraparatypestypes were studied from the following posteriorly than jects rather ventrally Suwallia places marin co woodacre I V 1955 SW nymphs cannot be separated at this time the hitchcock I1 male USNM lily pond alpine eggs of S sierra are distinctive by lacking a lake malaise trap 10iov V 9 VI 1970 6 males collar and having hexagonal reticulations fig 6 females CNC mendocino co same data 13 as holotype 18 IV 1985 1 male Suwallia sierra was found in the same major ETYMOLOGY this species is named for drainage basin of the central sierra nevada with edwin P pister of bishop california phil has 2 other Suwallia species however it is sepa- long been a supporter of rare species and en- rated spatially and temporally from both species dangereddangered habitats sweltsa pisteripinteri is presently S pallidulapallidula occurrs in middle elevation peren- in this category and needs to be better under- nial streams and emerges in june august and stood 199719971 NEW california chloroperlidaeCHLORO PERLIDAE 347

DIAGNOSIS the male genitalia ofofsjofsS pisteripinteri jr provided at the canadian national collec- are most similar to sweltsa tamalpa ricker tion CNC ottawa ontario canada is much sweltsa tornesitownesi ricker figs 9 10 and sweltsa appreciated OS flint jr made specimens resimabesima surdick figs 11 12 both S pisteripinteri available from the united states national and S tamalpa bear a sclerotized prong on museum USNM washington DC appreci- their ath9th tergum but it is small and delicate in ation is also expressed to the colleagues who S pisteripinteri fig 8 while it is large and bears were along when specimens were collected broadly sclerotized lateral bands in S tamalpa the excellent illustrations were done by jean ricker 1952 fig 140 the lateral outline of S tangerstanger leavitt the epiproctepiproct has a downward pointed projec- tion in S tamalpa but in S pipisteripinteristeri S resimabesima literature CITED and S tornesitownesi it is broad and flat in dorsal view the epiproctepiproct is pointed in S tamalpa enlarged JEWETT SGS G JR 1960 the stonestonefliesflies plecoptera of california bulletin of the california survey 6 apically in S resimabesima and S tow and nearly insect tornesitownesinesi 125 177 parallel S stehnstenn in pipisteripisteiipinteristeri the female subgenital plate RICKER WE 1952 systematic studies in plecoptera of S pisteripinteri is narrow at the tip as in S tamalpa indiana university publications science series 18 ricker 1952 fig 143 but not bifurcate or as 1 200 sharply pointed STARK BPB P SWS W SZCZYTKO AND RWR W BAUMANN 1986 north american stonestonefliesflies plecoptera systematics distribution and taxonomic references great basin acknowledgments naturalist 4638346 383 397 SURDICK RYR F 1985 nearctic geneiagenelagenera of chloroperhnaechloroperlinae we are grateful to BC kondratieff WD plecoptera chloroperlidaeChloroperlidae illinois biological mono- shepard BEBY stark and RFRY surdick for help- graphs 54154 1 146 1995 new western nearctic sweltsa plecoptera ing us with descriptions of these 2 interesting chloroperlidaeChloro perlidae proceedings of the entomological species KD alexander allowed us to use his society of washingtonofwasbington 9716197 161 177 SEM photograph of the sweltsa sierra egg special thanks are given to AW knight of the received 14 april 1997 university of california davis for his support accepted 9 septemberseptember19971997 and encouragement the help that DE bright great basin naturalist 574 C 1997 appp 348 351

HOMING IN EASTERN FENCE LIZARDS sceloporus UNDULATUS FOLLOWING SHORT DISTANCE translocation

eric W heinlaheinl2hem12 and shayna J Whitakerwhitakerl3whitaker1313

absrracabsrlaur r we conducted an experiment on eastern fence lizards sceloporus undulatesundulatus during august september 1995 near los alamos new mexico 1 to ascertain if lizards that were relocated short distances exhibited homing 2 to investigate a possible barrier to movement and 3 to determine the effect of transtranslocatinglocating individuals from a transplant area on lizards in a recipient area we relocated 15 of an estimated population of 39 95 CI 36 45 lizards an average distance of 46 m fourteen of 15 transtranslocatedlocated lizards returned to within 6816.816 81 stsy 1431 43 m of the original capture location movement distances did not vary F 0760.760 76153153 df P 03810.3810 381 between resident and translocatedtrans located lizards during the piepretreatmenttreatment period and did not vary for resident F 2869862 86 112 df P 011660.11660 1166 but varied between pretreatment and posttreatmentposttreatment periods for transtranslocatedlocated F 146514.6514 65 17 df P 000650.00650 0065 lizards translocatedTranslocated lizards did not affect the resighting probability of resident lizards F 096ogg0 96 114 df P 0340.340 34 but this may be related to low power 1 p 0150.150 15 and transtranslocatedlocated lizards moving out of the area quickly

key words barrier disturbance easternfenceeastern fence lizard sceloporus undulatesundulatus homing new mexico translocation

some reptile species may be relocated to and investigated the effect of transplants on mitigate habitat related conflicts or for humane resident lizards in a different area reasons dodd and seigel 1991 nevertheless the study was conducted on a 4355 m2ma area sceloporus sppapp may exhibit homing noble located in los alamos new mexico 3553 N 1934 mayhew 1963 weintraub 1970 guyer 10620106log 20 W at an elevation of 2165 in the 1978 ellis quinn and simon 1989 thus reduc- study site is divided into a south 1520 m2ma and ing the effectiveness translocationsoftranslocationsof translocations if eastern north 1900 m2ma area by a 55 x 17 m patch of fence lizards sceloporus undulatusundulates are trans dense vegetation which is bordered on the located it is unknown whether a subsequent southern portion of the north side by a 3 m increase in density in surrounding areas may wide arroyo each area is composed of moder- cause some individuals in the resident popula- ate to steep tallus slopes with a wide range of tion to be adversely affected eg see noble boulder sizes a nearly vertical canyon wall 1934 tubbs 1975 reinert 1991 gordon 1994 creates a boundary for approximately one half thick vegetation or open habitat may form of these areas the site also contains a 05 m barriers to dispersal and movements for east- wide trail running approximately south north ern fence lizards noble 1934 jones and droge which connects the 2 areas and may provide a 1980 tinkle 1982 the ability of animals to corridor for movements predominant vegeta- traverse the surrounding habitat matrix may tion in the 55 x 17 m wide interstitial area con- determine the number of animals reaching a sists of brome bromus sppapp yarrow achilla given distance from or returning to a source lanulanulosalosa apache plume fallugiaFallugia parparadoraparadoxaadoxa population however corridors may provide im- and ponderosa pine pinus ponderosa portant landscape components for dispersing we captured marked andor resighted east- animals noss 1983 inglis and underwood ern fence lizards during daily surveys that lasted 1992 approximately 151.5isls h during the mornings of this study was designed to determine if 14 17 20 25 and 28 august 1995 lizards lizards translocatedtranslocated 70 in across a 55 x 17 m were sexed measured from snout to vent SVL wide patch of vegetation would return to the and individually marked using canary yellow site of capture or remain in a different locale liquid paper8paperpapera the gillette co boston MA additionally we simulated an immigration event with a I1 x 15isem cm number on their dorsal

esileesiseslES 15l environmental science group MS j495 los alamos national laboratoryLaboi atory los alamos NM 87545 2pr3cntzpioscnt address U S fishfisli and wildlife service 2730 loker avenue west carlsbad CA 92008 resent3prcenicicsentsentt addreaddless university of new mexico department of biology albuquerque NM 87505

348 199711997 HOMING IN EASTERN FENCE LIZARDS 349 surface we assumed marking did not affect side of the canyon to within the area where we lizards noble 1934 jones and ferguson 1980 had repeatedly observed each individual or prior to initiating the experiment we con- within 10 m of the original point of capture for ducted mark resightdesight surveys and estimated a individuals that were not observed prior to daily population size of 39 95 CI 36 45 transtranslocatinglocating we also measured long distance hein and myers 1995 minimum daily move- movements for 2 lizards ID nos 2 and 18 that ment distances were determined during mark were observed twice during I1 survey resightdesight surveys conducted in the pretreatment all distances were normalized by log trans- period ie 14 28 august by measuring the formation prior to analyses we compared mean distance from the first sighting of an individ- distances moved and SVL between transplant ual to the next sighting on subsequent days and resident lizards during the pretreatment Resighting probabilities c were calculated by period using analysis of variance whereas SVL summing the number of times each resident in relation to distances moved was compared lizard was observed divided by the number of using regression PROC GLM SAS institute surveys past the initial capture and marking inc 1988 all other comparisons of distances during pretreatment cayecprecpye andd posttreatmentposttreatment that individual lizards moved were tested using apostccpostost periods we translocatedtranslocated lizards during a repeated measure analysis of variance PROC 20129 31 august 1995 and continued resighting GLM SAS institute inc 1988 we also tested lizards through 3 september 1995 we also whether transplants adversely affected resi- surveyed the study area on 19 september dents by comparing sightingreresighting probabilities 1995 for I1 h among resident lizards between pretreatment we randomly selected the south area as the and postposttreatmenttreatment periods using a repeated transplant population meaning recaptured indi- measure analysis of variance PROC GLM SAS viduals were relocated to the north area we institute inc 1988 because we used a repeated attempted to recapture all lizards transplants measures analysis of variance each lizard acted and residents and remark with liquid paperpapero as its own control and the normal between resident lizards were released at the site of experimental unit ie lizard to lizard variation recapture the north area was subdivided into from the error sum of squares was thus re- a grid of 4 equal area cells with each cell moved power 1 I1 P3 of tests was also calcu- approximately 475 m2ma we randomly selected lated for each comparison SAS institute inc I1 of the 4 cells to receive the first translocatedtranslocated 1988 lizard subsequent lizards were systematically placed in the next higher numbered cell RESULTS translocatedTranslocated lizards were placed in the center of each cell the shortest distance that lizards movement distances did not vary F 0760.76 were relocated was greater than the largest 153 df P 03810.381 1 Pp 0830.83 between the radii calculated from reported home range esti- resident and transplanted lizards during the mates 13013013.0 m turner et al 1969 15015.0 m mar- pretreatment period and did not vary between tins 1994 therefore transtranslocatedlocated lizards were periods for resident F 2862.86 112 df P believed to have been displaced outside the 011660.1166 1 Pp 0340.34 lizards but they varied normal range of their movements unmarked between periods for translocatedtranslocated F 146514.65 lizards captured on the south side were also 17 df P 000650.0065 1 P3 0910910.91 lizards addi- marked and translocatedtranslocated we measured the tionallytionally distances moved by lizards between straight line distance from each capture loca- north and south areas differed between pre- tion south to each release site north and the treatment and postposttreatmenttreatment periods F 158015.80 distance from each subsequent sightingreresighting to 119 df P 000080.0008 1 pP 1001.00loo SVL did the original point of capture until the lizard was not differ F 1891.89 1115 df P 01710.171 1 0 within 10 m of the capture location or the 0280.28 between transplant and resident lizards study ended straight line distances were used there was no relationship between SVL and to calculate Grifgriffinfirsmirss index griffin 1952 wein- distance moved F 1651.65 1634 df P 01070.107 traub 1970 which measures the directness of a I1 Pp 0790.79 between lizards during the pre- translocatedtranslocated animal s return ie homing path treatment period successful homing following translocation was thirteen of 15 7 female 8 male transtranslocatedlocated defined as moving from the north to the south lizards exhibited homing by moving to the 350 GREAT BASIN naturalist volume 57

TABLETABLL 1 summary of eastern fence lizards transtranslocatedtianslocatedlocated 75 m in los alamos new mexico during august september 1995

translocatedTranslocated homed no days Griffigriffinyss ID sex SVL mm distance m didistancestancea m to home index I1 F 77 4120 402 3 105 2 M 60 6580 1540 1 140 4 F 56 2380 1629 2 104 7 F 67 3880 940 1 104 10 F 66 5105 289 3 107 11 M 46 5970 500 21 102 15 M 56 5750 320 1 112 18 M 60 6340 185 1 139 42 F 66 4040 b 43 M 57 4935 880 2 117 44 F 62 4070 726 2 120 45 M 48 3905 1165 3 195 46 M 66 3750 169 3 109 47 M 52 5903 110 3 102 48 F 57 3120 509 1 164

distancedist niceulce fromh om the original01 iamigm il point of capturec iptui e pipriorlot101 to transtranslocatinglocating diciididdeclediddicl not demonstrateite homing but wasw is icresighted I1 time in the northern area

south side of the canyon within an average of cajcaprecprecpj g 0580.58oss s 0060.06 and posttreatmentposttreatment 6.81681 s 1.43 of original 05 681 sy 143 in the capture point aptcptpost 0490.49 sx 0050 periods for individual in 2 d sy 0250.25 table 1 translocatedTranslocated lizards resident lizards but this may be related to low moved an average of 7687.68 ssy 1471.47 and statistical power J1 p 00150.1515 because of a small 221722.17 in sys 4424.42 whereas resident lizards effect size 0090090.09 andor sample size n 15 moved an average of 6376.37 s 101.0iolo and 10010.0 rn ssy 1681.68 during pretreatment and post discussion treatment periods respectively one additional transtranslocatedlocated lizard was observed within 5 in of in our study the majority 14 of 15 of east- the original point of capture on 19 september ern fence lizards exhibited homing by return- 1995 griffingriffins s index averaged 1201.20 ssy 0070.07 ing to the south side of the canyon with most indicating that on average transtranslocatedlocated lizards J11I1 of 14 lizards returning to 10 in from the moved 121.2 times the relocated distance as they original capture location this finding agrees were returning to the capture location lizards with other studies that demonstrated homing 2 and 18 moved 43143.1 and 16416.4 in in 71 and 80 in lizards sceloporus sppapp that were translo min respectively catedbated 240 in noble 1934 150 in may- eight 4 female 4 male of 14 lizards were re- hew 1963 215 in weintraub 197051970 280 inm captured and transtranslocatedlocated whereas 7 3 female guyer 1978 and 200 inm ellis quinn and 4 male unmarked lizards were captured and simon 1989 male and female eastern fence relocated we recaptured and remarked 10 2 lizards homed equally well table 1 although female 8 male of 18 resident lizards and did we did not estimate home ranges the mini- not capture or sight any unmarked lizards in mum daily movement distances during the the resident area fifteen of 18 resident lizards pretreatment period indicate that all lizards were resighted an average of 2172.17 s 0290.29ogg were relatively sedentary however we cannot times during the experiment during the study rule out that some translocatedtranslocated individuals lizards were captured andor sightedreresighted on the may have been familiar with the northern area canyon floor tallus slopes and corridors ie and we suspect that lizards successfully homed trail and stream whereas no marked lizards because transtranslocatedlocated distances were relatively were captured or resighted and no unmarked short ie 65865.8 in no lizards were observed lizards were observed in the patch of vegeta- in the patch of dense vegetation it may have tion inhibited movements alternatively lizards sightingresightingRe probabilities did not differ F were observed on or near the small trail and 0960.96ogg 114 df P 0340.34 between pretreatment streamstreambedbed which suggests these features may 199711997 HOMING IN EASTERN FENCE LIZARDS 351

