Full Issue, Vol. 57 No. 3

Great Basin Naturalist

Volume 57 Number 3 Article 15

7-31-1997

Follow this and additional works at: https://scholarsarchive.byu.edu/gbn

Recommended Citation (1997) "Full Issue, Vol. 57 No. 3," Great Basin Naturalist: Vol. 57 : No. 3 , Article 15. Available at: https://scholarsarchive.byu.edu/gbn/vol57/iss3/15

This Full Issue is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. T H E

GREAT BASBASINI1 naturalistnaturalist

ale

VOLUME 57 ngN 3 JULY 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 youhgyoung university brigham young university provo UT 84602020084602 0200 provo UT 84602687984602 6879 8013785053801 378 5053 8017378801378668880173786688801 378 6688 FAX 8013783733801 378 3733 emailE mail nmshbllibyuedunmshbll1byuedu

associate editors J R CALLAHAN PAUL C MARSH museum of southwestern biology university of tentercentergenter 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 Biologicbiologicalajlainaln sciences university of university of nevada las vegas wisconsin whitewawhitewaterwhitewayterten whitewater WI15353190190igo las vegas NV 89154400489154 4004 JEFFREY J JOHANSEN PAUL T TUELLER department of biology john carroll university depdepartmentartmentartmont 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 VWVIVV 26506612526506 6125

editorial board berranjerran T flinders chaichaitmanchairmanrniananian 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 boboardard 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 thegreatthe great basin naturalnaturalistist for 1997 are 25 for individual sub- scriscribersbers 30 outside the united states and 50 formor institutions thetho ppriceriedriep 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 basinnatubasinNatunaturalistraliwt 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 houldshoulds contact the exchange librarian 6385 HBLL PO box 26889 brigham young university provo UT 8460268898466268898460284662 6889

editorial production staff joanne abel technical editor

copyright D 1997 by brigham young university ISSN 001736140017 3614 official publication date 31 july 1997 7977 97 750 22825 the great basin naturalist PUBLISHED AT PROVO UTAH BY BRIGHAM YOUNG university

ISSN 001736140017 3614

VOLUME 57 31 JULY 1997 no3nonoa 3

great basin naturalist 573 0 1997 appp 189 197

SPIDER WASPS OF COLORADO hymenoptera pompilidae AN ANNOTATED CHECKLIST

howard E evanslevanelevans1

ABSTRACT one hundred forty three species of pompilidae are recorded from colorado slightly more than half the number occurring north of mexico some of these occur principally at higher altitudes or in the northern part of the state this group includes 5 species of holarctic distribution others such as the tarantula hawks pepsis are prevalent across the southern third of the state and range south into new mexico and often into mexico still others are widely distributed wherever there is friable soil suitable for nesting certain genera are more or less restricted to preying upon certain spider taxa while others are generalists and a few are cleptoparasites of other pompilidae

key words hymenoptera pompilidae spider wasps distribution

spider wasps are ubiquitous insects occur- 11 caught in a snow driftdriffdrife at 13000 feet 3900 m ring wherever there are spiders they use these on mt rogers clear creek county this indi- arthropods to provision their nests employing vidual was undoubtedly blown there by the a single paralyzed spider per cell in colorado wind they are most plentiful on the eastern plains most colorado species are believed to be especially in sandy country along the valleys univoltine the adults active mid june through of the arkansas and south platte rivers how- early september only I1 species anopliusAnoplius ever they also occur widely in the western tenebrotenebrosussus cresson is known to overwinter two thirds of the state including the moun- as an adult all others are believed to overwin- tains I1 have taken 61 species around my ter as diapausingdiapausing larvae or pupae in their nest home in open ponderosa pine douglas fir cells woodland at 2300 m in larimer county the in this report I1 list 143 species from colo- holarctic species anopliusAnoplius nigerrimus scopoli rado slightly more than half the species known has been taken above timberline at 3600 rn on to occur north of mexico only minimal data trail ridge in rocky mountain national park are included concerning habitat and behavior evans 1951 A female of anopliusAnoplius tenebrotenebrosussus evans and yoshimoto 1962 reviewed the cresson in good condition in the university nesting behavior of species occurring in the of colorado collection is labeled as having been northeastern states and many of these same

department of entomology colorado state university fort collins CO 80523

189 190 GREAT BASIN naturalist volume 57 species and all genera occur in colorado the the largest spider wasp in our fauna and most recent catalog of north american hyme- one of the largest in the world is pepsis for- noptera krombeinKrombem et al 1979 includes further mosa say in which large females may have a references major papers on the systematics body length of 5 cm and a wing span of 9 cm of nearctic pompilidae are those of evans in contrast males of some of the smaller 1950 51 and townes 1957 although there species of ageniella have a body length of 2 have been several refinements in the classifi- mm with a wing span of about 3 mm cation in the decades since those papers were adult spider wasps of most species take published nectar at flowers and it is here that most are the colorado fauna includes 5 species that taken by collectors flowers with shallow also occur in eurasia these species range cocorollasrollas such as apiaceae umbelliferae are throughout the northern hemisphere and occur especially favored other genera that are fre- chiefly in the northern mountains Evageevagetestes quently visited include asclepias baccharis crassicorniscrassicrassicormscornis shuckardShucschuckardkard Ananophusadophusanopliusoplius nigernmusnigerrimus cleome euphorbia Mehmelilotuslotusiotus sapindus sol- scopoli arachnospilafumipennisarachnospila fumipenms zetterstedt idago and tamarixThmarix honeydew from extraflo- cahadurguscaliadurgusfasciatellusfascifasciatellusatellusatelius spinolaSpmolamoia and ceropalesCeropales ral nectaries and from aphids and other suck- maculata fabricius in contradistinction quite ing insects is also visited frequently especially a number of species characteristic of the deep on helianthus southwest and mexico enter colorado in the in this paper I1 accept the strictures of southernmost third of the state for example the menke 1990 though with misgivings as species ofofpepsisrepsisrupsispepsis and psorthaspis three species these result in the use of several unfamiliar have been reported only from montezuma names for well known species some of these novel county in extreme southwestern colorado synonymiessynonymicssynonymies are based on the discovery calopompilusCalopompilus pyrrhomelas walker priocneprocnepnocne by day 1977 of broken specimens with incor- locality mis fregonaoregona banks and apolusaporus luxus banks rect data that are claimed to represent recently several species characteristic of north american species others are based on the an unfor- eastern forests have appeared in colorado such rejection of secondary homonyms tunate procedure evidently part of the code as Priopriocnemispnocnermscnemis noratamiminoritaminorata banks and auplopusAuplopus now mellipesmelligesmellipes variharsatusvamtarsatus dalla torre similarly of zoological nomenclature I1 also accept the nomenclatural changes several typically west coast species have been taken here such as cryptocheilus hesperus suggested by shimizu 1994 based on his research and that of several european work- banks and diposondipogon seriserlseflsericeussericeoussenceusceus banks it is pos- ers most especially day 1981 arrangement sible that some of these species have been of the genera follows that in the catalog krom- introduced in commerce bein et al 1979 except in a few cases and with most spider wasps prepare simple nests in some changes in generic names list is the soil but a few make mud cells above- the based primarily on specimens in the collections ground A few oviposit on spiders directly of colorado state university collins the without taking them to a nest and some are fort university of colorado boulder and the parasites of other pompilidae of cer den- species ver museum of natural history some records opalsopales and Evageevagetestes most spider wasps are have been taken from the literature black in color with translucent or fumose wings A few have banded wings and some mostly SUBFAMILY PEPSINAE larger species have bright orange wings orange is an aposematic warning color in insects it genus calopompilusCalopompilus ashmead is believed that birds learn to associate this genus was called chirodamusChirodamus by color with undesirable qualities in this case a this townes 1957 but the type of genus is a potent sting and to avoid attacking such insects that south american species of doubtful affinities fifteen species occurring in colorado have with north american species wasps orange wings forming a complex of what are these are infrequently collected and nothing is often called mullerianmuiMullenan mimics there are other known of their predatory or nesting behavior examples of apparent mimicry species with yel- our I1 species has brilliant orange red wings low banding suggesting social wasps and others with orange patterns on the body suggest- pyrrhomelas walker montezuma co A single ing mutillidae velvet ants female was collected 7 august 1929 by 199719971 COLORADO SPIDER WASPS 191

PR franke in mesa verde national park recorded from colorado are rarely encoun- there is a female in the university of wyo- tered and nothing is known of their behavior ming collection from laramie WY col- there are several records of an eastern species lected 26 september 1964 by RE pfadt nebulosus dahlbom preying upon grass spi- the species was reported by townes 1957 ders of the genus agelenopsis from british columbia to arizona with no records from colorado or wyoming apache banks Huerhuerfanofano and larimer counties townes 1957 recorded this species genus pepsis fabricius only from southern arizona and texas

11 coloradensis banks boulder and these are the tarantula hawks of large clear creek larimer counties size and with bright orange wings females hunt for mygalomorph spiders tarantulas genus entypus dahlbom paralyze them by stinging and place them either in the spider s burrow or in a shallow these are wasps of moderate size most of nest dug in the soil hurd 1952 reviewed the the species having orange wings and resem- north american species and summarized data bling small tarantula hawks several of the on behavior our 5 species are mostly con- species have been found to prey on wolf spi- fined to the southern third of the state ders of the genus lycosa but so far as I1 am aware the nests have never been found angustimarginata viereck otero and pueblo counties aratus townes baca bent and otero counties fonnosafonnosaformosa formosa say bent co say described austaustrinusaustfinusausfinusninushinustrinus austrinusaustrinus banks bent boulder kio- this species from the arkansas valley and wa las animas and otero counties hurd recorded it from several unspecified fulvifulvicomiscomis cresson bent and larimer coun- sites in southern colorado but it has rarely ties been collected there in recent years tetanustexanus tetanustexanus cresson baca bent crowley mildeimiddei stal baca bent crowley otero prow- Huerhuerfanofano kiowa larimer las animas ers and pueblo counties otero prowers and pueblo counties pallidolimbata pallidolimbata lucas mesa and unifasciatus cressonicrescressionisoni banks boulder clear montezuma counties A male in the col- creek crowley el paso gunnison lari- orado state university collection labeled mer and weld counties ft collins 1112411 1 24 is surely mismislabeledlabeled unifasciatus unifasciatus say yuma co an thisbe lucas alamosaalamona baca bent crowley el eastern subspecies paso Huerhuerfanofano kit carson las animas otero and prowers counties several rather genus cryptocheilus panzer exceptional records are from boulder the members of this genus closely resem- douglas and weld counties all are males ble those of Entyentypuspus but average smaller As one taken 5 july the others in august in Entyentypuspus the females prey on wolf spiders they may have been strays from breeding of the genus lycosa the nests are multicellu- populations farther south lar and are dug from preexisting cavities in the soil genus hemipepsisHemipepsis dahlbom these wasps are superficially similar to attenuatesattenuatus banks boulder denver jefferson mesa species of pepsis and like them are predators larimer and otero counties on mygalomorph spiders williams 1956 hesperus banks crowley fremont lincoln and otero counties ustustulateustulataulata ustulataustulateustulata dahlbom archuleta baca idoneum birkbirkmannimanni banks bent conejos bent el paso fremont Huerhuerfanofano las Huerhuerfanofano kit carson otero prowers mesa animas montezuma and otero saguache sedgwick and weld counties counties severini banks baca bent boulder clear creek crowley genus priocnessus banks denver fremont huer fano larimer las animas logan morgan compared to the previous 3 genera these otero prowers sedgwick and weld coun- are smaller slender bodied wasps the 2 species ties 192 GREAT BASIN naturalist volume 57