have been used as corridors between the 2 ELLIS QUINN BAB A AND CAC A SIMON 1989 homing behav- areas corridors may provide important land- lorior of the lizard sceloporus jarrovi journal of her- scape components for dispersing animals noss petpetologyology 231468314623 146 152 GORDON DRD R 1994 translocation of species into conser- 1983 inglis and underwood 1992 vation areas a key for natural resource managers we did not detect an effect on the resightdesight natural areas journal 143114 31 37 ing probabilities of resident lizards by translo GRIFFIN DRD R 1952 bird navigation biological review 2735927 359 393 catedbated lizards but our test had poor power 1 I1 GUYER C 1978 comparative ecology of the short horned 3 olsois0.15 a low 009 015 because of effect size 0090.09 lizard phrynosoma douglassidouglassi and the sagebrushsage brush and small sample size n 15 if the effect lizard sceloporus graciograciosussus in southeastern idaho size had been large eg 0450450.45 which might unpublished master s thesis idaho state university imply the biological significance of an immi- pocatello 56 appp gration event was high then the power of this HEIN EWE W AND OB0 B MYERS 1995 comparison of mark recapture and mark resightdesight methodologies for test would have strong 080 we esti- been ie 0800.80 mating the size of a small population los alamos sightedreresighted 15 of 18 residents I1 time during national laboratory LA UR 95 4355 los alamos the experiment suggesting transtranslocatedlocated lizards NM 35 appp did not cause resident lizards to emigrate INGLIS G AND AJA J UNDERWOOD 1992 comments on some designs however lizards were capable of moving large proposed for experiments on the biological importance of corridors conservation biology 6 distances in a short amount of time and the 581 586 transtranslocatedlocated lizards spent relatively little time JONES SMS M AND DLD L DROGE 1980 home range size 2 d among the residents the amount of time and spatial distributions of two sympatric lizard for translocatedtranslocated lizards to home was shorter species sceloporus undulatusundulates holbrookiaHolbrookia maculata than studies sceloporus in the sandhillssandhills of nebraska Herpetologicherpetologicaherpetologicala 36 that displaced sppapp 127 132 greater distances than our study noble 1934 JONES SMS M AND GWG W FERGUSON 1980 the effect of ellis quinn and simon 1989 but similar to a paint marking on mortality in a texas population of study with shorter 125 m displacement dis- sceloporus undulatesundulatus copela 19808501980 850 854 tances weintraub 1970 MARTINS EPE P 1994 phylogenetic perspectives on the evolution of lizard territorialityterntonality pages 117 144 in we did not detect any deleterious effects of in LJL J vitt and EXE Z pianka editors lizard ecology transtranslocatinglocating lizards on the resident lizards historical and experimental perspectives princeton however if small scale habitat disturbance university press princeton NJ 403 appp causes fence lizards to emigrate into neighbor- MAYHEW WW 1963 biology of the granite spiny lizard ing areas resident lizards in these areas may sceloporus orcuttiorcuttsorcutti american midland naturalist 6931069 310327310 327 be affected translocatingTranslocating eastern fence lizards NOBLE GKG K 1934 experimenting with the courtship of may cause residents to display aggressively or lizards natural history 34534 5 15 attack noble 1934 which may affect survival NOSS RFR F 1983 A regional landscape approach to main- and reproduction vinegar 1975 similarly tain diversity bioscience 3370033 700 706 artificial crowding may affect sceloporus REINERT HKH K 1991 translocation as a conservation strat- sppapp egy for amphibians and reptiles by some comments reducing growth andor survival rates tubbs concerns and observations Herpetologicherpetologicaherpetologicala 473573634735747 357 363 1975 consequently future studies should SAS INSTITUTE INC 1988 SASSTAT user s guide release investigate whether transtranslocatedlocated or resident 6036.036 03 version SAS institute inc carygary NC lizards are affected eg increased aggression TINKLE DWD W 1982 results of experimentalofexperimental density manip- lower by ulation in an arizona lizard community ecology 63 or survival immigration events 57 65 TUBBSTUBBSAAAA 1975 effects of artificial crowding on behavior acknowledgments growth and survival of juvenileofjuvenile spiny lizards copeiacopela 48204 820 823 this report is contribution number LAURLA UR TURNER FBEB RIR I1 JENNRICH AND JDJ D WEINTRAUB 1969 95435495 4354 from alamos national laboratory home ranges and body sizes of lizards ecology 50 los 1076 1081 we thank myers for OB providing resources VINEGAR MBM B 1975 life history phenomena in two pop- for the study and reviewing an earlier draft of ulationsulations of the lizard sceloporus undulatusundulates in south- the manuscript western new mexico american midland naturalist 9338893 388402388 402 CITED WEINTRAUB JDJ D 1970 homing in the lizard sceloporus literature orcuttiorcutts animal behavior 1813218 132 137 DODD CKC K AND RAR A SEIGEL 1991 relocation repatria- received 28 february 1997 tion and translocation of amphibians and reptiles accepted 28 april 1997 are they conservation strategies that worlwork herpeto lodicalogica 4733647 336 350 cleatcreatgleatgreat basin naturalist 574 0 1997 appp 352 354

NEW VARIETY OF astragalus conjuncturconjunctusCONJUNCTUS S WATSON FROM BENTON COUNTY washington

stanley L welshlwelsh1welshel florence caplow2caplowe and kathryn beckabeck2

ABSTRACT discussed are lelationshipsrelationships of the species within astragalus section Conjuncti A conjunctconjunctusconjuncturus var rickardrickardsrickardurickardiirickrockarduii welsh beck & caplow var nov is proposed

key words astragalus conjunctconjunctusconjuncturus Conjunctconjuncticonjunctsi washington

in 1984 a collection of plants from the han- hanford reservation US department of ford atomic energy plant taken by gary baird energy fig 1 arrived at the herbarium at BRY among the rattlesnake ridge is within an area estab- specimens were several collections of an astra- lished as the fitzner eberhardt arid lands galus obviously belonging to the section con ecology reserve a high quality native shrub junttijuncti as proposed by barneby 1964 the steppe environment section is characterized by having stistipulesstipuledpules the benton county plants were subse- connate at the lowermost nodes and greatly quently compared with materials obtained on shortened lower intermternodesinternodesnodes with the upper loan from oregon state university OSC one or few elongating the leaves thus disposed including WILU through the kindness of dr in a subbasal tuft barneby 1964 isely 1996 aaron listenliston the OSC and WILU collections commonly most of the plant height is attrib- include specimens of all species of section utable to elongated pedpedunclespeduncleduncles and racelesracemes conjuncti and demonstrate the considerable the hanford specimens have erect sessile pods range of variation within the individual species similar to those of all other members of the main variation within the species complex in- Conjunctconjuncticonjunctsi except for A leibergii in which the volves flower size pod lengthwidthlength width ratio and pods are stipitate degree of elongation of the upper interinternodesnodes however the plants from hanford have the stipe of A leibergii easily distinguishes it strigulose pods merely strigose vesture other- from all other taxa within the conjuncti and wise banner reflexed through about 45 and from such look alikes as A sheldoniisheldosheldonienii rydberg rather short cylindric calyces thus they do barneby and A reventus A gray both of which not fit exactly within any of the species outlined occur adjacent to the range of the conjunct i in the atlas of north american species of astra- phalanx these latter both relegated to section galus barneby 1964 dr barneby graciously leventireventi arrecti subsection leventireventi arrecti examined the plants and indicated their close have similar overall habit but lowermost stip relationship with A conjunctconjunctusconjuncturus S watson the ules are distinct present writers concur though the collections A single specimen from OSC lawrence 99 by baird from 1984 and more recent collections 26 june 1917 is similar to the benton county taken by kathryn beck and florence caplow materials it is from wasco county oregon 12 in 1995 represent a slight northward extension miles southeast of the dalles near rice sta- of that species into washington both the baird tion in a bunchbunchgrassgrass prairie and transition scab- and beck and caplow materials represent rel- land with the notation protected from grazing atively uniform plants taken in 2 main locali- this season it is in fruit only the pods are ties in benton county washington one south- strigulose as inn the benton county plants and west of kiona and the other north northwest bear the notations A reventus and A hoohoodianusdianus of benton city on rattlesnake ridge in the the specimen s features are mainly those of A

IMLM L bean litelife science museum and department of botany and range science brigham young university provo UT 84602 23963 squalicumsqualicuniSqualsquallSquali cunieuniicum lake road bellingham WA 98226

352 199719971 NEW VARIETY OF astragalus CONJUNCTconjunctusconjuncturUS 353 conjunctconjunctusconjuncturus sens lat and the collection is ten- tattativelyively identified as belonging to the benton county plants here regarded as var rickardrickardiiii hanford franklin should revisited rsarvaficn certainly the collection site be yakima A key to the species of Conjunctconjuncticonjunctsi differing considerably from that published by barneby richland pasco 9.9 0 P wallakg 1964 is presented below following the species benton wallawalia i i key is a description of A conjunctconjunctusconjuncturus and a key i washington 1 to the closely related taxa as herein interpreted klickitatKlicki tat

errgonorrgon i section conjuncti umatilla dallas I1 pods and ovaries stipitate the stipe at least 3 mm bnhetheshe dallesdaliesdailes ameapeAA 1 gilliam morrow long plants of kittitasKittitas chelan and douglas coun- aft sherman ties washington A leibergleibergiileibergnii MEM E jones wasco

1 pods and ovaries sessile plants of various distribution in oregon washington and adjacent idaho fig 1 portions of washington and oregon showing 2 pods evidently villous sometimes glabrous in locations of astragalus conjunctconjunctusconjuncturus var rickardii welsh age calyx 10 11 15 mm long the teeth 262 6 beck & caplow 464 6 676 7 757 5 mm long plants within and near the columbia gap wasco and hood river counties oregon and klickitatKlickitat co washington A hoohoodianushoodiamisdianus howell flowers ascending to spreading at anthesis the 45 2 pods glabrous or strigulose calyx and teeth of axis 3 4 12 15 cm long in fruit bractsbraats 2 454.5 various length plants variously distributed mm long pedicelspedicels 1 454.5 mm long bractbracteoleseoles 3 banner recurved through ca 45045 oblanceo- 2 calyx 7 858.5 12 mm long the tube st57575.7 late or broadly rhombic oblanceolate emar- 6 929.2gg mm long 252.5 4 mm wide cylindric or ginate 16 25525.525 5 mm long 676 7 10 hunnununnmm wide subcylindric strigulose pilosulous the teeth calyx teeth 131.31 3 3 4 mm long pods 5 8 mm 13 3 16 255 thick plants of benton co washington and 131.3 4 mm long subulate flowers 25525.5 transmontane oregon mostly above 610 m mm long whitish with keel tip and other petals where its range approaches the following tipped with purple the banner recurved oregon from wasco to baker co southeast through ca 45 pods erect sessile oblong ellipeglip into southwest idaho A conjunctconjunctusconjuncturus S watson sold to narrowly oblong ovoid straight or through 900goo 3 banner strongly recurved 90 slightly 12 25 mm long 5 8 mm oblong oblanceolate rhombic oblanceolate incurved elliptic oblong ovate or somewhat quad thick obcompressed glabrous or strigulose ratelybately ovate cuneate usually deeply notched subbilocular the septum to 141.4 mm wide ovules 13213213.213 2 14 20720 7 mm long 5 989.89 8 mm wide 23 30 calyx teeth 24242.42 4 272.72 7 515.15 1 mm long pods 45454.54 5 6 11 mm thick plants of klickitatKlickitat yakima 1 calyx tube cylindric to subcylindric 494 9 929 2 mm and kittitasKittitas counties crossing to the south long pods glabrous 5 8 mm thick plants wide- bank of the columbia in sherman co oregon spread var conjunctusconjuncturus A reventifonnisreventzformis rydberg barneby conjunct 1 calyx tube campanulate to subcylindric 494 9 626 2 astragalus conjunctconjunctusconjuncturus S watson proc amer mm long pods strigulose 434.34 3 5 mm thick plants acad arts 17 371 1882 basalt milkvetchmilkvetch local in benton co washington and wasco co moderate acaulescentsubacaulescentsub or shortly caules- oregon var rickardrickardiinckardiiii cent perennial 15 65 cm tall from a superfi- pubescence cial branching caudex strigulose astragalus conjunctconjunctusconjuncturus var conjunctconjunctusconjuncturus basifixed stems erect or ascending several to numerous in bushy clumps stipulesStistipuledpules 3 10 11 A reventus var conjunctconjunctusconjuncturus S watson ME mm long at least the lowermost connate jones phaca conjuncta S watson piper lium sheathing leaves 6 10 30 cm long mostly conjunctconjuncturconjunctumum watson rydberg in a subbasal cluster leaflets 9 13 25 31 Pedpedunclespeduncleduncles 10 33 cm long racelesracemes 7 to 17 3 23 mm long linear oblong elliptic lanceo- 20 flowered the axis 3 4 12 15 cm long in late or subsubfiliformfiliform obtuse acute or retuse fruit pedipedicelscels 1 4544.55 mm long calyx 7 858.5 12 the terminal one continuous with the rachis mm long the tube 57575.7 6 929.2gg mm long cylin- pubescent below and above Pedpedunclespeduncleduncles 10 33 dric or subcylindric the teeth 131.3 3 4 mm cm long racelesracemes 7 to 17 20 flowered the long flowers 16 25525.5 mm long pods 12 25 354 GREAT BASIN naturalist volume 57 mm long 5 8 mm thick glabrous ovules washington wasco co see lawrence 99 cited 23 30 type in john day valley oregon J above howell in may 1880 and on sterile rocky Bunchbunchgrassgrass sagebrush community at 450 to ridges in baker county by W C cusick 1881 1070 m on the hanford atomic energy site holotype GH isotopesisotypesisotypes ORE WS paraparatypestypes benton co washington and bunchbunchgrassgrass seabscab GH ORE land community wasco co washington meadows brushy slopes grasslands sage- relationships within the conjuncti are prob- brush desert and pine forests on basaltic bed- lemalematicaltical all taxa being closely alike the taxon rock at 485 to 1555 m from the blue moun- proposed here appears to share features of tains baker co west to the deschutesDeschutes river both A conjunctconjunctusconjuncturus and A reventiformis the and south to the malteurmalheur valley steens moun- calyx tube proportions are similar to those of tain oregon and east to owaheeowyhee co idaho the only slightly disjunct A reventifonnisreventiformis but the calyx measurements are smaller than for us rickard astragalus conjunctconjunctusconjunctur var ii that taxon furthermore proportions of the welsh beck & caplow var nov calyx tube length width ratio within A conjuncconjunct similis A conjuncti var confconjconjunctunchi in habituhabiru tus in a strict sense differ only in degree from sed in leguminibusleguminibus pubescentpubescentibusibus et angustioaugustio those of var rickardrickardiiii the presence of pubes- ribusbibus et florinusfloribusfloribus minorminoribusibus generageneralitergeneralizerliter differt cence in pods within the genus rises and falls Pedpedunclespeduncleduncles 5 26 cm long racelesracemes 10 to but is apparently uniform in the benton county 19 flowered the axis 4 13 cm long in fruit plants pod pubescence is herein considered pedipedicelscels 1 252.5 mm long calyx 7 9 mm long diagnostic and seems to be correlated with the tube 494.9 626.2 mm long campanulate the relatively shorter calyces and narrower pods teeth 151.5lsis 3 mm long flowers 13213.2 14 20720.7 some specimens assigned to A reventiformis mm long pods 13 20 mm long 434.3 5 mm from nearby yakima county caplow & beck thick strigulose ovules 15 20 95075 96003 and baird 633 have pods ab- TYPE washington benton co TIINTUN solutelysolutely and proportionately as narrow as those r26e s30 NWSW on northeast facing slopes of the proposed new variety the pods in those of rattlesnake mountain with sagebrush and specimens are however glabrous sandberg bluegrass at ca 1036 m 29 may 1995 the taxon is named in honor of dr bill kathryn beck & florence caplow 95083 rickard one of those responsible for estab- holotype BRY isotopesisotypesisotypes NY US WTU WS lishmentlishment of the fitzner eberhardt arid lands additional SPECIMENS PARATYPES wash- ecological reserve ington benton co rattlesnake hills ca I1 mi NW ofofbennettbennett ranch 18 may 1984 G baird literature CITED 794 north slope of horse heaven hills along mcbee 12 may 1984 G baird 734 ratt- barneby RCR C 1964 atlas of north american astragalus rd of the new york botanical garden 13 18 may memoirs lesnaketlesnake hills above snively basin 1 1188 1984 G baird 775 horse heaven hills near isely D 1996 astragalus L in native and naturalized chandler butte 14 april 1995 K beck & F leguminosae fabaceae of the united states unpub- caplow 95022 rattlesnake mountain 14 may lished manuscript iowa state university ames 1995 K & F caplow 95038 beck horse received 7 may 1997 heaven hills near chandler butte 22 may accepted 11 june 1997 1995 K beck & F caplow 95064 all BRY great basin naturalist 574 0 1997 appp 355 358

COYOTE ACTIVITY PATTERNS IN THE SIERRA NEVADA

john A shivikl2shivik12Shivik12 michael A jaegersjaeger3 and reginald H BarrBarrettbarrettiettletti1

key words activity patterns california caniscams latranslatranolatrans coyote sierra nevada

the winter months in high mountain regions MATERIALS AND METHODS of the sierra nevada make survival difficult for predators that exist there high productiv- we conducted our research in the 105 km2 ity during the spring and summer enables coy- watershed surrounding the university of cali- otes canis lalatranslatranotrans to survive and reproduce fornia sagehen research station approximately but little is known about how these animals 13 km north of truckee california in the tahoe overwinter in mountainous areas hawthorne national forest elevation ranges from 1880 1970 suggested that coyotes make altitudinal to 2620 m the area is characterized by long migrations but recent data gantz 1990 gese cold winters and warm dry summers with et al 1996 shivik et al 1996 indicate that nightly temperatures falling below OC often coyotes can remain in the high mountains at all times of year most of the annual precipi- throughout the year tation 91 cm falls as snow during winter temporal rhythms eg innate behavioral forested areas are dominated by jeffrey pine rhythms such as diel cycles and seasonal repro- pinus jeffrejeffreyjeddreyyii and white fir abies concolor ductive cycles may influence or help elucidate brush fields contain deerdeerbrushbrush ceanothus coyote activity patterns we hypothesized that velutinus and greenleaf manzanita arctosta- coyotes vary activity levels throughout the year phylos patula sagebrush artemisia triden as a result of these seasonal biological require- tata dominates on lower dry slopes small ments we related coyote activity patterns to stands of lodgepole pine pinus confortacontortacontorta var the seasons that are likely to influence these murmurryanaryana and aspen populus tremultremuloidesoides patterns we analyzed coyote activity in the occur near springs meadows and streams sagehen basin of the sierra nevada because red fir abies magnificamagnified mountain hemlock seasonal activity patterns in this seasonally cold tsuga mertensianamerten siana and western white pine and snowy climate provide insight into how pinus monticola dominate at higher eleva- coyotes allocate energy in order to survive and tions morrison et al 1985 reproduce we trapped coyotes using steel legholdbeghold traps previous analyses of coyote activity used with offset padded jaws and short anchor movement data to determine activity levels chains to minimize trapping injury and stress gipson and sealander 1972 andelt and gip- hawthorne 1970 olsen et al 1986 coyotes son 1979 smith et al 1981 gese et al 1989 were immobilized physically with a pin stick however some authors argued that using dis- and then vet wrap or electrical tape the sex tance traveled is not a good measure of coyote weight age gier 1968 and general condition activity knowlton et al 1968 laundre and were recorded for each captured coyote shivik keller 1981 the literature presents a paucity 1995 during the course of the study we made of analyses examining seasonal trends in coyote 18 captures of 16 coyotes during 3 trapping activity using activity sensitive collars espe- periods cially in mountainous areas therefore we con- radiocollarsRadio collars with signal pulse varying activ- ducted a study using activity sensitive radio ity switches model 400 telonics 932 E impala collars to monitor seasonal coyote activity in a ave mesa AZ 85204669985204 6699 were fitted to each mountainous region of the sierra nevada coyote these collars transmitted a 75 pulse

departmentldepartment of environmental science policy and management 151 hilgard hall university of california berkeley CA 94720 2presentpresent address department of biology colorado state university fort collins CO 80523 SUSDA national wildlife research center 151 hilgard hall university of california berkeley CA 94720

355 356 GREAT BASIN naturalist volume 57

minmm signal when the collar was moving and a used ANOVA to determine whether differences 50 pulseminpulsemmpulpuiseminsemm signal when it was stationary for in activity were apparent at different times of over I1 minmm hand testing of the collars indi- day during the biological seasons cated that the internal switches were quite sen- for all analyses the sample unit was the sitive and that a very small amount of move- individual coyote for example in the ANOVA ment was required to set the collar into active of the percent activity by season and day cate- mode therefore for purposes of this study a gory all activities of coyote f040 during each nonactive coyote was one that had not moved season and day category were reduced to a sin- for over I1 minmm and was probably sleeping gle average to avoid pseudopseudoreplicationreplication hurl- concurrent with attempts to locate each ani-ani bert 1984 if a significant difference in mean mal we monitored collared coyotes for activity activity level was detected we performed mul- during eight 4 hr radio tracking sessions per tiple comparisons using tukey tests statistical week shivik et al 1996 coyotes were moni-moni assumptions were assessed using residual plots tored once per hour in 4 hr blocks that brack- kirby 1993 eted sunrise middle of the day sunset and middle of the night each coyote was moni- RESULTS AND discussion tored for approximately 2 minmm each hour during the 4 hr tracking session for analysis data for the 12 coyotes 7 males and 5 females were divided into the following 6ghrhr categories monitored 1368 activity rates were calculated 11 morning 0400 and 1000 day 1000 for the 4 time of day categories xY 342 per and1600and 1600 evening 1600 and2200and 2200 and season x 114 per coyote from 2150 obser- 11 nightmght2200night220029002200 and f 0400 A coyote s percent vations on individual coyotes x 538 per activity during each block was the basic de- season mean activity varied between seasons pendent variable we calculated the estimate P 00010.001 but there was no evidence for a of percent activity by dividing the number of difference in activity by sex P 0630.63 and no times the animal was recorded as active by the interaction between sex and season P 0192olgg0.192 total number of times it was heard during the because we did not detect a difference in activ- block of monitoring if the number of samples ity between sexes we did not partition out the was 2 effects of sex in the remaining analyses coyote the influence of seasonal factors such as activity during the breeding season was signif- weather patterns cannot be easily separated icantly less than the pup P 00010.001 and dis- from changes due to internally controlled persal P 00110.011 seasons and activity was behavioral rhythms however to survive and less during the pre pup than the pup P reproduce coyotes must pursue certain behav- 00290.029 season fig 1 activity significantly iorslors ege g forming pair bonds maintaining ter- varied by day category only during the disper- riritories and feeding pups regardless of envi- sal season P 00330.033 when activity peaked ronmentalronmental conditions therefore we used the during the evening Tx 0600.60ogo sys 0050.05 and biological season as the basis for examining was lowest during the day x 0420.42 sxaxs coyote activity through time Laundrlaundre6 and 0040.04 keller 1981 smith et al 1981 data were our results are consistent with those of divided into the following seasons for analysis other researchers who found that coyote activ- breeding 1 I1 january 15 march pre pup 16 ity varied by time of day even when previous march 30 april pup rearing 1 I1 may 31 july studies used different methods to rate activity and dispersal 1 I1 august 31 december data and involved different degrees of pseudorepli were collected 1 august 1993 31 july 1994 cation gipson and sealander 1972 andelt and we assessed activity levels by animal within gipson 1979 shivik and crabtree 1995 fur- seasons during the regularly scheduled morn- therthermoremore other studies did not show differing evening midday and midnight tracking ences in activity between males and females sessions and arcsinearcsmearceme transformed the activity gipson and sealander 1972 andelt and gip- rate for each coyote before analysis zar 1984 son 1979 suggesting that activity rates of because seasonal activity could be influenced males and females do not differ drastically by the sex of the coyote we analyzed data using however laundrelaundralaundr6 and keller 1981 indicated a 2 way ANOVA we hypothesized differences that females travel less than males during the in activity between sex and season and also pup season leading us to hypothesize that 199711997 NOTES 357

07 breeding and pre pup seasons but because deer are absent from this study area in winter 06 T the carcass mechanism for decreased activity I1 proposed by bekoff and wells 1980 may not T r i apply to sagehen coyotes 05 the ultimate mechanism for reducing energy 1 T expenditures during winter may be that reduced 04 A winter activity probably improves chances for 0 individual survival because activities such as z 03 pup rearing are not occurring coyotes are able LU to reduce their levels of activity during winter behavioral plasticity allows coyotes to 02 this survive and reproduce in mountainous areas even when carrloncarrionearrion ege g from winterwinterkillkill anguungu 01 lates is not available thus the proper cur- rency for examining coyote ability to remain in 00nn i areas with a seasonally reduced prey base and harsh weather conditions is the amount of time ev o909oa relegated to social and reproductive behaviors seasons when behaviors SEASON in activity intensive such as pup rearing are not occurring coyotes fig 1 seasonal activity of coyotes in the sagehen basin reduce energy expenditures and exist inm areas of the sierra nevada bars represent I1 standard error with seasonally limited food supplies means were calculated using each coyote as the sample unit after estimating individual coyote activity within each acknowledgments season the breeding season was 1 january 15 march n 7 pre pup was 16 march 30 april n 7 pup was 1 may 31 july n 11 and dispersal was 1 august 31 we thank RKR K bloom MLM L klavetter BWB W december n 12 merkle DSD S pilliod V schliecher L scinto J terenzi and LAL A wood for their valuable assistance in the field this study was possible overall female activity is not reduced but that because of the support of V aretchearretche KJ jones travel by female coyotes is limited to a smaller and JLJ L kent of the truckee ranger district area during the pup season differences be- tahoe national forest provided administra- tween what the distance traveled metric and tive and logistic assistance M reynolds resireslresi- absolute activity measure actually represent dent manager of sagehen creek research sta- contribute to confusion regarding seasonal tion procured housing and other support we changes in coyote activity lanudrelanudr6Lanudre and keller thank JWJ W laundrelaundralaundr6 for his review of the manu- 1984 female coyotes that are nursing pups script this work was supported by the denver would be considered active by our method wildlife research center under cooperative and not active when using distance traveled agreement contract 12 34 74 0235 CA with data A correlationeoncon elation is evident however between the university of california berkeley the cali- distance traveled and absolute activity and fornia agricultural experiment station project therefore both distance traveled and motion 54541010 MMSS and university of california berke- sensitive radiocollarradiocollar data are useful for elaminexaminexamin- ley wildlife graduate student funds ing diel and seasonal coyote activity noting that each method is sensitive to different behaviors literature CITED coyotes in grand teton national park rest ANDELT WFWE AND PS GIPSON 1979 home range activity more during winter months because coyote reli- and daily movements of coyotes journal of wildlife ance on carrioncarrlonearrion reduces the need to hunt small management 4394443 944 951 mammal prey bekoff and wells 1980 simi- BEKOFFBEKOFE M AND MCM C WELLS 1980 social ecology and be- larly in yellowstone national park coyotes havior of coyotes scientific american 242130242 130 148 reduce activity as available carcass biomass GANTZ GRG R 1990 seasonal movement patterns of coyotes gese in the bear river mountains of utah and idaho increases et al 1996 during our study unpublished master s thesis utah state university coyotes were also less active during winter logan 67 appp 358 GREAT BASIN naturalist volume 57