terminatestermiterminatusnatus terminatestermiterminatusnatus say archuleta bent thomisidae cells are separated by barriers boulder elbert garfield gunnison huer of debris bits of soil wood seeds dead insects fano jefferson larimer las animas mesa the particles being carried with the help of the pitkin and weld counties beards members of this genus have rarely been collected in colorado being more partial genus Priopriocnemispnocnermscnemis schiodteschiodtz to deciduous forests this is a large genus of small wasps that iracundiracundusus townes boulder co a single female prey upon a great diversity of ground dwelling taken by CC lanham at nederland spiders nests are multicellular and are often 2500 in dug from preexisting cavities in the soil in lignicoluslignicolous evans larimer co type locality and aequalisquallsquails banks larimer co a single female only records 21 km W of livermore taken 21 km W of livermore at 2300 in 2300 in elevation sayi nigrior townes larimer co a single female cornica say baca denver garfield kiowa collected by david leatherman at estes larimer mesa teller and weld counties park 2400 inm townes 1957 recorded the germana cresson delta and laplatalaplaca coun- species from colorado but without a spe- ties cific locality kemm kasbauerwasbauerWasbauer larimer co 2 males taken serisericeussericeousceus banks larimer co a series of both in a malaise trap 21 km W of livermore sexes taken in a malaise trap in hewlett 23002300min gulch 18 km NW fort collins 1900 in noratamiminoritaminorata banks larimer and el paso counties genus minageniaMinagenia banks navajo navajo banks garfield larimer mesa although townes 1957 placed this genus mineral and routt counties in the subfamily ceropalinae I1 regard it as a notha notha cresson boulder garfield huer probable derivative of Priopriocnemiscnemis these are fano LalaplatalaplacaPlata larimer morgan and routt delicate wasps that do not build nests but counties this and the preceding species oviposit directly upon lycosid spiders which are broadly sympatric and differ in details are left in place kaston 1959 of the male termterminaliamaliamallamaila navajo in the past has been considered only subspecifically congrua cresson larimer co fort collins distinct from notha but the two are clearly montismontisdorsadorsa dreisbach bent co hasty full species genus ageniella banks fregonaoregona banks montezuma co mesa verde national park this is the first of 3 genera of unusually scitulascatulascvtula felictarelicta banks larimer co fort collins slender body form in which the females normally amputate some or all of the spiders legs genus pate caliadurgus before transporting them to the nest species this is a small genus with a single uncom- of ageniella use a great variety of small spiders mon species in colorado prey is reported to as prey and so far as studied place them in be orb weaving spiders that are flushed from short burrows dug from preexisting cavities in their webs the nest is a shallow burrow in the soil sixteen species are reported from the soil colorado fasciatellusfasciatellus alienatusahenatusalienatesaheabealiealleailenatus smith logan and pueblo accettaaccepta cresson baca bent boulder huer counties fano larimer mesa moffat otero prow- fasciatellusfasciatellus excoctus townes larimer co ers pueblo and weld counties genus agenioides fox boulder and larimer coun- diposondipogon fox ties females of this genus have paired tufts of arapaho evans larimer co type locality and bristles on the maxillarmaxillaemaxillae hence the name only records 21 km W livermore 2300 in diposondipogonDipogon two beard they nest in hollow arcuataarcuate banks boulder larimer logan and twigs and some will accept wooden trap nests montezuma counties cells are provisioned with small spiders of var- arizonicaarizonica arizonicaarizonica banks otero co 28 km S ious genera but most commonly crab spiders lajunta 199719971 COLORADO SPIDER WASPS 193 conflictconflictsconflictaa banks bent boulder garfield lari- nignigrellusrellus banks bent boulder chaffee mer montezuma morgan and weld coun- denver douglas el paso and larimer ties counties cupida cresson larimer co poudre canyon 1600 in townes 1957 SUBFAMILY pompilinaepompilidaePOMPI LINAE euphorbiae viereck alamosaalamona bent boulder genus aporus conejos crowley Huerhuerfanofano larimer apolus spinola montezuma otero saguache and weld members of this genus prey on trapdoortrap door counties spiders ctenizidae using the spider s burrow fulgifrons cresson larimer co fort collins as a nest languid cresson co a single female larimer luxus banks montezuma co several males taken 8 km NE oflivermoreof livermore taken in a malaise trap at arriola by T mintaka brimley bent crowley delta huer marquardt this is primarily a species of fano prowers and weld larimer counties the western coastal states neglectsnegneglectalecta banks boulder and larimer counties genus psorthaspis banks placita sonorensissonorensis townes delta co is this this is another genus of predators on trap- an unusual range extension for this species door spiders the species show strong sexual A single female was collected by UN dimorphism the females of our species being lanham 5 mi N of 30 june 1938 delta patterned with orange similar to many mutil- reynoldreyreynoldireynoldsireynoldsnsi banks bent and animas coun- fenfeynoldi las lidae while the males are all black a presumed ties example of mullerian mimicry rufescentrufescensrufescens banks boulder and larimer counties nigricepsnigriceps banks montrose co cravanuravan A semitincta banks bent boulder Huerhuerfanofano species of the deep southwest larimer mesa montezuma and otero sansanguineasanguinedsanguinedoguineaguined smith baca and otero counties counties genus Evageevagetestes Lepeletier genus Phanphanageniaagenia banks members of this genus are nest parasites of this is a small genus of wasps in which the other pompilidae females seek out and dig females utilize spiders of several kinds and into nests of other spider wasps destroy the place their paralyzed prey in oval mud cells egg and lay an egg of their own on the spider built beneath stones or logs asignus dreisbach alamosaalamona bent Huerhuerfanofano bombycina cresson boulder larimer and larimer prowers weld and yuma coun- montezuma counties ties calefactuscalefactus evans larimer co A single female genus auplopusAup lopus spinola of this evidently rare species was taken by the author 10 km E of PO on 9 this is a large genus of worldwide distribu- livermore august 1996 on foliage of helianthus tion the females make mud cells in protected annausannuus is the first colorado record places by carrying water from pools or pud- this of a species previously from dles and making pellets from dry soil the pel- known california lets are carried in the mouthpartsmouthparts and applied arizona texas and questionably montana to the nest cells by use of the tip of the crassicrassicomiscomis conconsimilissimilis banks boulder abdomen as a trowel spiders of diverse groups larimer mesa mineral park and are used as prey teller counties crassicorniscrassicomiscrassicomiscornis crassicomiscrassicorniscrassicomiscornis shuckardShucschuckardkard larimer architectarchitectsarchitectusus architectarchitectsarchitectusus say arapahoe boul- co fort collins der delta fremont jefferson larimer hyacinthinushyacinthinus cresson alamosaalamona baca boul- montezuma and weld counties der conejos gilpin Huerhuerfanofano jefferson caerulescens subcortical is walsh boulder larimer mesa otero and weld counties co ingenuusingeningenuitsingenuityuitsuttsuus cresson alamosaalamona bent el paso mellipesmelligesmellipes variitarsatusvariffarsatus dalla torre larimer larimer morgan otero prowers and co fort collins yuma counties 194 GREAT BASIN naturalist volume 57 mohave banks alamosaalamona baca bent boulder oregon evans grand and larimer counties cheyenne Huerhuerfanofano larimer moffat quinquenotatus hudi evans alamosaalamona conejos montezuma and weld counties Huerhuerfanofano larimer mesa park saguache padpadrinusrinus padpadrinusrinus viereck bent boulder and weld counties delta jefferson larimer mesa moffat quinquenotatus quinquenotatus say alamosaalamona montezuma routt washington and weld boulder el paso larimer saguache and counties weld counties carvusparvus cresson baca bent boulder delta slowisnowi viereck baca bent boulder crowley douglas eagle el paso jackson jefferson delta el paso Huerhuerfanofano kiowa larimer larimer moffat montezuma morgan las animas mesa montezuma prowers otero routt washington and weld coun- and weld counties ties genus poecilopompilus howard subangulatus banks boulder Huerhuerfanofano jefferson larimer mesa moffat montezuma these are moderate sized wasps that are park pueblo and weld counties frequently banded with black and yellow polistes genus much like social wasps of the genus agenioideus ashmead like members of the preceding genus they these are small wasps of delicate build are specialists on orb weaving spiders and nesting so far as known from inside crevices nest in simple burrows in the soil in the soil algidusaegidus coquilletticoquillcoquilletteetti provancher mesa co col- biedbiedermanibiedermannermani banks bent larimer and weld orado national monument counties algidusaegidus willistoniwillistoni patton alamosaalamona baca bent birkbirkmannimanni banks larimer mesa and otero crowley larimer las animas monte- counties zuma prowers and weld counties humilis cresson alamosaalamona boulder el paso interruptintermptusinterruptusus interruptinterruptusus say baca bent kiowa and larimer counties this is a predator las animas montezuma otero prowers on orb weaving spiders often around pueblo and weld counties buildings genus tachypompilus ashmead genus Sericosericopompiluspompilus howard these are large wasps of mostly reddish this is a small genus of slender wasps that brown coloration they prey largely on wolf prey on diverse spiders and nest in the soil spiders lycosa and dolomedes and place usually from the side of a rodent burrow or the their paralyzed prey in shallow burrows in burrow of another wasp sandy or powdery soil often close to buildings gustatusangustatusan cresson bent boulder el paso prowers weld and yuma counties ferrugiferruginousneus ferrugiferrugineusferruginousneus say arapahoe boul- apiapicaliscalis say baca bent and prowers coun- der larimer and jefferson counties ties unicolor carinuscerinus evans bent crowley and neotropicneotropicalisalis cameron otero co hawley otero counties unicolor unicolor banks co km genus larimer 32 episyronEpisyron schiodteschiodtz N fort collins this western subspecies these small wasps prey exclusively on orb has also been taken twice in wyoming weaving spiders araneidaearaneidanAraneidae which they flush green river and 30 miles NE ofoflaramielaramie from their webs and to a simple sting carry genus nest dug in the soil so rapid are their actions anopliusAnoplius dufour that george and elizabeth peckham 1898 subgenus notiochares banks called I1 species the tornado wasp this is a small subgenus of mostly black or biguttatus biguttatus fabricius baca bent bluish wasps of moderate size it is primarily a larimer and otero counties neotropical group with I1 species that is occa- biguttatus californicuscaliforcalif nicus banks alamosaalamona chaf- siosionallynally taken in colorado the prey consists fee huerfanoHuerfano larimer and montezuma of wolf spiders that are most often taken in tall counties grasses in wet places 199711997 COLORADO SPIDER WASPS 195 lepidus atramentariusatramentarius dahlbom baca lari- variety of other ground dwelling spiders are mer and prowers counties also used fifteen species are reported from colorado subgenus lophopompilus radoszkowskiRadoszkowski brevihirtabrevihirta banks alamosaalamona bent el paso prowers saguache and weld counties this is another small subgenus of rather an inhabitant of sand dunes large wasps in this case holarctic in distribu- clysteraclystera banks archuleta baca bent boulder tion relatively large spiders of several fami- crowley delta gilpin gunnison kiowa lies are used as prey nests are dug from flat larimer mesa and otero counties soil or from the sides of preexisting holes cylindriccylindricusus cresson prowers co carlton aethiops cresson arapahoe archuleta bent estellina banks delta and larimer counties boulder conejos crowley denver fre- frafraternusternus banks kiowa and logan counties mont Huerhuerfanofano jefferson larimer las insinsolentinsolensolens banks alamosaalamona bent boulder cone- animas montrose otero saguache weld jos delta gunnison Huerhuerfanofano larimer and yuma counties mesa moffat morgan otero prowers pueblo atrolatrox dahlbom boulder and larimer coun- washington and weld counties ties leona cameron larimer co 8 10 iniknikm E of cleora banks alamosaalamona baca delta mesa livermore PO 3 females taken 4 5 august moffat prowers weld and yuma coun- 1994 on euphorbia marginatemarginatamarginata and a male ties largely confined to the vicinity of taken 20 august 1996 on foliage of heli- streams and lakes anthus annausannuus first colorado record of a species described from mexico and re- subgenus arachnophroctonus ported from texas new mexico and ari- howard zona marginatusmargimarginattisnattisnatus say cheyenne costilla paso species of this subgenus are inhabitants of el kiowa larimer moffat morgan sedg- open country where the soil is reasonably fri- wick weld and yuma counties able simple nests are dug in the soil and pro- percituspercitus evans boulder douglas primarily mercitus bent eagle visioned with wolf spiders lycosi- jefferson morgan prowers dae larimer and counties acapulcoensis cameron baca bent chey- rectangularis rectangularrectangularisis dreisbach larimer enne otero prowers and pueblo counties and yuma counties americanusamericanosamericanus ambiambiguusambiguousguus dahlbom mesa co splendentsplendens dreisbach bent costilla el paso nigritus dahlbom alamosaalamona baca bent larimer lincoln logan morgan otero boulder cheyenne conejos costilla prowers and weld counties crowley delta denver douglas elbert subcylindricsubcylindricusus banks arapahoe archuleta el paso garfield Huerhuerfanofano larimer las baca bent boulder cheyenne delta animas logan mesa morgan otero Huerhuerfanofano larimer lincoln otero and prowers weld and yuma counties this phillips counties species has been called relarelativesrelativustivus fox for subtruncatus dreisbach bent elbert lari- many years mer otero and weld counties semicinctus dahlbom alamosaalamona baca bent tenebrosustenebrosus cresson boulder clear creek conejos costilla el paso Huerhuerfanofano douglas eagle gilpin jackson jefferson kiowa las animas morgan otero prow- larimer mesa mineral moffat monte- ers weld and yuma counties this species zuma montrose morgan routt and teller has been called marmarginaliamarginalisginalis banks for counties many years truntruncatestruncatuscatus dreisbach bent larimer prowers semirufmsemirufussemirufus cresson baca bent larimer mesa and weld counties and weld counties subgenus anopliusAnoplius dufour subgenus Pompipompilinuslinus ashmead these are small black or bluish wasps that this subgenus is closely similar to the pre- nest primarily in niches such as under stones ceding but the species are smaller in size or logs prey consists of various ground wolf spiders are the most common prey but a dwelling spiders 196 GREAT BASIN naturalist volume 57