GESE EM OJ RONGSTAD AND WR MYTTON 1989 1984 home range size of coyotes a critical review changes in coyote movements due to military activity journal of wildlifeofwildlife management 4812748 127 139 journal of wildlife management 5333453 334 339 MORRISONORRISON MLM L ET AL 1985 natural history of vertebratesofvertebrates GESECESE EME M RLR L RUFF AND RLR L CRABTREE 1996 forag- of sagehen creek basin nevada county california ing ecology of coyotes caniscams latranslalatranotrans the influence university of california berkeley 16 appp of extrinsic factors and a dominance hierarchy cana- OLSENLSEN GHG H SBS B LINHART RAR A HOLMES GJG J DASCH AND dian journal of zoology 7476974 769 783 CBC B MALE 1986 injuries to coyotes caught in padded GIER HTH T 1968 coyotes inm kansas kansas state college and unpadded steel foothold traps wildlife society agricultural experiment station bulletin 393 118 bulletin 1421914 219 223 PP SHIVIKIIVIK JAJ A 1995 factors influencing space use and activ- GIPSON PS AND JAJ A SEALANDER 1972 home range and ity of sagehen basin coyotes unpublished master s activity oftheodtheof the coyote caniscams latransfrustrorlatranslatrano frustrorfrustbrusttorror arkansas thesis university of california berkeley 162 appp proceedings of the annual conference of the south- SHIVIKIIVIK JAJ A AND RLR L CRABTREE 1995 coyote activity eastern association of game and fish commissions levels in relation to presence of california gulls at 268226 82 95 mono lake california california fish and game HAWTHORNE VM 1970 movements and food habits of 812281 22 28 coyotes in the sagehen creek basin and vicinity SHIVIKIIVIK JAJ A MMM M JAEGER AND RHR H BARRETT 1996 coyote unpublished master s thesis university of nevada movements in relation to the spatial distribution of reno 78 appp sheep journal of wildlife management 6042260 422 430 HURLBERT SHS H 1984 Pseudopseudoreplicationreplication and the design of SMITHkllthgjjrGJ JR CARYGARY AND OJ RONGSTAD 1981 sam- ecological field experiments ecological monographs pling strategies for radio tracking coyotes wildlife 5418754 187 211 society bulletin 9889 88 93 KIRBY KNK N 1993 advanced data analysis with SYSTAT van ZARVR JHJ H 1984 Biostatistical analysis and2nd edition prentice nostrand reinhold new york 475 appp hall new jersey 718 appp KNOWLTON FFEE PE MARTIN AND JCJ C HAUG 1968 A telemetric monitor for determining animal activity received 18 november 1996 journal of wildlife management 3294332 943 948 accepted 3 april 1997 LAUNDRLAUNDRE9 JW AND BLB L KELLER 1981 home range use by coyotes in idaho animal behaviour 2944929 449 461 great basin naturalist 574 0 1997 appp 359 362 US distribution OF THE COEUR DALENEYALENE salamander PLETHODON idahoensis SLATER AND SLIPP

albert G wilson jrjrejr11 evelyn M wilson2wilsonn craig R groves3 and richard L Wallacewallace4wallacec4

key words amphibia salamander plethodon geographic distribution rocky mountains

the coeur dalene salamander plethodon and fall working within and around the geo- idahoensis is the only lungless salamander graphic range of P idahoensis as it was known plethodontidae known from the northern previously lynch 1984 we surveyed regions rocky mountains discovered in 1939 on the accessible by road or trail in idaho and mon- idaho panhandle slater and slipp 1940 this tana up to 2000 in elevation and attempted to terrestrial amphibian has since been encoun- visit and veivelverify all previously reported localities tered in northwestern montana and southeast- we located salamanders by digging by displac- ern british columbia teberg 1964 holmberg ing surface objects and by night searches with et al 1984 flashlight descriptions of localities are listed prior to this study the status and exact loca- with the idaho natural heritage program tions of some historic P idahoensis collection boise and the montana natural heritage pro- groves sites were unknown 1988 moreover gram helena voucher specimens have been the northern rocky mountains in the US hahad deposited in CASGAS in the charles R conner nuss- not been well surveyed for amphibians museum CM washington state university baum et al 1983 raising the possibility that pullman and in herpetological collections at additional localities might exist to assist con- the university of idaho ulUI efforts directed toward this species servation A locality is here defined as an area of occur- groves 1988 we conducted surveys docu- rence 05os050.5 kmkin from another such area it menting its occurrence south of canada herein appears that at least 55 localities were recorded we report results of the study for P idahoensis before the present study we we compiled locality data from reviews in found vague and incomplete data for some his- brodie 1970 brodie and storm 1970 and collection sites and suspect they may be lynch 1984 from unpublished records of toric for which have idaho and montana wildlife agencies from synonymous with localities we personal correspondence with individuals hav- more complete information the canadian ex- ing field experience with P idahoensis and tent of this species has yet to be completely from the collection records of the following documented three british columbia locali- museums california academy of sciences san ties exist in the kootenai valley within 60 kmkin francisco CAS museum of vertebrate zool- of the US canadian border orchard 1991 ogy university of california berkeley ameri- one hundred thirty two new localities were can museum of natural history new york discovered during our field surveys these in- field museum of natural history chicago clude the following 3 range extensions 39 km museum of comparative vertebrate zoology due NNE of libby lincoln county montana harvard university cambridge on 9 may 1988 CM 8916189 161 484423n4804423n fieldwork was conducted from 1987 through liyiqiyw1151915w 792 in elev 656.5gs km due W of 1994 primarily during wet weather in spring victor ravalli county montana on 25 may

departmentidepartment of zoology washington state university pullman WA 99164423699164 4236 present address department of life sciences spokane falls community college MS 3180 3410 west fort wright drive spokane WA 99224528899224 5288 address correspondence to this author college ofveterinary medicine washington state university pullman WA 99164701099164 7010 present address peone pines veterinary clinic 4717 north newport highway mead WA 99021 sldaho31daho department of fish and game 600 south walnut street box 25 boise ID 83707 present address the nature conservancy 2060 broadway avenue suite 230 boulder CO 80302 department of biological sciences university of idaho moscow ID 83843

359 360 GREAT BASIN naturalist volume 57

1140 0 0 il MONTANA e

ileklekre r

0 0 0 50

L I1 km 4 80

A 0 0 r 0 k

joe

0 0 0 C 0 4 19 0 w a t e r e S 0

I1 if 0

fig 1 USU S distribution of the coeur dalene salamander plethodon idahoensis historic localities are represented by open circles new localities by closed circles 199719971 NOTES 361

1987 CM 8916389 163 462529n4602529n distributions of salamanders by limiting time 1550 in elev and 32 km due SEIMWof lowell for foraging and reproduction lynch 1981 idaho county idaho on 15 may 1989 UI IA not surprisingly P idahoensis occupies drain- 6271627 1 460435n4600435n ils11511516161151616nv1616 NV 780 in elev ages with the longest local annual frost free in all these sites extend the species known periods the kootenai valley which contains range 44 km NE 87 km E and 16 km S respec- the species northernmost populations has the tively teberg 1965 brodie 1970 lynch 1984 longest frost free season of any forested region we found P idahoensis in forested moun- in northwestern montana or southeastern tainous regions between 500 and 1550 in eleva- british columbia caprio 1965 ross and sav- tion most localities occur in valleys draining age 1967 farleyparley 1979 the west slope of the bitterroot range of idaho the association of P idahoensis with rocky the southernmost occur in the clearwater river micromicrohabitathabitat is consistent with reports of other drainage fig 1 the species has a scattered terrestrial amanderssalamanderssalamanderisal talus and rock outcrops distribution in the clark fork and kootenai favor occurrence of these forms by providing river drainages of montana we commonly en- seasonal shelter and oviposition sites herring- countered the salamander in talus and other ton and larsen 1985 Ramotramotmkramotniksamotniknik and scott 1988 rocky debris exposures of fractured bedrock bury et al 1991 such retreats have historically are present at almost all localities and we often insulated P idahoensis from the harsh climate observed P idahoensis using fractures as retreats and recurrent wildfireswildfires of the northern rock- all localities are affiliated with springs seep ies arno 1980 nussbaum et al 1983 As has ages or damp areas along streams been proposed for other plethodons herring- except for the existence of some marginal ton 1988 we suggest that protection of under- lowland populations in xerophytic habitats ground refugia may be the key to preserving distribution of P idahoensis is confined to tem- populations of this species in the future perate mesophytic forest types daubenmire 1978 the upper altitudinal limits of the sala- acknowledgments mander coincide with the transition between temperate mesophytic and subalpine commu- this study was supported by the USDAUS DA nities in the northern rockies arno 1979 forest service bureau of land management habeck 1987 and exceed those of most west- and natural heritage programs of idaho and ern plethodons nussbaum et al 1983 montana we thank the cited museums joel plethodon idahoensis is part of a disjunct chavez lowell diller phillip dumas james coastal biota inhabiting the portion of the rocky lynch ronald nussbaum stanley orchard mountains in which maritime climatic influ- and kenneth teberg for sharing locality data ence is strongest arno 1979 johnson 1987 other assistance was provided by daniel bivens cooper et al 1991 annual precipitation in francis cassirer patrick mullen nancy tyler regions occupied by P idahoensis averages and richard wallen manuscript reviewers 50 140 cm the highest levels occur on the included keith aubry robert herrington west slope of the bitterrootsBitterroots where the sala- lawrence jones and john larsen mander is most abundant pacific northwest river basins commission 1969 through much literature CITED of its range P idahoensis occupies drier terrain than do other western plethodons at the same ARNOAKNO SES F 1979 forest regions of montana USDA forest service research paper INT 218 intermountain latitudes dumas 1956 herrington 1985 wilson forest and range experiment station ogden UT et al 1995 this species affinity for stream 39 appp margins and seeseepagespages contributes to its survival 1980 forest fire history of the rocky mountains in comparatively andaridarld habitats wilson and journal of forestry 7846078 460 465 BRODIE EDE D JR 1970 western salamanders of the genus larsen 1987 plethodon systematics and geographic variation much precipitation in the northern rocky Herpetologic a 2646826 468 516 mountains occurs as snow finklin 1983 and BRODIE EDE D AND RMR M STORM 1970 plethodon vandykesvandykeivandy kei regions inhabited by P idahoensis have colder americanameiamel icanlean society of lebthyologistsichthyologists and herpetolo- more prolonged winters than those inhabited gists catalog of american amphibians and reptiles 91191.191gli 1 912 s by most of the salamander western convenerscongenerscongeners BURY RBR B PS CORN AND KBK B AUBRY 1991 regional pat- nussbaum et al 1983 freezing weather affects terns of terrestrial amphibian communities in oregon 362 GREAT BASIN naturalist volume 57

and washington pages 341 350 in LEL F ruggiero insecta coleoptera unpublished master s thesis KBK B aubry ABA B carey and MHM H huff editors university of idaho moscow 268 appp wildlife and vegetation of unmanaged douglas fir LYNCH JEJ E JR 1984 reproductive ecology of plethodon forests USDA forest service general technical idahoensis unpublished master s thesis university of report PNW GTR 285 pacific northwest research idaho moscow 59 appp station portland OR LYNCH JEJ F 1981 patterns of ontogenetic and geographic CAPRIO JMJ M 1965 average length of freeze free season variation in the black salamander aneidesflavipuncaleidesaneides flavipunc folder 83 montana agricultural experimental sta- tatus caudata plethodontidae smithsomansmithsonianSmithsoman cont- tion bozeman ributions to zoology 32413243941 1 53 COOPER SV KE NEIMAN AND DW ROBERTS 1991 NUSSBAUM RAR A EDE D BRODIE JR AND RMR M STORM 1983 forest habitat types of northern idaho a second amphibians and reptiles of the pacific northwest approximation USDA forest service general tech- university press of idaho moscow 332 appp nical report INT 236 intermountain forest and ORCHARD SAS A 1991 amphibian population declines in range experiment station ogden UT 143 appp british columbia pages 10 13 in CAC A bishop and daubenmire R 1978 plant geography academic press KEK E pettit editors declines in canadian amphibian new york 338 appp populations designing a national monitoring strat- DUMAS PC 1956 the ecological relations of sympatry in egy canadian wildlife service occasional paper 76 plethodon dunnidunmbunni and plethodon vehiculumvehwulum ecology canadian wildlife service ottawa ontario 3748437 484 495 PACIFIC NORTHWEST RIVER BASINS commission 1969 FARLEYPARLEY ALA L 1979 atlas of british columbia university climatological handbook columbia basin states pre- of british columbia press vancouver 136 appp cipitation volume 2 environmental science ser- FINKLIN AIA I1 1983 weather and climate of the selway bit vices administration washington DC 262 appp terrottterroot wilderness university press of idaho moscow RAMOTNIK CAC A AND NJN J SCOTT JR 1988 habitat require- 144 appp ments of new mexico s endangered amanderssalamanderisalamanderssal GROVES CRC R 1988 status and distribution of the coeur pages 54 63 in RCR C szaro KEK E severson and DRD R dalene salamander plethodon vandykeivandykesvandy kei idahoensis patton editors management of amphibians reptiles in idaho unpublished report to the idaho depart- and small mammals in north america proceedings ment of fish and game boise 39 appp of a symposium flagstaff AZ USDA forest service HABECK JRJ R 1987 present day vegetation in the north- general technical report RM 166 rocky mountain ern rocky mountains annals of the missouri botan- forest and range experiment station fort collins ical garden 7480474 804 840 CO herrington RER E 1985 the ecology reproductive biol- ROSS SHS H AND CNC N SAVAGE 1967 idaho earth selencescience ogy and management of the larch mountain sala- geology fossils climate water and soils idaho earth mander plethodon iwseuilarsellicarsellilarlafselli burns with comparisons science series 111 1 271 to two other sympatric plethodons unpublished doc- SLATER JRJ R AND JWJ W SLIPP 1940 A new species of pletho- toral dissertation washington state university pull- don from northern idaho occasional papers of the man 102 appp department of biology college of puget sound 1988 talus use by amphibians and reptiles in the 383838 38 43 pacific northwest pages 216 221 inCT RCR C szaro KEK E TEBERG EKE K 1964 an extension into montana of the severson and DRD R patton editors management of known range of plethodon vandykeivandyvandykeskei idahoensis her amphibians reptiles and small mammals in north petologica 1928719 287 america proceedings of a symposium flagstaff AZ 1965 range extensions of the salamander pletho- USDA forest service general technical report RM don vandykeivandykesvandykei idahoensis copela 19652441965 244 166 rocky mountain forest and range experiment WILSON AGA G JR AND JHJ H LARSEN JR 1987 activity and station fort collins CO diet in seepage dwelling coeur dalene salamansalsaiamandersdeisders herrington RER E AND JHJ H LARSEN 1985 current status plethodon vandykeivandykesvandykei idahoensis northwest science habitat requirements and management of the larch 6221162 211 217 mountain salamander plethodon larlarsellarsellilarselltcarsellisellilt burns bio- WILSON AGA G JR JHJ H LARSEN JR AND KRK R mcallister logical conservation 3416934 169 179 1995 distribution of van dyke s salamander HOLMBERG RGR G NPDN PD ANGERILLI AND LJL J LACASSE plethodon vandykeivandykesvandykei van denburgh american mid- 1984 overwinteringOverwintering aggregations of leiobunumLeiobunum land naturalist 134388134 388 393 paesslerpapaesspaesslenpaessleriessienesslenlerilerl in caves and mines arachnida opiliones journal of Arachnology 1219512 195 205 received 8 january 19919977 JOHNSON PJ 1987 larval taxonomy biology and biogeog- accepted 16 june 1997 raphy of the genera of north american byrrhidaeByrrhidae great basin naturalist 574 0 1997 appp 363 365

reproduction IN THE WESTERN CORAL SNAKE micruroides EURYeuryxanthuseukyxanthusXANTHUS ELAPIDAE FROM ARIZONA AND SONORA mtxicoMEXICO

stephen R goldberglgoldbergeGoldbergl

key words reproduction micruroides euryeuryxanthusxanthus western coral snake arizona maicomalcomexico

the western coral snake micruroides eury cut into sections at 5 gmwimlim slides were stained xanthus kennicott 1860 ranges from central with harris hematoxylinhaematoxylin followed by eosin arizona and southwestern new mexico to counterstain testes slides were examined to southern sinaloa mexico and occurs from sea determine the stage of the male cycle epi level to 1770 in stebbins 1985 information didymidesdidymides and vasa deferentiadeferdeferentialentia were examined on the biology of this species is in roze 1974 for sperm slides of kidney sexual segments there are only anecdotal accounts of repro- were examined for secretory activity ovary duction in M euryeuryxanthusxanthus funk 1964 behler slides were examined for the presence of yolk and king 1979 stebbins 1985 ernst 1992 deposition williamson et al 1994 rossi and rossi 1995 data on the male micruroides euryeuryxanthusxanthus degenhardt et al 1996 according to lowe testicular cycle are presented in table 1 tes- et al 1986 the M euryeuryxanthusxanthus reproductive ticular histology was similar to that reported cycle is tied to summer rains egg laying occurs by goldberg and parker 1975 for the colu- in july august with hatchhatchlingslings appearing in brid snakes Masticmasticophisophis taenitaeniatusatus and pituophis summer shaw 1971 assumed breeding melanoleucusmelanoleucus in the regressed testes seminif- occurred in the spring with egg deposition in erous tubules contained spermatogoniaspermatogonial and late spring or early summer roze 1996 re- sertolibertoli cells in recrudescence there was reported that in sonora mexico oviductal eggs newal of spermatogenic cells characterized by were found in M euryeuryxanthusxanthus in the and2nd half spermatogonial divisions primary and sec- of may and the end of july egg laying ondary spermatocytes and spermatids may have stretched from the end of july to september been present in spermiogenesis metamermetamor the purpose of this report is to provide infor- phasingphosing spermatids and mature sperm were mation on reproduction in M euryeuryxanthusxanthus present males undergoing spermiogenesis I1 report on data from 56 micruroides eury were found in all months examined april xanthus 35 males mean snout vent length november table 1 epididymides and vasa SVLSVLI 382 mm 41941.9 s range 320 493 mm deferentiadeferdeferentialentia of spermiogenic males contained 21 females mean SVL 371 mm 49749.7 s sperm the smallest spermiogenic male mea- range 315 497 mm from arizona and sonora sured 320 mm SVL only 2 males with re- mexico in the herpetology collections of the gressed testes were found I1 each in june 493 university of arizona UAZ tucson and the mm SVQSVL and august 330 mm SVL no males natural history museum of los angeles county with recrudescent testes were noted the sex- JACMLACM los angeles appendix counts were ual segment of the kidney was enlarged and made of oviductal eggs or enlarged follicles contained densely staining secretory granules 3 mm diameter the left testis epididymis in spermiogenic males mating coincides with and vas deferens and part of the kidney were hypertrophy of the kidney sexual segment saint removed from males the left ovary was re- bironsgirons 1982 moved from females for histological examina- the smallest reproductively active female tion tissues were embedded in paraffin and enlarging eggs measured 356 mm SVL to

department of biology whittier college whittier CA 90608

363 364 GREAT BASIN naturalist volume 57

tahleTABLETAHLL 1 monthly distribution of conditionsofconditions in seasonal the time frame for egg laying suggested by testicular cycle of micruroides euryeuryxanthusxanthus values shown lowe et al 1986 for arizona july august are the numbers of males with testes exhibiting each of aieale and roze 1996 for sonora the 2 conditions none weiewere in recrudescence mexico end of july to september quinn 1979 reported that a month N regressedreglRegi essed speimiogenesisspermiogenesis few females of the texas coral snake micrurus april 1 0 1 fulviusfulvous tener laid eggs in may but most were Mmayay 3 0 3 deposited in june september newborn M eury june 3 1 2 xanthus measured 190 200 mm total length july 2 0 2 15 al august 11 1 10 and weighed 151.5 g lowe et 1986 september 9 0 9 spermiogenesis occurred in all months in october 5 0 5 which testes of micruroides euryeuryxanthusxanthus were novemberNovembeibel 1 0 1 examined april november this may suggest that spermiogenesis is continuous and that males are capable of breeding throughout the year this would support and 1995 avoid the possibility of using immature females rossi rossi who stated that mating may occur the fall as I1 included only micruroidesmicrurotdes euryeuryxanthusxanthus 4of in well as in spring quinn 1979 similarly re- this size or larger in my analysis of the female ported an extended reproductive cycle one female from 31 mamayy period of spermiogenesis all months except may and 356 mm SVL contained 3 enlarging eggs 4 5 june july in micrurus tener reproductive cycle mm diameter one female 440 mm SVL from micrurusfulviusfulviusfulvous the of M euryeuryxanthusxanthus appears to fit into 23 june contained 6 oviductal eggs 6 7 mm saint girons 1982 category of diameter which may represent the largest birons spermatogenesis mixed type A in which there is mating in fall published clutch size for this species two in size and spring with a long period of spermiogenesis females I1 from 30 may 497 mm SVL and 1 from 13 july 357 mm SVL contained folli- acknowledgments cles in early vitellovitellogenesisgenesis yolk granules pre- other examined sent the females were not I1 thank charles H lowe department of i eproductivelyreproductively active this included 1 female ecology and evolutionary biology university from may 374 mm SVL I1 from june 370 ofarizonaof arizona and robert L bezy natural his- mm SVL I1 from july 386 mm SVL 2 from tory museum of los angeles county for per- september SVLs 381 mm 393 mm 2 from mission to examine micruroides euryeuryxanthusxanthus october SVLs 363 mm 493 mm funk 1964 and jeffrey feng whittier college for techni- reported 2 oviductal eggs in a single M eury cal assistance estella J hernandez assisted xanthus from pima county arizona collected with histology 20 july stebbins 1985 williamson et al 1994 rossi and rossi 1995 and degenhardt et al literature CITED 1996 reported that clutches contained 2 3 eggs my female sample size is too small to BEHLER JLJ L AND FWEW KING 1979 the audubon society speculate on what proportion of the female field guide to north american reptiles and amphib- ianslans alfred A knopf NY 743 appp produces each year ians population eggs however degenhardt WG CWC W PAINTER AND AHA H PRICE 1996 my finding of I1 may and I1 june females with amphibians and reptiles of new mexico university inactive ovaries may suggest that not all females of new mexico press albuquerque 431 appp reproduce annually the female from july with ERNST CHC H 1992 venomous reptiles of north america smithsonian institution press washington DC 236 inactive no yolk deposition may have ovaries PP already deposited eggs it will be necessary to FUNK RSR S 1964 on the reproduction of micruroides eury examine more female M euryeuryxanthusxanthus before a xanthus kennicott copela 19642191964 219 conclusive statement can be made quinn 1979 GOLDBERG SRS R 1995a reproduction in the western patch found oviductal eggs in only 274 3 female nose snake salvadora hexahexalepislepis and the mountain in patchnosepatch nose snake salvadora grahamitegrahagrahamiaemiae colubridae micrurus fulviusfulvous tener from texas only a por- from arizona southwestern naturalist 4011940 ilg119 120 tion of the female population breeds in other 1995b reproduction in the lyre snake trimortnmorprimor snakes from the north american desert gold- phodonphonon biscutatus colubridae from arizona south- berg 1995a 1995b1995c1995b 1995c 199619971996 1997 western naturalist 4033440 334 335 my 1995c reproduction in the banded sand snake limited data on the micruroides eury chilomeniscuschilomemscusChilomeniscus linctuscinctus colubridae from arizona xanthus ovarian cycle appear compatible with great basin naturalist 5537255 372 373 199711997 NOTES 365

1996 reproduction in the saddled leaflearleafnoseleafhosehosenose snake ico pages 157 172 in W bucherlbucheri and EEE E buckley phyllorhynchus browmbrownibrownl and the spotted leaflearleafnoseleafhosehosenose editors venomous animals and their denomsvenoms volume snake phyllorhynchus decurtatusdecurtatus from arizona jour- II11 venomous vertebrates academic press NY nal of herpetology 3028030 280 282 STEBBINS BCR C 1985 A field guide to western reptiles and 1997 reproduction in the western shovelnose amphibians houghton mifflin boston 336 appp snake chionactisChionactis occipitalis colubridae from cali- williamsonwilliamwilllamWlLLIAM SON MAM A PW HYDER AND JSJ S applegarthAPPLE garthCARTH fornia great basin naturalist 578557 85 87 1994 snakes lizards turtles frogs toads and salaman GOLDBERG SRS R ANDANDWSWS PARKERPARKEB 1975 seasonal testicu- ders of new mexico sunstone press santa fe NM lar histology of the colubrid snakes Masticmasticophisophis tae 176 appp matus and pituophis melanoleucusmelanoleucus Herpetologic a 31 317 322 received 15 february 1997 LOWE CHC H CRC R SCHWALBE AND TB JOHNSON 1986 accepted 2 april 1997 the venomous reptiles of arizona arizona game and fish department phoenix lis115pp115 appp QUINN HRH R 1979 reproduction and growth of the texas coral snake micrurus fulviusfulvous beneretenere copela 1979 453 463 ROSSI JVJ V AND R ROSSI 1995 snakes of the united states APPENDIX and canada keeping them healthy in captivity volume 2 western area krieger publishing company specimens examined from herpetology collections at malabar FL 325 appp the natural history museum of los angeles county romROZE JA 1974 micruroides schmidt western coral snake LACM and the university of arizona UAZ sonora catalogue of americanamerlean amphibians and reptiles mexico LACM 16056 104325 UAZ 9358 27079 35214 1631163163.11 1634163.4163 4 44875 76 45202 203 45904 arizona cochise UAZ 1996 coral snakes of the americas biology iden- 14435 14616 39679 39681 45843 4637846378468345004646834 50046 tification and denomsvenoms krieger publishing company gila 43287 pima UAZ 9360 14429 31 14433 14436 malabar FL 328 appp 14438 39 14441 42 14445 46 14451144511461814618 19 27075 SAINT GIRONS H 1982 reproductive cycles of male snakes 29658 34459 35829 36557 37831 40874 42608 45011 and their relationships with climate and female repro- 46420 47344 47432 47484 48342 48784 finalfinaipinal UAZ ductive cycles Herpetologicherpetologicaherpetologicala 38538 5 16 14432 14434 14444 santa cruz UAZ 39677 yavapai SHAW CEC E 1971 the coral snakes genera micrurus and UAZ 14443 39669396694574845748 micruroides of the united states and northern mex great basin naturalist 574 0 1997 appp 366 368

SPREAD OF PHALARIS arundinacea ADVERSELY IMPACTS THE endangered PLANT HOWELLIA AQUATILIS peter lesicallesica1vesical

key words exotic invasion endangered plant phalaris arundarundinaceaarundmaceamacea howellisHowhowelliaellia aquaaquatilistilis natural area montana

invasive exotic species are considered one flowers and fruits large temporary declines of the primary threats to native communities in abundance occur following years when ponds mooney and drake 1986 and are a major con- fail to dry by late summer lesica personal cern to natural areas managers bratton 1982 observation seeds germinate in montana in harty 1986 exoticaexotics often displace native domi the fall growth begins in the spring and plants nants sometimes altering community function grow up through the water column producing as well as composition vitousek et al 1987 leaves and flowers at or below the surface they are also implicated in the decline of rare flower production starts in june and contin- species but such cases have rarely been docu- ues until drying occurs usually late august mented for plants huenneke and thomson howelliahowellisHowellia aquaaquatilisaquatifistilistifis occurs in ephemeral ponds 1995 lesica and shelly 1996 here I1 present or the margins of shallow permanent ponds evidence that phalaris arundinacea L reed dominated by emergent macrophytes such as canarycanarygrassgrass an aggressive rhizomatous grass equisetumfluviatileequisetum fluviatile L slumsium suave walt and with native and exotic genotypes is gradually the tussock forming carex vesicariavesicaria L these displacing native marsh vegetation including species do not usually form a closed canopy the endangered plant howellishowelliaHowellia aquaaquatilistilis gray and H aquaaquatilistilis generally grows in the spaces between these taller plants it is known histor- SPECIES descriptions ically from northern california western oregon western and eastern washington north- phalaris arundinacea poaceae is a strongly ern idaho and northwest montana however rhizomatous perennial grass native to north- many of these populations are thought to be ex- ern europe and northern north america dore tirtirpated howelliahowellisHow ellia aquaaquatilistilis is considered threat- and mcneill 1980 it has been cultivated for ened or endangered throughout its range in forage and hay in europe since at least the the pacific northwest lesica and shelly 1991 early 1800s schoth 1929 much of the P and is listed as threatened under the federal arundinacea now found in north america is endangered species act thought to be derived from cultcultivarsivars introduced for agriculture dore and mcneill 1980 apfel- STUDY AREA AND METHODS baum and sams 1987 and many of these cul tivarsaivars have been selected for vegetative vigor I1 conducted my study on the nature con- alderson and sharp 1994 phalaris arundinacea serservancyvancy s swan river oxbow preserve along forms dense monomonoculturescultures up to 2 m high in the swan river 5 kmkin south of the town of montana it is capable of invading native wet- swan lake in lake county montana how lands causing declines in native species apfel- ellia aquaaquatilistilis occurs in marshy areas adjacent baum and sams 1987 to a large oxbow slough phalaris arundinacea howellishowelliaHowellia aquaaquatilistilis campanulaceae is an is found throughout many of the wetlands of annual aquatic plant dependent on yearly the preserve where it likely invaded from recruitment from the seed bank seeds of H plantings on the adjacent swan lake national aquaaquatilistilis require an aerobic environment to wildlife refuge germinate lesica 1992 but plants require an phalaris arundinacea occurs on the edge of aquatic environment to grow and produce 2 shallow bays ca 100 m apart of a large river