dreisbachidreisbachi evans archuleta boulder elbert paraulaparvula banks aiaAlaalamosaalamonaalamosalmosal boulder Huerhuerfanofano el paso fremont Huerhuerfanofano larimer jackson larimer and teller counties mesa morgan otero sedgwick and silvisilvivagavaga evans alamosaalamona boulder and lari- weld counties mer counties hispidulushispidhispidulousulus dreisbach elbert co running creek field station 2120 in subgenus arachnospila kincaid illinoensis robertson boulder delta garfield these are black wasps averaging larger than larimer and weld counties those of the preceding subgenus females prey imimbellisbellis banks boulder jackson larimer mesa upon various ground dwelling spiders and and mineral counties make simple nests in friable soil ithaca banks larimer co nests along streams building short galleries among stones arataarcta cresson alamosaalamona boulder conejos nigerrimus scopoli larimer co stevens denver el paso grand gunnison hins- gulch 22 km NW fort collins 2200 in dale Huerhuerfanofano larimer mesa moffat papago banks bent garfield kiowa and otero montrose park san miguel and teller counties counties tolucatoluba cameron bent and larimer counties fumifumipennisjumipennispennis eureka banks alamosaalamona arapahoe virginiensis cresson recorded from colo- boulder gilpin jefferson larimer ouray rado without further data by evans 1951 and routt counties scelesta cresson alamosaalamona arapahoe genus evans bent hesperopompilus boulder chaffee conejos delta denver this is a small genus of rare little studied eagle el paso gilpin Huerhuerfanofano jackson species most of them confined to the south- jefferson kiowa larimer las animas west or mexico lincoln mesa moffat otero park saguache teller weld and yuma orophilus evans fremont and larimer coun- counties ties subgenus anoplochares banks genus arachnospila kincaid nothing is known of the behavior of the single subgenus ammosphexAmmosphex wilcke species of this subgenus occurring in colorado A european species is known to these are small wasps more often taken in attack burrowing wolf spiders using the spi- wooded areas than on the plains the prey der s burrow as a nest consists primarily of small wolf spiders this provancher and the following 2 subgenerasubgenera are holarctic in alicataapicata bent boulder garfield distribution gilpin jefferson larimer and weld counties gularisanangularis angularis banks boulder delta jefferson larimer las animas mesa teller genus aporinellus banks and weld counties these are minute wasps all species exten- anomala anomala costilla dreisbach bent sively patterned with pale pubescence records douglas larimer mesa montezuma mor- indicate that jumping spiders salticidae are gan and pueblo counties the usual burrows dakota prey nests are simple in dreisbach boulder larimer and the ground teller counties imbecillaimbecilla imbecillaimbecilla banks chaffee lake lari- basalisbabalis banks alamosaalamona bent delta jackson mer and teller counties larimer otero and weld counties luctuosa cresson boulder conejos custer comcompletuscomplexuspletus banks bent chaffee delta huer lake larimer montezuma park and fano jackson larimer mesa montezuma summit counties washington and weld counties michiganensis dreisbach grand jefferson mediansmemedianusdianus banks alamosaalamona bent delta jeffer- and larimer counties son larimer teller and weld counties occidentoccidentalisoccidentalistalis dreisbach adams boulder cos- taeniolatus rufus banks baca bent boulder tilla denver elbert el paso larimer larimer teller and weld counties proba- mesa mineral park rio blanco and bly no more than a color form of the fol- weld counties lowing 199719971 COLORADO SPIDER WASPS 197