ithehe natuiendtuiematuie conseconservancycy 32 south ewing helena MT 59601

366 199719971 NOTES 367 oxbow designated north and south marshes 47053n47 53n 1135rw11351w1135rW 945 in elevation that also 80 A ecotone support colonies of H aquaaquatilistilis these colonies so 0 experience the same hydrologic regime that u controls the sometimes significant annual fluc- 22 40 tuationstuations in abundance of H aquaaquatilisaquatifistilistifis see 20 lesica 1992 but have different depths and 0 distribution of dominant vegetation in 1987 I1 1 1 1 1 1 1 1 30 B established a permanent post in the center of ecotone 25 the P arundinacea colony in the north marsh 0 20 distance from the center post to the edge 0 the 15 of the solid P arundinacea sward was measured 2.2 0 Z 10 for 6 radii at 20 intervals 1 I1 interval was 30 0 5 by a hand held compass measurements were 0 1991 I1 y I1 I1 repeated in and 1996 data were analyzed 30 c wholewhoie by repeated measures analysis ofofvariancevariance with 25 marsh each radius treated as an independent measure 20 of the growth of the sward U total area estimates 15 for the 110 of the colony 1987 and 2 portion in 10 1996 were calculated as the sum of the geo- 5 metrically derived areas of each arc section 01 1 1 T I1 in 1988 I1 established 2 permanent transects 86 89 90 91 92 93 94 95 96 to monitor the abundance of H aquaaquatilistilis and year P arundinacea 40 in in the north marsh and 32 in in the south marsh both transects began fig 1 percent canopy cover of A phalaris arundinacea in colonies of P arundinacea and extended in the plansphalaris marsh ecotone B H aquaaquatilistilis in the eco- through carex vesicariavesicaria equisetum fluviatile tone and C howellishowelliaHowellia aquaaquatilistilis in the entire north marsh marsh that supported colonies of H aquaaquatilistilis from 1988 to 1996 I1 used the line intercept method mueller dombois and ellenberg 1974 to estimate cover by 1996 the south marsh of P of solid P arundinacea swardsawards and H aquaaquatilistilis in cover arun dinacea remained stable over the course of the clusters of ifflff aquaaquatilistilis stems were treated as study and was change if they were the crowns of shrubs and the dis- there no apparent in tance overlapped by the line was recorded the canopy cover of H aquaaquatilistilis in the ecotonal when the line crossed a single strand it was zone the the extent solid recorded as 1 cm the minimum distance re- in north marsh of the corded transects were read annually in mid P arundinacea sward intercepted increased for july from 1988 to 1996 4 of 6 radii between 1988 and 1996 and this difference was marginally significant fg 10 2.86 FP 0.10001010 power of this test isi lim- RESULTS AND discussion 286986 010 the ited by the small sample size so P 0100.10 is a phalaris arundinacea colonies consist of a reasonable significance level taylor and ger- monoculture core area surrounded by a periph- rodette 1993 the estimated area of the sam- ery where rametscamets occur among the marsh dom pled P arundinacea sward was 155 m2ma in 1988 inantsimants C vesicariavesicaria E fluviatile and S suave and 209 m2ma in 1996 an increase of 35 in 9 yr in the north marsh cover of P arundinacea my study provides indirect evidence that increased from 20 to 95 in 9 yr in the 3 m howelliahowellisHowellia aquaaquatilisaquatifistilistifis is being displaced by phalaris wide ecotonal zone adjacent to the core area arundinacea in the north marsh the size of at the beginning of the transect in this same the P arundinacea monoculture increased in period canopy cover of H aquaaquatilisaquatifistilistifis declined the north marsh over the course of the study steadily to near zero cover along the entire while the canopy cover ofhofjofhH aquaaquatilistilis declined transect fluctuated but did not show a similar to near zero in the area of invasion even in overall declining trend fig 1 phalaris arun 1996 when abundance reached a 9gyryr high in dinacea was completely absent from the other the entire marsh the fact that H aquaaquatilistilis de- end of this transect in 1988 but was common clined in the presence of P arundinacea while 368 GREAT BASIN naturalist volume 57 increasing elsewhere in the marsh does not acknowledgments prove but strongly suggests interference by P arundinacea bernie hall brian martin anne morley phalaris arundinacea forms dense swardsawards and alfred schuyler assisted with fieldwork that likely produce deeper shade than the more susan winslow and 3 anonymous reviewers open native marsh vegetation furthermore made helpful comments on the manuscript P arundinacea monomonoculturescultures produce deep continuous thatch like litter that decomposes literature CITED slowly to decom- compared the patchy quickly grass produced by ALDERSON J AND WC SHARP 1994 varieties in posing litter the native vegetation the united states USDASCSUSDA SCS agriculture hand- this dense litter layer may inhibit the growth book no 170 washington DC of seedlings bergelson 1990 thereby exclud- ANDERSON DED E 1961 taxonomy and distribution of the ing H equaaquaaquatilisaquatifistilistifis howellishowelliaHowellia aquaaquatilistilis is however genus phalansphalarisphalanx iowa state journal of science 36 capable of persisting in mixtures of P arundiakundi 1 96 APFELBAUM SIS I1 AND CEC E SAMS 1987 ecology and con- neceanecee and native marsh trol of reed canary grass natural areas journal 7 phalaris arundinacea is capable of displac- 69 74 ing native wetland vegetation apfelbaum and BAKER HG 1972 migration of weeds pages 327 347 in H valentine editor taxonomy phytogeography sams 1987 but the rate of invasion is liable to DDH and evolution academic press london depend on individual site characteristics in BARRETT SCHS C H 1983 crop mimicry in weeds economic the north marsh P atundinaceaarundinacea is increasing botany 3725537 255 282 at ca 3 per year toward the center as well as BERGELSON J 1990 life after death site preemption by poapoo around the margins on the other hand extent the remains of boa anduaannua ecology 71215771 2157 2165 BRATTON S P 1982 the effects of exotic plant and animal P of arundinacea in the south marsh did not species on nature preserves natural areas journal increase along the transect over the 9 yr of the 23323 3 13 study DORE WG AND J MCNEILL 1980 grasses of ontarioontano it is generally acknowledged that P arundiakundi monograph 26 research branch agriculture canada ottawa ontario faceanacea is native to the northern portions of HARTY FM 1986 exoticsexotica and their ecological ramifications north america however introduced cultcultivarsivars natural areas journal 642064 20 26 are now common throughout much of north HUENNEKE LEL F AND JKJ K THOMSON 1995 potential inter- america obscuring the geographic range and ference between a threatened endemic thistle and plant biology of 1961 an invasive nonnativenormative conservation habitat native ecotypesecotypes anderson 941642394169 416 423 dore and mcneil 1980 it is generally believed LESICA P 1992 autecology of the endangered plant that invasive populations of P arundinacea are howellishowelliaHowellia aquaaquaticsaquatilisaquatihstilis implications for management and derived from nonnativenormative agricultural cultcultivarsivars reserve design ecological applications 241142124112 411 421 LESICA P AND S SHELLY 1991 sensitive threatened and sams 1987 although JSJ apfelbaum and the mor- endangered vascular plants of montana montana phphological variability of the species makes dis- natural heritage program occasional publication crimination between native and nonnativenormative pop- no 1 helena ulatulationsions impossible anderson 1961 genetic 1996 competitive effects of Centacentaureaubaweagea macumaculosamaculoselosa introgression from cultivated into native eco on the population dynamics of arabis jefecundacunda bulletin of the torrey botanical club 123111123 iiiili111 121211 types resulting in more aggressive weeds has MOONEY HAH A AND JAJ A DRAKE EDITORS 1986 biological been demonstrated in other species baker invasions of north america and hawaii springer 1972 barrett 1983 and may also be occurring verlag new york in P arundinacea MUELLER DOMBOIS D AND H ELLENBERG 1974 aims grom and methods of vegetation ecology john wiley & howelliahowellisHow ellia aquaaquatilisaquatifistilistifis is currently known komhombomfrom 7 sons new york areas in the pacific northwest and P erundina SCHOTH HAH A 1929 reed canary grass USDA farmers cea is present at most of these sites results of bulletin 1602 my study although limited in scope suggest TAYLOR B L AND T gerrodette 1993 the uses of statistical power in conservation biology the faquitavaquita and that this aggressive exotic could reduce popu- northern spotted owl conservation biology 74897 489 500 lations of H aquaaquatilistilis throughout much of its VITOUSEK PM LRL R WALKER LDL D WHITAKER D range although eradication of P arundinacea MUELLER DOMBOIS AND PA MATSON 1987 biolog- may not be possible preventing establishment ical invasion by myrlcaMyricamyricajayamyncafayajaya alters ecosystem development in hawaii science 238802238 802 804 in uninvaded wetlands and controlling spread in in others will be necessary to protect this rare received 20 january 19919977 species accepted 12 may 1997 great basin naturalist 574 0 1997 appp 369 370

UTAH REPTILES OCCURRING ONLY IN SOUTHERN UTAH

wilmer W tannerlnannerltanner1

key words reptiles utah mojave desert

in 1935 dr vasco M tanner published a list arizona elegansdelegans philipi klauber 1964 col- of reptile species that were known to occur in lected by WW tanner near lone rock kane utah the list included 40 species 12 of which county utah were known only from washington county A trionyx spinspiriferaspiniferaifera emoryiemoryy agassiz 1977 col- previous study by woodbury 1931 listed 39 lected by CYCX roby and EC loveless near known species for utah and his recently de- the virgin river approximately 4 miles north scribed subspecies crotalus confluentconfluentusus con- of st george utah color the tanner study listed for the first time phyllorhynchus decurtatusdecurtatus perkperkinsiinsi klauber the worm snake leptotyphlops humilis for utah 1995 collected by russel bazette on road 12 and noted that the southwestern corner of mile north of arizona utah state line beaver utah is a habitat distinctly different from other dam slope utah parts of the state actually the southwestern these species have all entered through the corner is a part of the mojave desert and the southwestern corner of utah arizona elegansdelegans reptile fauna extends from this corner of utah and xantusiaXantusia vigilis and perhaps others may southwest into the deserts of arizona nevada have been in utah for a long time but not pre- and california both the woodbury and tanner viously observed the remaining species may reports list probable species that may occur in be recent entrants why new species are enter- southern utah tanner lists diposaurusdipsosaurusDiposaurus d dor- ing is explainable on the basis of temperature salis uta graciosa heterodon nasicusnasicus micrurus increases an examination of utah climatic euryeuryxanthusxanthus and kinosternonflavescenskinosternon flavescentflavescensflavescens both records for the past 50 yr utah state weather list holbrookiaHolbrookia maculata approximarisapproxinwris but there station logan utah provided by donald T is still some doubt that it occurs in southeastern jensen director indicates that there has been utah in spite of collecting done since these a slow warming of this desert area in recent reports were published none of woodbury s years these records place the slow increase in or tanner s probable species have been found minimum and not maximum daily temperatures in the state the new reptile immigrants except for trionyx since these early lists were published many are all evening or night foragersforagers and would reptile specimens have been added to collec- benefit from warmer and perhaps longer warm tions including 7 to be added to the utah list evenings the continued expansion of the range xantusiaXantusia vigilis baird collected by margaret by night foragersforagers indicates that we may still be starey 1940 near castle cliff beaver dam in the realm of postglacial warming wells and slope washington county utah jorgenson 1964 although published temper- arizona elegansdelegans eburebumataeburnatanata klauber 1946 col- ature records for southeastern utah are not lected by LM klauber near st george utah available the occurrence of reptile species crotalus scutulascutulatustus kennicott 1947 collected such as thamnophis cyrcyrtopsistopsis elaphe gutgutlatalata by AM woodbury on beaver dam slope and probably holbrookiaHolbrookia maculata now in the crotalus miltmiltchellichelli pyrrhus cope 1960 col- four corners area of utah suggests that this lected by WW tanner and GW robison west area has also been invaded by reptiles from of castle rock on beaver dam slope the south and southeast

1ml1mam L bean life science museum brighambnghambangham young university provoprove UT 84602

369 370 GREAT BASIN naturalist volume 57

recently a specimen of tantilla p utahensis literature CITED was taken by dr stanley L welsh at granite creek dolores triangle near the utah colo TANNER VM 1935 western worm snake siagonodon humilis baird and girard found in utah utah rado in border and supports the record reported academy of science arts and letters 1226712 267967 270 by tanner 1966 for colorado two other TANNER WW 1966 A evaluationreevaluationre of the genus tantilla species xantusiaXantusia vigilis and lampropeltis cali- in the southwestern united states and northwestern fornia have extended their range into the mexico Herpetologic a 222134222 134 152 WELLS PK AND CDC D JORGENSON 1964 pleistocene wood upper colorado basin thus even within utah rat biddensmiddens and climatic changes in the mojave desert reptiles seemingly are expanding their desert a record of juniper woodlands science 199 ranges and those requiring a more mesic habi- 996 998 tat such as charina bottaibottae and lampropeltis WOODBURY AM 1931 A descriptive catalog of the rep- pyropyromelanamelana have been restricted to the moun- tiles of utah university of utah bulletin 2151215 1 129 tain habitats received 14 may 1997 accepted 5 june 1997 great basin naturalist 574 0 1997 appp 371 372

BOOK REVIEW birds of zion national park and vicinity is somewhat disconcerting to discover that these roland H wauer utah state university locations are not listed on the map when press logan UT 8432278008432278001997843227800.199784322 780019977800 1997 1495149514.95 referring to the map the reader or visiting softcover birder must understand that lacustrine habitats such as kolobdolob reservoir blue springs lake birds of zion national park and vicinity is a and grafton ponds are located outside the park revised edition of the booklet by the same boundaries yet many significant sightings espe- name published in 1965 by roland wauer and cially of shorebirdsshorebirds have occurred at these dennis L carter whereas the first edition sites and are referenced in the book as part of contained references to 233 bird species this the park s vicinity in addition grafton ponds new version contains an annotated list of 252 are actually water treatment ponds that are fully accepted species plus another 35 not fully fenced and generally inaccessible to the general verified public a fact not mentioned by the author while retaining howard roirollinRolliisillss colorful jerome had a key to the gate full page paintings of birds associated with 4 one of the most worthwhile contributions of zions principal bird communities this edi- by the author is that he lists resident and tion is enhanced by 34 pen and ink sketches migratory birds by the habitat in which they of birds by mimi hoope wolf eight color are most likely to be found and then indicates photographs characterize typical zion national the best time of year for observing them if the park scenery although a picture of the virgin visitor is limited for time and wishes to maxi- river with associated vegetation would have mize the birding experience this information been preferable to the one depicting a riparian is invaluable ie more birds may be found community where no water is visible in the late spring and early summer than at the book is appropriately dedicated to any other time of the year jerome gifford who kept volumes of meticu- it is gratifying to note that mr wauer has lous records of all birds in the vicinity of zion utilized the extensive information gathered from national park for a number of years there is 25 christmas bird counts CBCs in the park hardly a notation to any bird listed therein with- this writer was privileged to participate in many out some reference to his records unfortu- of them with only 1 exception steller s jay nately jerome died in 1988 before his records high and low numbers are listed for each could be published As dr william behle men- species observed on the CBC as well as the tioned in the book s foreword mr wauer has number ofyears any given species has occurred very successfully incorporated all of jerome s it is this attention to detail that makes this outstanding records into this new book much more than just a dust gathering book for A useful map of the park and surrounding the library shelf vicinity at the beginning of the book shows the following statements regarding species numbered locations to which the reader can abundance are instructive this is the most refer when visualizing where any given bird commonly seen owl in the zion park aredarea can be observed this is especially important great homed owl this is one of zionziots s most to a visiting birder inasmuch as many of these characteristic summer birds white throated sites can be reached only by trails shown on swift this is one of zionziols s most characteris- the map locations such as carpenter hill tic birds western wood pewee no other maloney hill and grafton are referred to as bird is so representative of the zion park aredarea places to find band tailed pigeons bushtitsBushtits canyon wren this is one of the park s most and song sparrows respectively however it wide ranging songsongbirdsbirds and is present at all

371 372 GREAT BASIN naturalist volume 57

elevations and habitats solitary vireo this book is that it demonstrates the importance of is one of zion canyonganyoncanyons s most numerous and keeping complete and accurate records obvious songsongbirdsbirds in summer black headed girds ofzionofzion national park and vicinity has grosbeak contributed significantly to our understanding in addition to jerome s records and the of the distribution and abundance of bird CBC data the author gained firsthand experi- species within and around one of the most ence by banding almost 6000 birds during the beautiful national parks in this country the more than 3 yr he spent in the park many author is to be commended for his efforts in sightings of the birds of zion and for that making this information available matter southern utah are the results of mr wauer s commitment to keeping complete and merrill webb accurate records in addition he utilized nest- science teacher ing records from various observers including provo high school his own an outstanding contribution of this provo UT 84604 dhddhdo

T H E

GREAT BBASINA S I1 N naturalistIS F m ao ff

I1I1 hl

A t

I1 N D E X

VOLUME 57 1997

BRIGHAM YOUNG university great basin naturalist 574 0 1997 appp 374 381 INDEX volume 57 1997

authorindexAUTHOR INDEX addis john S 93 goldberg stephen R 85 273 363 andrews tom 21 gomez josejosg M 74 253 atkinson eric 116 griffith JS 238 atkinson melonie 116 groves craig R 116 359 austin dennis D 278 gryska andrew D 338 baker william L 21 hall linnea S 124 barrett reginald H 355 hastriter michael W 281 barton susan H 93 hayes jack PE 83 bauer selena B 231 heaton hoyt J 253 baumann RW 209343909343209 343 heckmann richard A 44 178 beaver donald L 184 hein eric W 348 beck kathryn 352 henderson norman 315 berg louis N 11 hepworth dale K 11 bissonette john A 278 hoffman scott 88 blank robert R 57 hoffhaglehoffnagleHoffhaglehagie timothy L 66 bottorff RL 343 holloway harry L jr 268 bunnell stephen T 263 houseman richard M 209 bunnell S dwight 263 hoy JA 259 bursey charles R 273 hubert wayne A 338 huxman kimberly A 38 caplow 352 florence huxman travis E 38 cheam hay 273 cicero 104 carla inchaustiInchinchaustyausty victor H 44 178 cieminski karen L 327 clarkson robert W 66 jacobi GZ 283 cole kenneth L 315 jaeger michael M 355 conover michael R 278 jimenez jaime E 74 crawford john A 220 220 ebersole james J 50 keegan thomas W eckel EMPM 259 kilgore melissa 149 elliott JC 259 kuyper kristin FE 1 evans howard E 189 lei simon A 155 163 172 fairbanks W sue 149 lesica peter 368 flake lester D 327 lugoortizlugo ortiz CR 283 foutz michael 178 frederick glenn 116 markle douglas FE 142 frederick terry 116 mccafferty WPWE 283 frost herbert H 90 mcclure mark FE 278 fuentes marcelino 74 mihuc janet R 245 mihuc timothy B 245 geluso keith 83 morrison michael L 124

374 199711997 INDEX 375

nowak robert S I1 stamer marc R 38 stromberg JC 198 ottenbacher michael J 11 oconnor candace S 83 tanner wilmer W 70 79 366 tonnesen alex S 50 paul don 263 trent james D 57 phelps JM III111 238 trost charles H 245

ramieranne brigitte M 21 walker lawrence R 155163155 163 robinson anthony T 66 wallace richard L 359 roth jan J 184 webb menillmerrillmen illiiilii 371 ruas claudeteclaudeceClaudete 178 welsh stanley L 352 ruas paulo 178 whitaker shayna J 348 ryan michael G 21 white clayton M 263 whitmore R craig 131 schuppscbupp eugene W 7425374 253 whitmore robert C 131 servheen gregg 116 wilkison richard A 238 shafer david S 315 wilson albert G jr 359 shelley rowland M 231 wilson evelyn M 359 shepherd jay 116 yandell ulla I1 shivik john A 355 young james A 57 shoemaker craigcraigaCrcraiganalgaaigAA 268 young michael K 238 simon david C 142

KEY WORD INDEX

taxa described as new to science in this volume appear in boldface type in this index

abundance 11 broadcast tape recordings 116 dace acanthocephala 273 bromus tectoriumtectorumtectorum 253 kendall warm springs 338 activity patterns 355 brood rearing 220 deer affinity 209 brown trout 238 mule 327 age structure 50 bryophytes 259 white tailed 278 altered andesite soils I1 degraded rangeland 253 amphibia 359 calcium 268 den sites 124 analysis california 85 343 355 densities gradient 21 inyo county 231 population 116 antelope 149 white mountains 124 desert 327 antilocapra americana 149 canis latranslalatranotrans 355 oasis 131 arizona 273 363 cestoda 273 salt 57 astragalus conjunctconjuncturconjunctusus 352 chionactisChionactis occipitalis 85 determination chloroperlidaeChloroperlidae 343 colorimetric 268 baitfishbaitfish introductions 142 classification 21 155 diel activity 238 327 baja california sur mexico 131 cnemidophorus diet 338 bald eagle 184 sonorae 273 differentiation barrier 348 tigris 273 genetic 1 beaver river utah 11 coleogyne ramosissima iss155163155 163 morphological I1 benthic macroinvertebratesmacroinvertebrates 245 colorado 21 diplostomum 44 178 biogeography I1 northwestern 184 dispersal bird records 131 colorimetric determination 268 insect 38 black billed magpie 184 conductance 57 mammalian seed 74 blackbrushblackbrush 172 conjuncti 352 seed74253seed74seed 7425374 253 black greasewood 57 conservation 66 distribution 11 116 189 281 bothriocephalus biology 104 geographic 259 acheilognathi 66 cutthroat trout 11 disturbance 50 348 bristlecone pine 21 douglas fir 116 376 GREAT BASIN naturalist volume 57

eastern fence lizard 348 howellishowelliaHow ellia aquaaquatilistilis 368 morphological differentiation I1 ecology hymenoptera 189 mortality 338 recreation 50 hypsiglena tortorquatatorquatequata 79 mosses 259 ecotype 172 movement 238 electric transmission towers 263 idaho116245327idaho 116245327ilg116 245 327 mule deer 327 electrophoresis insect dispersal 38 eulegemuleg6mulege mexico 131 protein 1 insular populations 104 mustela erminea 83 elevation 172 introduced fish 178 myofibrogranuloma 268 endangered plant 368 introductions endemic species I1 baitbaitfishfish 142 darceusnarceus americanosamericanusameric anus 231 ephemeroptera 283 invasion 368 native 11 ephydatia muellenmuelleri 93 inyo county california 231 natural area 368 eriogonumeriogonunz iodine bush 57 nematoda 273 lobbiilobbia 1 nesting roburobustumstum 1 juniper expansion 74 peregrine falcon 263 eunapiusfragiliseunapiusEunapius fragiasfragihs 93 juniperus prairie falcon 263 exclosures 245 monomonospermamonospermysperma 50 nevada exotic 368 occidentoccidentalisoccidentalistalis 74 lucky strike canyon 155 expansion osteosperma 253 163 juniper 74 sheep range155Range 155 range131278range131range 131278131 278 kendall warm springs dace spring mountains 155 eye fluke 44 338 new mexico 283 348 nicaragua 79 falco laboratory release 142 nonnativenormative species 142 mexicanusmexic anus 263 lagomorphslagomorpha 253 northwestern colorado 184 peregrinus 263 Lemmislemniscuslemmiscuscus curcurtatustatus 83 fathead minnow 142 life cycle oncorhynchus flammulated owl 116 parasite 44 clarki 178 floods198floods 198 Lincollincolnrss sparrow 104 mykiss 238 fossil pollen 315 little colorado river 66 ordination 155 freshwater slsponges 93 livestock grazing 104 oregon142Oregonoregon142220142220142 220 frugivoryfrugivorafrugivory 74 lizard otusflammeolusotus flamflammeolusglammeolus 116 eastern fence 348 genetic differentiation I1 lower coleogyne ecotone 163 pankratpackrat biddensmiddensmiddens 315 geographic distribution 359 lucky strike canyon nevada parasite life cycle 44 germination 172 155163 parasitism 66 cilacliagila cypha 66 patterns golden eagle 184 macromacroinvertebratesinvertebrates activity 355 gradient analysis 21 benthic 245 peregrine falcon nesting 263 grazing mammalian seed dispersal 74 perognathus carvusparvus 83 impacts 315 mammals 74 peromyscus trueithuei 83 124 livestock 104 management 11 50 245 phalaris arundinacea 368 greasewood meleagris gallopavogallopavo intermediainterintermedialmedia photosynthesis 57 black 57 220 pine great basin 209 melospiza bristlecone 21 lincolniilincollincolnianii 104 ponderosa 116 habitats 116 melodia 104 pinus association 104 metacercariae 178 edulisadulis 50 availability 149 mexico 7936379 363 ponderosa 116 riparian 198 baja california sur 131 pinyon mouse 124 selection 149 eulegemuleg6mulege 131 pinyon juniper woodland 50 use124220use 124220124 220 micrositemicrolitemicrosite preference 50 plant winter 149 micruroides euryeuryxanthusxanthus 363 endangered 368 helminthshelminthes 273 biddensmiddens plecoptera 209 343 high altitude 83 pacpackratpankratkrat 315 pleistocene 209 hiltoniusHiltonius 231 minnow plethodon 359 holocene vf fetationgetationgevegetationtation fathead 142 pollen history 315 mojave desert 163172366163 172 366 fossil 315 home range 238 montana 93 259 368 pompilidae 189 homing 348 montane meadow 104 ponderosa pine 116 199711997 INDEX 377 ponds 327 sevier river utah 11 trout population densities 116 sheep range nevada 155 brown 238 populusfremontiipopulus fremonfremontiifremontiatiitil 198 shorebirdsshorebirds 245 cutthroat 11 porifera 93 sierra nevada 355 rainbow 238 prairie falcon nesting 263 siphonaptera 281 tunga monositusmonositus 281 prechill 172 sites tylotustylobusTylobus 231 predation 245 characteristics 220 frederickfredfredericksonfredericksomfredericksoniericksomsoni 231 presettlement vegetation 315 den124den 124 utahensis 231 pronghomprongpronghornhom 149 327 snake protein electrophoresis 1 western coral 363 urban areas 278 pseudocrossidium 259 western shovelnose 85 utah 209 278 366 crinitumcrinitum 259 soils 163 beaver river 11 homschuchianum 259 altered andesite 1 raft river mountains 209 obtusulum 259 song sparrow 104 sevier river 11 replicatumreplicatum 259 southwestern utah 11 southwestern 11 revolutumrevolutum 259 species virgin river 11 pseudotsuga menziemenziesiimenziesiasii 116 endemic 1 washington county 231 interactions 142 radio telemetry 124 220 inventory 283 vegetation 21 raft river mountains utah nonnativenormative 142 holocene 315 209 spider wasps 189 presettlement 315 rainbow trout 238 sponges zones155zones iss155 range freshwater 93 virgin river utah 11 expansion 131 278 spongilla lacustrislacnstnslacustris 93 land degraded 253 spongillidae 93 walleye 268 restoration 253 spring mountains nevada washington 352 records 155 washington county utah 231 bird 131 stizostedion vitreum 268 wastewater 327 recreation ecology 50 stonestonefliesflies 209 343 water regeneration 50 strawberry reservoir 44 potential 57 reproduction 85 338 363 structure use efficiency 57 reptiles 366 age50age 50 watering frequency 172 rhinichthys Suwallia 343 weevilsweevily culusosculusesculusoscultisoscoltiscultis 66 pipisteripinteristeri 343 yucca38yucca 38 osculusesculus therthennalisthermalisthennalismalis 338 sierra 343 western coral snake 363 rio grande wild turkey 220 sweltsaSweltsa 343 western shovelnose snake 85 riparian habitats 198 white mountains california rocky mountains 21 359 tamarixtamanyramany chinenchinensissis 198 124 rotenone treatment 44 tape recordings white tailed deer 278 broadcast 116 winter sagebrush 184 teiidae 273 habitat 149 salamander 359 telemetry population 184 salix goodingii 198 radio 124 220 woodland salmo trutta 238 temperature 172 pinyon juniperjumper 50 salt desert 57 towers wyoming 178 salvelinus namaycush 178 electric transmission 263 sceloporus undulatesundulatus 348 trampling 50 yucca weweevilsweevilyevils 38 scyphophorusscyphophorous yuccasyuccae 38 translocation 348 yucca whippleiwhipplerwhzppleiwhippleipiel 38 seed dispersal 74 253 transpiration 57 seedling 50 zoogeography 209 378 GREAT BASIN naturalist volume 57