taeniolatus taeniolatus dalla torre bent boul- robinsorobinsoniinii stigmaticastigstigmaticalmatica banks larimer co taken der delta larimer and pueblo counties at the same locality as the preceding and unionsunionis dalla torre bent boulder delta doubtless no more than a color form douglas Huerhuerfanofano jefferson larimer rugatarogata townes bent delta huerfanoHuerfano larimer mesa moffat montezuma otero and montezuma otero and weld counties weld counties for many years this species was caedfasciatuscalledfasciatus smith literature CITED yucatanensis cameron bent boulder delta douglas Huerhuerfanofano larimer mesa moffat DAY MCM C 1977 notes on some pompilidae hymenop- montezuma otero and weld counties tera of incorrectly reported type locality entomolo fistsgists monthly magazine 11271112 71 74 genus paracyphononyx gribodoghibodo 1981 A revision of pompilus fabricius hyme- nopteranop tera pompilidae with further nomenclatorial these large black wasps do not build nests and biological considerations bulletin of the british but attack wolf spiders in their burrows the museum natural history entomology 42142 1 42 EVANS HEH E 1950 51 A taxonomic study of the nearctic wasp larva developing on the spider as a para- spider wasps belonging to the tribe pompilliPomPomppompilmipompilinipiliniilmiliml hyme- sitoid noptera pompilidae transactions of the american nereus lepeletierlepelletierLe baca boulder entomological society 7513375 133 270 7620776 207 361 fufunereus peletier bent 7720377 203 340 paso fano delta el Huerhuerfano larimer las EVANS HEH E AND CMC M YOSHIMOTO 1962 the ecology animas mesa montezuma otero and and nesting behavior of the pompilidae hymenoptera prowers counties of the northeastern united states miscellaneous publications of the entomological society of ameiamelamer SUBFAMILY ceropalinae ica 3673 67 119 HURD PD JR 1952 revision of the nearctic species of genus ceropalesCeropales latreille the pompilid genus pepsis hymenoptera pompilidae bulletin of the american museum of natural members of this genus are parasites of other history 9826198 261 334 pompilidae females are attracted to nesting KASTON BJB J 1959 notes on pompilid wasps that do not females then slip in and lay an egg in the dig burrows to bury their spider prey bulletin of the book lungs of the spider the Ceroceropalespales egg brooklyn entomological society 5410354 103 113 KROMBEIN KV HURD SMITH AND B hatches before that of the host and the larva FDPD JR DR BDD BURKS 1979 catalog of hymenoptera in america consumes the host egg and then the spider north of mexico pages 1523 1570 in volume 2 brevibrevicorniscornis patton bent boulder el paso aculeata smithsonianSmith soman institution washington DC prowers MENKE ASA S 1990 nomenclature of north american kiowa larimer otero and weld pompilidae sphecos 2018901820 18 19 counties PECKHAM GWG W AND EGE G PECKHAM 1898 on the instincts elegansdelegans aquilonioaquilonia townes larimer and weld and habits of the solitary wasps wisconsin geological counties and natural history survey bulletin 2 245 appp SHIMIZU A 1994 phylogeny and classification of the fam- elegansdelegans elegansdelegans cresson prow- bent larimer ily pompilidae hymenoptera tokyo metropolitan ers I1 and weld coulcounties university bulletin of natural history 212 1 142 maculatamaculatafraternafratenafratema smith boulder grand huer TOWNES HKH K 1957 nearctic spider wasps of the subfamsubram fano jackson jefferson LalaplatalaplacaPlata larimer liles Peppepsinaepepsinatepepsmaesinae and ceropalinae bulletin of the united and montezuma counties states national museum 2091209.1209 1 286 cresson paso WILLIAMS FX 1956 life history studies of pepsis and nigripes baca bent crowley el Hemihemipepsispepsis wasps in california hymenoptera pom- kiowa larimer las animas logan mor- pilidaepil idae annals of the entomological society of gan otero and prowers counties america 4944749 447 466 robinsorobinsoniinii robinsorobinsoniinii cresson larimer co 21 received 3 1997 km W 2300 m january livermore accepted 17april17 april 1997 great basin naturalist 573 C 1997 appp 198 208 GROWTH AND survivorship OF FREMONT cottonwood GOODDING WILLOW AND SALT CEDAR SEEDLINGS AFTER LARGE FLOODS IN CENTRAL ARIZONA

JC StromStrombergstromberglstromberg1stromberghbergibergl1

ABSTRACFABSTRACT during winter 1993 arizona experienced regional river flooding floodwatersFloodwaters at the hassayampaHassayampa river eroded floodfloodplainsplains and created a 50 m wide scour zone available for colonization by pioneer plant species the slow rate and long duration of the floodwater recession allowed establishment of spring germinating native trees mainly fremont cottonwood populus fremonfremontnfremontiifremontiatiitiltn and goodding willow mahxsahxsalix gooddingii as well as summer germinating species including the introduced salt cedar tamarix chinenchinensissis and related species goodding willow and fremont cottonwood seedlings showed zonation in the floodplainfloodplain while salt cedar was equally abundant in zones with saturated and dry surface soils floodplainFloodplain elevation and soil moisture influenced shoot growth rate to different degrees among the 3 species for example goodding willow seedlings were significantly taller in areas with saturated soils than dry surface soils fremont cottoncottonwoodswoods were taller in the dry surface soil areas and salt cedar were equally short in both soil moisture zones other factors that differentially influenced abundance 01or growth rates included competition with herbaceous species mebmelilotusMehlotusiotus sppapp an introduced plant locally preempted salt cedar establishment and herbivoreherbherbivoryivory selective browsing by livestock at 1 river site reduced the natural height advantage of the native tree species I1 draw on the results of this descriptive field study to suggest ways in which stream flows and floodfloodplainplain land use can be managed to lestorerestore ecological conditions that favor native tree species over the introduced and widespread salt cedar

key words riparian habitats floods populus fremonfremontiifremontnfremontiatiitiltn Salixallxsalix gooddingii tamarix chmensiscbinensis

fremont cottonwood populusfremontiipopulus fremonfremontiifremontiapremontiitil and by fremont cottonwood goodding willow and goodding willow salix gooddingii are pioneer herbaceous vegetation stromberg et al 1993 species that establish episodically after winter larger floods of longer duration in contrast spring flood flows along rivers of the desert have been observed to facilitate establishment southwest stromberg et al 1991 everitt 1995 of various species of tamarix stevens and winterspringWinter spring floods scour competing vegeta- waring 1985 ohmart et al 1988 griffin et al tion deposit and rework alluvial sediments and 1989 provide supplemental moisture during the short during winter 1993 arizona experienced period in spring during which the seeds dis- regional river flooding although instantaneous perse and germinate salt cedar tamarix chi peaks were on par with other recent large nensisbensis and related species an invasive riparian floods the 1993 event ranked as one of the shrubby tree native to eurasia also is adapted most severe in state history when collectively to establish after flood disturbance but estab- considering magnitude duration and volume lishes more opportunistically than fremont house 1995 the 1993 floods caused much cottonwood and goodding willow brock 1994 geomorphic and vegetational change includ- salt cedar initiates seed dispersal later in the ing channel widening huckleberry 1994 and season but disperses its seeds over a longer mass wasting of floodfloodplainsplains supporting velvet period of time warren and turner 1975 thus mesquite prosopis velutina woodland and depending in part on timing and duration fremont cottonwood goodding willow forest floods may either preclude or enhance salt the floods also created extensive habitat for cedar establishment for example salt cedar riverine marshland and young stands of cotton- seedlings were scarce after a small spring flood wood and willow stromberg et al 1997 I1 in the hassayampaHassayampa river in 1991 because the undertook this study with the objective of de- narrow band of germination space created by terterminingmining how abundance distribution growth the rapidly receding floodwatersfloodwaters was colonized and survivorship of seedlings and vegetative

I1gentelcentergenter toiforfoifot environmental studies arizona state university tempe AZ 85287321185287 3211

198 199719971 NATIVE TREE survivorship AFTER FLOODING 199 sprouts of fremont cottonwood goodding wil- TABLE 1 density stemskastemshastemsha of stems 2 cm of native and low salt cedar and various shrubs were influ- exotic pioneer species in forest stands at 3 central arizona study riversnversavers enced by the 1993 floods and by hydrologic events in subsequent years goodding fremont and bonplandBonpland salt cottonwood willows cedar STUDY SITES hassayampaHassayampa river 287 412 45 study sites are located on perennial reaches santa maria river 312 416 195 3 sonoran is of central arizona rivers one site date creek 45 60 5 in the arizona nature conservancy s hassa yampa river preserve in northwest maricolamaricopaMaricopa county elevation ca 600 in the hassayampaHassayampa river has a mean annual flow rate of 050.5 msnam3s 1 lishmentlishment of plot markers rebar that were at the morristown gage USGS 9516500 removed or buried during floods floodplamfloodplainFloodplamplainpiam located about 4 km downstream of the pre- topography and changes in plot surface eleva- serve surface sediments are predominantly tion due to sediment deposition and scour were sand surface water has an electrical conduc- determined based on repeat cross sectional tivity of 600 700 uscmusam the area was grazed surveys of the hassayampaHassayampa river transects by cattle to 1987 but the land use at prior main stromberg et al 1997 plots 1 I1 x I1 in were the preserve is now ecotourismecotourism the santa sampled for density and height of woody seed- maria river study site is in the alamo lake lings and vegetative stem sprouts 3 5 times wildlife management area elevation 365 in per year during 1993 1994 and 1995 shoot paz in mohave and la counties and is grazed height was measured for the tallest individual by feral burro and trespass cattle mean annual plot plants assumed dead 1 per species per were flow rate of the santa maria is 2 nainalna1naa at the if absent from plots although live root or stem bagdad gage USGS 9424900 located about fragments may have been dispersed from plots 5 kmkin upstream of the study reach the date in floodwatersfloodwaters the nonparametricnonparametncnon kruskal is state land leased by parametric creek site on arizona wallis test was used to test for significant dif- creek ranch elevation 880 in date date ferencesferences in survivorship of the 1995 flood be- creek is a tributary to the santa maria river tween 1993 seedling cohorts of fremont cotton- and is gaged floodplainfloodplain is not the date creek wood goodding willow and salt cedar spear- grazed by cattle from november to march man rank correlation analysis was used to deter- floodplainsFloodplains of all 3 rivers are vegetated pri- whether seedling survivorship of the 1995 marily by fremont cottonwood goodding wil- mine flood pooled values for all 3 tree low forests velvet mesquite woodlands burro using species varied with distance from the 1993 channel brush Hymenhymenocleaoclea monogyramono gyra scrublandsscrublands and edge SPSS for windows was used for all sta- seep willow baccharis salicifoliasalicifolia stands brown tisticaltical analyses 1982 other woody species include arizona tis all 3 50 randomly 1 X 1 ash fraxinus velutina bonplandBonpland willow salix at riversrisers located in and bonplandianabonplandiand coyote willow salix exigiaexiexiguagua plots were sampled in june october of 1994 1 soil the arrow weed tessoriatessariaTessaria sericea and screwbean within or 2 moisture zones in mesquite prosopis pubescentpubespubescenscens salt cedar had scoured flood channel fifty plots were in a lower density than fremont cottonwood or zone with saturated surface soils or shallow willows at study sites at all 3 rivers table 1 standing water 3 cm and 50 were inm a zone based on sampling of saplings and trees stems with slightly higher surface elevation 10 30 2 cm at breast height in 1994 with the cincm higher and dry or damp surface soils as of point quarter method 20 points per river june 1994 at date creek the saturated soil zone was not extensive and plots were sam- METHODS pled only in the dry surface zone woody seedlings in the plots were sampled for density at the hassayampaHassayampa river data were collected and for height and browse status browsed or at a total of 100 permanent plots distributed unbrowsed of the tallest individual per species along 8 transects established at the preserve in plants were classified as browsed if any of the 1987 plots were located at known distances shoots had been eaten if there were 20 along the transect line allowing for relstabreestab individuals of fremont cottonwood goodding 200 GREAT BASIN naturalist volume 57 willow or salt cedar in the 50 plots stem 250 height and browse status were measured in Z c0ca 200 additional random plots to increase sample E size to 20 individuals per species per zone 41 150 herbaceous cover by species was visually estimated in the plots using gnddedgoddedgridded plot 0 namesframes to reduce sampling error CO one way analysis of ANOVA with a 50 variance C 0 post hoc sheffe s multiple comparison test was C D 0 used to detect statistical difference in stem density n 50 plots and stem height n 20 T individuals per species between fremont cot- ondjfmamjjasondjfmamjjasondjfmamjjN I1 I1 M 1 M I1 1aaa 1fmamjjas0ndjfmamjj tontonwoodwood goodding willow and salt cedar with- 1993 1994 1995 in zones of each riverriven independent sample in riverriven fig 1 mean daily flows in the hassayampahassayarnpaHassayampa river at the student s t tests were used to compare density morristown gage during water years 1993 1994 and n 50 plots and stem height n 20 of each 1995 species between the saturated and dry surface zones of the hassayampaHassayampa and santa maria rivers additionally plots within the dry surface zone at the hassayampaHassayampa river were subse- elevation of the scour zone relative to 1993 quently divided into those with 50 and stromberg et al 1997 the santa maria also 50 cover of sweet clover melilotus albus had large flood peaks instantaneous dis- and M officinaofficinalisofficinahshslisils and densities of salt cedar charges in the santa maria peaked in 1993 at and fremont cottonwood were compared be- 347 msm3s 1 on 8 january 440 msnrsarsursm3s 1 on 9 febru- 1 tween the 2 groups with student s t tests ary and 356 na on 20 february 1993 seedling cohort RESULTS TEMPORAL AND SPATIAL distribution AS surface flow the hassayampaHassayampa river floodwatersfloodwaters receded in 1993 woody plants germinated in exposed the 1993 flood in the hassayampaHassayampa river had moist soils in the following sequence a 25 yr recurrence interval with instantaneous fremont cottonwood march april goodding willow flow rates peaking at 745 1 on 8 january italraira april may salt cedar may september 1993 this was followed by flood peaks of 218 arrow weed july september and seep wil- na 1 on 17 january 328 na 1 on 9 februarfebruaryy low july september fig 2 trace amounts and 439 m3s 1 on 20 february surface flow ms 0010010.01 stemmstemm2stemma2 of bonplandBonpland willow were was above average during and ooi spring summer recorded from date creek data not shown of 1993 and was through august present at the but this species was not present at the has morristowntown gage dry Morns located in a frequently sayampa river site reach of the hassayampaHassayampa river fig 1 flood woody seedlings showed zonation in the waters in 1993 had an estimated depth of 303.03 0 floodplainsfloodplains of the hassayampaHassayampa and santa maria inm on low floodfloodplainfloodplamplamplainpiam terraces and a velocity of rivers table 2 fremont cottonwood was sig- I1 212.12 1 in s floodwatersFloodwaters enlarged the channel nificnificantlyantly more abundant in dry surface zones from about 3 to 50 in wide and caused a net while goodding willow coyote willow and lowering of the floodplainfloodplamfloodplamplainpiam surface throughout arrow weed were significantly more abundant 1994 instantaneous flow rates never exceeded in zones with saturated soils salt cedar and 2 na 1 during 1995 instantaneous flow rates seep willow were equally abundant in both peaked at 566 arsm3smsnrs 1 15 to 20 yr recurrence zones within the dry surface zone at the has interval in february flows remained high in sayampa river however salt cedar seedlings spring and summer of 1995 but diminished varied significantly t test P 0050.05 in density more rapidly than in 1993 surface flow was depending on cover of yellow and white sweet present at the morristownMornstownmown gage through june clover fig 3 salt cedar seedlings were nearly 1995 the 1995 floods deposited sediment on absent in plots with high sweet clover cover large portions of the floodplainfloodplain raising the but averaged 15 stemsm2 in plots with little 199719971 NATIVE TREE survivorship AFTER FLOODING 201