TABLE OF CONTENTS volume 57

no I1 january 1997 articles on the taxonomic status of eriogonum roburobustumstum polygonaceae a rare endemic in western nevada kristin FE kuyper ulla yandell and robert S nowak I1 distribution and abundance of native bonneville cutthroat trout oncorhynchus clarki utah in southwestern utah dale K hepworth michael J ottenbacher and louis N berg 11 natural variability of vegetation soils and physiography in the bristlecone pine forests of the rocky mountains brigitte M ranne william L baker tom andrews and michael G ryan 21 dispersal characteristics of the yucca weevil scyphophorusscyphophorous yuccasyuccae in a flowering field of yucca whippleiwhipplerwhipplei travis E huxman kimberly A huxman and marc R stamer 38 evaluation of fish diplostomatosis in strawberry reservoir following rotenone application a five year study victor H inchaustiInchinchaustyausty and richard A heckmann 44 human trampling effects on regeneration and age structures of pinus edulisadulis and juniperus monospermamonospennamonospermymonomonos pennasperma alex S tonnesen and james J ebersole 50so physiologyEcoecophysiology ofthe temperate desert halophytes ailenalienAllenallenroeaallenrolfearofaroea occidentoccidentalisoccidentalistalis and sarcobatus vermicavermicu latus james D trent robert R blank and james A young 57 asian tapeworm bothriocephalus acheilognathi in native fishes from the little colorado river grand canyon arizona robert W clarkson anthony T robinson and timothy L hoffnagle 66 dorald M allred 1923 1996 wilmerwiimerwilmerewilmerwwllWilmerWW tanner 70 notes dispersal of juniperusofjuniperus occidentoccidentalisoccidentalistalis western juniper seeds by frugivorous mammals on juniper mountain southeastern oregon eugene W schupp josejosg M gomez jaime E jimenez and marcelino fuentes 74 type locality restriction hypsiglenaofofhypsiglena torquatatorquatetorquata guntherguinther wilmer W tanner 79 elevational records for mammals in the white mountains of california keith geluso candace S oconnor and jack PE hayes 83 reproduction in the western shovelnose snake chionactisChionactis occipitalis colubridae from california

I1 stephen R goldberg 85 book review the western san juan mountains their geology ecology and human history rob blair editor scott hoffman 88 the natural history of a mountain year four seasons in the wasatch range claude T barnes herbert H frost 90 199719971 INDEX 379

no 2 april 1997 articles freshwater sponges porifera spongillidae of western montana susan H barton and john S addis 93 boggy meadows livestock grazing and interspecific interactions influences on the insular distribution of montane lineolLincollincolnrss sparrows melospiza lincollincolniifincolniilincolnianiinil altial coldcoidticola carla cicero 104 density distribution and habitat of flammulated owls in idaho craig groves terry frederick glenn frederick eric atkinson melonie atkinson jay shepherd and gregg servheen 116 den and relocation site characteristics and home ranges of peromyscus truel in the white mountains of california linnea S hall and michael L morrison 124 late fall and early spring bird observations for eulegemulegemuleg6 baja california sur mexico robert C whitmore and R craig whitmore 131 Interinterannualannual abundance of nonnative fathead minnows pimephales kromelaspropromelasmelas in upper klamath lake oregon david C simon and douglas E markle 142 winter habitat selection by reintroduced pronghorn on antelope island great salt lake utah melissa J kilgore and W sue fairbanks 149 classification and ordination coleogyneofcoleogyneof communities in southern nevada simon A lei and lawrence R walker 155 biotic and abiotic factors influencing the distribution of coleogyne communities in southern nevada simon A lei and lawrence R walker 163 variation in germination response to temperature and water availability in blackbrushblackbrush coleogyne ramosissima and its ecological significance simon A lei 1721 72 diplostomiasis in native and introduced fishes from yellowstone lake wyoming victor H inchaustiinchaustyInchausty michael foutz richard A heckmann claudeteclaudeceClaudete ruas and paulo ruas 178 winter survey of raptorsraptores with notes on avian scavengers in northwestern colorado donald L beaver and jan J roth 184 errata 187

no 3 july 1997 articles spider wasps of colorado hymenoptera pompilidae an annotated checklist

I1 howard E evans 189 growth and survivorship of fremont cottonwood goodding willow and salt cedar seedlings after large floods in central arizona JC stromberg 198 zoogeographic affinities of the stonestonefliesflies plecoptera of the raft river mountains utah richard M houseman and richard W baumann 209 brood rearing habitat use by rio grande wild turkeys in oregon thomas W keegan and john A crawford 220 distribution of the millipedmillimillipedeped tylobolusTylobolus utahensis chamberlin with remarks on T fredericksonifredericksoni causey Spirospirobolidabolida spirobolidae rowland M shelley and selena B bauer 231 contrasting movement and activity of large brown trout and rainbow trout in silver creek idaho michael K young richard A wilkison JM phelps III111 and JS griffith 238 380 GREAT BASIN naturalist volume 57 shorebird predation on benthic macromacroinvertebratesinvertebrates in an irrigation reservoir janet R mihuc charles H trost and timothy B mihuc 245 lagomorphslagomorpha and the dispersal of seeds into communities dominated by exotic annual weeds eugene W schupp hoyt J heaton and josejosg M gomez 253 pseudocrossidium obtusulum pottiaceae bryopsida new to montana with a key to north american species in the genus PM eckel JAJ A hoy and JCJ C elliott 259 stick nests on a building and transmission towers used for nesting by large falcons in utah stephen T bunnell clayton M white don paul and S dwight bunnell 263 effects myofibrogranulomaofofmyofibrogranuloma on serum calcium levels in walleye stizostedion vitreum

I1 craig A shoemaker and harry L holloway jr 268 notes helminthshelminthes from the sonoran spotted whiptail cnemidophorus sonorae and the western whiptail cnemidophorus tigris sauriasauna teiidaetendaeteddaeoehTenoen dae from southern arizona with comments on abbreviateabbreviata terrapenisterraterrapemspenis nematoda physaloptendaephysalopteridae stephen R goldberg charles R bursey and hay cheam 273 range expansion of white tailed deer odocoileus virginianusvirgzmanusvirginianuslanus into urban and agricultural areas of utah maikmarkmalk FE mcclure john A bissonette michael R conover and dennis D austin 278 establishment of the tingidtungid flea tunga monositusmonositus siphonaptera pulicidae in the united states

I1 michael W hastriter 281

no 4 october 1997 articles mayfly fauna of new mexico WP mccafferty crlugoCcolugoR lugoortizlugo ortiz and GZG Z jacobi 283 holocene vegetation and historic grazing impacts at capitol reef national park reconstructed using packratpankrat biddensmiddens kenneth L cole norman henderson and david S shafer 315 mule deer and pronghorn use ofofwastewaterwastewater ponds in a cold desert karen L cieminski and lester D flake 327 observations on the reproduction sources of mortality and diet of the kendall warm springs dace

I1 andrew D gryska and wayne A hubert 338 two new species of chloroperlidaecbloroperlidaeChloroperlidae plecoptera from california RWR W baumann andandrlandalRL bottorff 343 homing in eastern fence lizards sceloporus undulatusundulates following short distance translocation eric W heinhemheln and shayna J whitaker 348 new variety of astragalus conjunctconjunctnsconjunctusconjuncturnsus S watson from benton county washington

I1 stanley L welsh florence caplow and kathryn beck 352 notes coyote activity patterns in the sierra nevada john A shivik michael M jaeger and reginald H barrett 355 USU S distribution of the coeur dalene salamander plethodon idahoensis slater and slipp albert G wilson jr evelyn M wilson craig R groves and richard L wallace 359 reproduction in the western coral snake micruroides euryeuryxanthusxanthus elapidae from arizona and sonora mexico stephen R goldberg 363 199711997 INDEX 381 spread ofot phalaris arundinacea adversely impacts the endangered plant howellishowelliaHowellia aquaaquatilistilis peter I1 I1 I1 lesica 366 utah reptiles occurring only in southern utah wilmer W tanner 369 book review birds ofofzionzion national park and vicinity roland H wauer merrill webb 371 the shrub research consortium and the great basin environmental education center are sponsoring the loth WILDLAND SHRUB SYMPOSIUM

august 12 14 1998 snow college ephraim utah

shrubland ecotonesEcotones is the theme of the symposium which will feature a mid symposium field trip to the great basin experimental range and to hybrid zones in salt creek in the uinta national forest contributed papers and posters are invited and should address the following topics

succession within and between the role of boundaries in biology communities management and restoration of diversitybiodiversitybio various shrubland communities hybrid zones and their interfaces with other edge effects communities other subjects on shrubland biology

proceedings of the symposium will be published by the USDA forest service and distributed to each registrant participants interested in making an oral or poster presentation should send the title and an abstract of 200 words or less before january 15 1998 to durant mcarthur at the address below registrationpreregistrationpre materials and information concerning lodging can be obtained from dave lanier

dr E durant mcarthur dave lanier shrub sciences laboratory great basin environmental 735 north 500 east education center provo UT 84606 150 east college avenue phonephone801377801 3775717377 5717 ephraim UT 84627 emaiemalemailemallemaliE maimalmailmaiimali1 SEMCARTHsemcarthurou1U RCU 1 phonephone801283801 2837261283 7261 s22amhs fswaattmail emailE mail daveldavelstormsnowedustorm snow ed u COM information FOR AUTHORS the great basin naturalist welcomes prepreviouslyviouslyvinously VOUCHER SPECIMENS authors are encouraged to unpublished manuscripts pertaining to the biologi- designate properly prepare label and deposit ccalal 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 1989 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 chronological order use et al after name of 84602020084602 0200 an accompanying cover letter must first author for citations having more than two include phone numbers of the author submitting authors the manuscript and FAX number and emailE mail acknowledgments under a centered main address when applicable the letter must also pro- heading include special publication numbers when vide information describing the extent to which data appropriate text illustrations or have been used in other papers literature CITED also under a centered main or books that are published in press submitted or heading lists references alphabetically in the fol- soon to be submittedsubmitted elsewhere authors should lowing formats adhere to the following guidelines manuscripts not so prepared be returnedfor forbbrmor may returned revision mack GD and flake 1980 habitat relation- manuscript preparation great LD in general the ships of waterfowl broods on south dakota basin naturalist follows recommendations in stock ponds journal of wildlife management scientific style and format the CBE manual for publishers 44695 700 authors editors and ath6th edition sousa WPWE 1985 disturbance and patch dynamics council of biology editors inc 11 south lasalle on rocky intertidal shores pages 101 124 in street suite 1400 chicago IL 60603 USA PHONE STA pickett and PSES white editors the ecolo- 3122010101312 201 0101 FAX 3122010214312 201 0214 we do however gy of natural disturbance and dynamics differ treatment of patch in our entries in literature cited academic press new york authors may consult vol 5511 no 2 of this journal coulson RN and 1984 for specific instructions on format these instruc- JA witter forest entomology ecology and management john wiley tions guidelines FOR manuscripts SUBMITTED 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 of the great basin naturalist for changes TABLES are double spaced on separate sheets and designed to fit the width either a single column TYPE AND DOUBLE SPACE all materials including of literature cited table headings and figure legends or a page use lowercase letters to indicate foot- avoid hyphenated words at the righthandright hand margins notes use WordwordperfectPerfect s italics feature for words to be photocopies OF FIGURES are submitted initially printed in italics use standard bond 22x28x28 cm witwith the manuscript editors may suggest changes leaving 25 cm margins on all sides lettering on figures should be large enough to SUBMIT 3 COPIES of the manuscript 5 copies of withstand reduction to one or two column width fish manuscripts and the original on a 35 inch disk originals must be no larger than 22 X 28 cm utilizing 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purpose methods results cover letter explaining any duplication of and conclusions of the research it is followed by information and providing phone numbers 6 12 key words listed in order of decreasing FAX number and emailE mail address importance to be used for indexing 3 copies of the manuscript 5 copies of fish TEXT has centered main headings printed in all papers and WordwordperfectPerfect 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 issn0017133n 001736140017 3614 GREAT BASIN naturalist volvoi 5757.57 no 4.4 octoberoctoberi9971997

CONTENTS articles mayfly fauna of new mexico WPWE mccafferty crlugocolugo ortiz and GZ jacobi 283 holocene vegetation and historic grazing impacts at capitol reef national park reconstructed using packratpankrat biddensmiddens kenneth L cole norman henderson and david S shafer 315 mule deer and pronghorn use ofwastewaterof wastewater ponds in a cold desert 11 I1 karen cieminski and lester D flake 327 observations on the reproduction sources of mortality and diet of the kendall warm springs dace andrew D gryska and wayne A hubert 338 two new species of chloroperlidaeChloro perlidae plecoptera from california RW baumann and RL bottorff 343 homing in eastern fence lizards sceloporus undulatusundulates following short distance translocation eric W hein and shashaynaanayna J whitaker 348 new variety of astragalus conjunctconjuncturconjunctusus S watson from benton county washington stanley L welsh florence caplow and kathryn beck 352 notes coyote activity patterns in the sierra nevada john A shivik michael M jaeger and reginald H barrett 355 US distribution of the coeur dalene salamander plethodon idahoensis slater and slipp albert G wilson jr evelyn M wilson craig R groves and richard L wallace 359 reproduction in the western coral snake micruroides euryeuryxanthusxanthus elapidae from arizona and sonora mexico stephen R goldberg 363 spread ofphalarisofphalaris arundinacea adversely impacts the endangered plant howelliahowellisHowellia aquaaquatilistilis peter lesica 366 utah reptiles occurring only in southern utah wilmer W tanner 369 book review birds ofofzionzion national park and vicinity roland H wauer merrill webb 371 index to volume 57 373