140 at the end of the 1993 growing season POFR november fremont cottoncottonwoodwood seedlings 120 1 had a mean density of 20 stemsm2 in the has SAGO sayampa river flood channel as a whole fol- loo100 TACH lowed by salt cedar 13 stems and goodding 13 willow 9 stems values had declined to about BASA E 4 stemsm2 for each 1994 80 species by october TESE the 1995 floods caused high mortality of these remaining 1993 seedling cohorts in hassa Z 60 yampa river study plots fremont cottonwood and goodding willow seedlings had 96 40- mortality and salt cedar had 100 mortality between species differences in survivorship 20 were not statistically significant at P 0050.05oos however there was a significant correlation 01 r 0500.50oso P 0050.05 n 28 between woody 0 6 12 181 8 24 30 36 seedling survivorship and distance of the plot months past jan 119931 1993 from the edge of the channel as of 1993 fremont cottoncottonwoodswoods and goodding willows that fig 2 density of woody seedlings established after floods in 1993 and 1995 in the hassayampahassayarnpaHassayampa river POFR did survive were mainly in narrow bands along populus fremonbremonfremontiifremontnfremontiatiitiltn SAGO salixsahysaby gooddinggooddingiigooddingnii TACH the edge of the scour zone with some surviv- tamany tamarix chinenchinensischmensissis BASA baccharisbocBacchenschans salicisahcifohasalicifoliafolianoila TESE ing despite deposition of up to 1 in of sediment tessenaTessariasafia sericea GROWTH RATES AND BROWSE RATES on average in the hassayampaHassayampa river floodplainfloodplain seedlings of native tree species goodding willow and fremont cottonwood were taller than or no sweet clover fremont cottonwood seed- salt cedar and native shrub species seep willow lings were equally dense in plots with low and and arrow weed fig 5 among tree species high values for sweet clover cover yellow and goodding willows were significantly taller than white sweet clover are introduced bienbiennialsnials fremont cottonwood and salt cedar during their and had high cover in 1993 and 1994 in the ist and and2nd growing seasons in the saturated dry surface zone of the hassayampaHassayampa river soil zones at the hassayampaHassayampa and santa maria preserve rivers fig 6 in dry surface zones at river sites ABUNDANCE AND SURVIVORSsurvivorshipHIP at the the 3 species did not differ significantly in hassayampaHassayampa river fremont cottonwood was height after I1 growing season after 2 growing the most abundant species in the dry soil zone seasons native tree species were significantly after I1 growing season and goodding willow taller than salt cedar in the dry surface zone at was most abundant in the saturated soil zone the hassayampaHassayampa and santa maria rivers but fig 4 after 2 growing seasons fremont cot- not at date creek date creek was the only tonwood and salt cedar seedlings had declined study site with high rates of browse by live- to approximately equal densities in the dry soil stock with 89 of the fremont cottonwood zone as had gooddinggooddmg willow and salt cedar seedlings and 5 of the salt cedar seedlings in the saturated soil zone at the santa maria classified as browsed as ofjuneof june 1994 in most river fremont cottonwood and gooddinggooddmg wil- cases browsing resulted in loss of the terminal low seedlings showed trends of being more shoot browse rates were 5 per species at abundant in the dry surface and saturated soil the other 2 rivers zones respectively salt cedar seedlings how- fremont cottoncottonwoodswoods were significantly taller ever were significantly more abundant than in the dry surface zone than in the saturated either native tree species in both zones at the soil zones at both the hassayampaHassayampa and santa santa maria river after a single growing season maria rivers after I1 and 2 growing seasons and densities did not differ significantly between table 3 in contrast goodding willows tended species after 2 growing seasons seedling den- to be taller in the saturated soil zone but dif- sities did not differ significantly between species ferenferencesces between zones were significant only at at date creek the hassayampaHassayampa river during the ist growing 202 GREAT BASIN naturalist volume 57

a 1 0.0 0.0 saria X 000 000 000 mamw010.1 tessoriatessaria standard 0 0 0 0 1a maerSAEXy 1 1 1 1 Tesrue a m 1 t ei VI 000 0 000.00 0 000 00ee0 001

BASA almasmeans D 000.00 01 0000.0 06 000 s W 000 0 000 0 0 2 ciTESE 1 1 1 1 1 aremol M folia s CO OS H 000.00 03 000 19 000 salicifoliaa 0 0 0 0 salici j shown 0 s0 0.1 5 S 001 001ea 010.10 010 001 BASA ci 1 1 1 1 1 U values K baccharis

1 1 C CO CO 11 i 1 19940 CQ a r D r i 04t 03 03 01 tamarixthmarix in ri 04 moisture 7 5 6 4 a 1 1 1 1 1 S TACH 1 T 0 e aqcq m 0 0 0 0 S 1- ro co in i in zone f s 53 45 74i 71t- 45 S S in f october CO andmrm .0 1 1.7 1.4tf ard by 0 TACH 17 26 14 24 030.3 S S a ln C lbzw 003tm s TACH 1 1 1 1 1 seasonsb0 moisture 0 S 00 0 r l tl 00 S DC 78 mm969.6 515.1rl 74 08 SQ B wms 0 96 51mlIL SAGO 11 siswlm ce t oi 0 0 11 12 2 ajobjo 1 1 1 SAGO0 1 by 1creek chinen X growing 125c 149 116S s N 1 M ii cl 020.2 mg575.7 000.00 121.2 000 U u U 02em 57 00 lbizm1 1 DC 0 0 10 0 0 jg Q SAGO 1 1 1 1 1 gooddingi zaledatezataodd f in to Q twoamz tamarixthmarix9 lf 04 000 26 000 si t 18500 CO creek 0 r 1 0 0 i 1 S boandmcm U M 3 rt 800oo 6to 800oo 800oo in 1 1 5 X X 1 1 1 salix POFR 1 1 3 h t oo i i i i zeledatezela 44 04 07 01 04 0 00 rir 67 in rt t SMS pi co128 71i 141fi D 55 SAGOgl 0 0 0 0 cac2 in boarbolmPOFRfc 1 1 1 1 1

ca 1 c2 T I i i andarm 0 13 S ard pi rl 11 252.5 100 POFRfc 2 10210.2 i 01 i 102 85c i river allowserrors ii 01 0 0 2 sseSs e s SSM tiialm 333 goodding v s 2 in h- d in to xariamahiamaria 5 7 5 6 000 000 000.00 000 1 1 1 1 1 00 TACH 0 x 0 standardtat3 I1 U 0 0 0 0 N COI CO T t maem 1 1 1 col 0 SAEXM 1 ff C ao&o river fremon in52 53in 400 41 47t pa CO t p2 CO a 00ee0.00 000 00ee0.00 03 santa1 salixS 3 0 0 0 0 ca e c3 mariag seasonaseamona0 populus it s IS 11. meansJ almaim 11 r i r- HR U .2 000.00 01 010.1ri 07 00o POFR 2 M 00 el01 s co 0 00oo 00oo CT santa 0 bc 0 8 9 8 8 9 VI 0 0 0 0 1 1 1 1 1 1 1 1 heoeTESE 1 1 ia mago C SAGO W of are 21 CO i M growing 77 66 000 03 010.1 52 000 V 103weewem 60 80 tiiugad 0 0 el010 in 0 preserve 0 w cohorts ab&b HR S shown5 i E p1pa ai fremon 8 caaonecaoc i i aqcq 001 1 0.4 S M t- ea010.1 04 04 070.7 02 bo 0 c 01 04 eb07 river CO 0 0 0 0 0 s S s values preserve BASA 1 1 1 1 1 s seedling 0 0 01 0 0 D CO CO 3 o in P S M OS LI IH 9 6 6 5 6 m 1 1 1 1 0 oi POFR 1 at populus 121.2 09 13 13 06 bolz 1 ablm 1 l sl CL eigpig i1ia ooo 19945 0 0 w oo i 1 y 00 in in ID Q CTS 1 zones 2 i78 45 600 35co 46 i T 1 to nin i river seasons 1993 hassayarnpa11 s S U of tt in 1 co 0 llrt juneC 444.4 75 59ai 47 03 hwwh tat1 W os t 10 cl moisture jaj2 1 3 a f 0 te oj 1 1 1 1 1 for hf a TACH yampa cohortst E f 00 CO 10 oi 1141 leq108zem pherwe 09 growing 343i 6662761- amoS hassayampa laothe 11 1 0 CO M 0 heightboaboznoz bo3 0 between M w 1994 C Hassaa at CO HRrt SM X seedling ffi i 2 ffiafi 1994 D i i afiU 4 06 57 020.2ci 05in 01 stemsa andaxeaam HRtf SM DC Q s october thezae ab02 UTACH 0 0 in 0 0 0 a Q difference 1 1 1 K soils soils june SAGO 1 1 se 005 co eiE i 1 s at 1 c1ca i CO CO CO I 1993 in v 32 381 030.3 37 02 3 in OS f CO 00 03 CO sis after surfaceg surface surface 1 0 0 r 01 CO P SAEXM TABLE oj 3 ns S oker gl C w significant at F saturated V saturated1 0 measured of0 g S a bmeasured E i chinen naaezone & & laanthan zones ei S fl 11iS S is CO r t D dry olidryarmbab2 dry 595.90 030.3 06 01rt e 0 Q co Q rt in59mm 03em 06 5 N zoizol nom2noma 0 0 0 0 POFRfc 1 1 1 1 1 exigiaexigua less

moisture 0 CO CO 1 CO PL 12 590 01 23 38338.300 r i in co s CO 0 salix valuesmazzas densityS c t between

mcm0 andC D HR SM cates s g a S rt w w stem 0 HR SM5 DCu 4 cindicatesvindicates1 r B w Q difference soils soils sgCindi hemeTESE lazda u S w S w 2 M 0 0 0 yb surface 1 surface 1 surface ca TABLE 3 3 n 3 OT saturated VI saturated1 VI significant

sericeaseficea1 errors 3 S dry aj dryAb j zlydryib S u Q &0 0 Q 199711997 NATIVE TREE survivorship AFTER FLOODING 203

50 lings were an order of magnitude more abun- high cover dant than vegetative sprouts low cover CIJCM 40 E discussion AND 0 management implications S 30 salt cedar was subdominant to fremont cot-

D tonwood and goodding willow in mature for- 0 stands at the 3 rivers included in this study 0g 20 est rivers in S to maintain or relegate salt cedar to a sub- 1 flows I0 dominant role stream and floodfloodplainplain U0 10 lands need to be managed to produce ecological conditions that favor native species this study identified several factors that influence

0 1 establishment rates of salt cedar vs native tree POFR TACH species FLOOD TIMING MAGNITUDE DURATION AND fig 3 stem density of 1993 seedling cohorts of populus STAGE DECLINE RATE the large magnitude fremonfremontnfremontiifremontiapremontiitiltn POFR and tamarix chinenchinensissis TACH in plots and long duration of the 1993 and 1995 winter with 50 cover of Mehmelilotuslotusiotus n 32 mean species fluvial cover of 76 7 and 50 n 18 mean cover or 5 1 floods created extensive surfaces avail- as of june 1994 values shown are means and standard able for colonization by woody pioneer plant error bars species that do not tolerate competitive herbaceous cover the long slow decline in the river stage flow rates did not decline to base levels season salt cedar height did not differ signifi- until several months after the flood peaks cantly between zones allowed germination of woody species with a variety of temporal regeneration niches exten- 1995 seedling cohort sive scour prevented herbaceous species from rapidly colonizing the sediments that seedlings of all 3 tree species again germi- moist exposed throughout the and thus nated in 1995 in the moist scoured zone ex- were summer posed by the slowly receding floodfloodwaterswaters in germination sites were available for springspangspnng the hassayampaHassayampa river floodplainfloodplamfloodplamplainpiam densities of germinatinggermmatmg native trees fremont cottonwood the 1995 seedling cohorts ranged among the 3 and goodding willow as well as for salt cedar peaks species from I1 to 4 stemsm2 as of october 1995 a species with biseasonal seed dispersal compared to 9 20 stemsm2 for the 1993 cohorts in june july and august september horton as of november 1993 dry surface soils were 1957 1977 warren and turner 1975 relatively more abundant and saturated soils stream flows on regulated rivers are being less abundant in 1995 than 1993 partly due to naturalized min several ways to facilitate estab- floodplainfloodplain aggradation in 1995 fremont cotton- lishmentlishment of fremont cottonwood and other wood seedlings were the most abundant of the native species stanford et al 1996 poff et al 3 tree species in 1995 and outnumbered salt 1997 the same could be done to reduce estab- cedar by a ratio of about 212 1 as of october 1995 lishmentlishment of salt cedar scouring flows should seedlings of all 3 species tended to have shorter be released during early spring in potential re- stems in 1995 than in 1993 but the difference cruitmentcruitment years for cottoncottonwoodswoods and willows between years was significant at FP 0050.05oos0 05 only the flood flows need to be of sufficiently high for goodding willow fig 7 magnitude and duration to rework sediments densities of vegetative sprouts were greater and create bare surfaces lying within about I1 m in 1995 than during 1993 there were 0250.250 25 of the base flow alluvial groundwater table 0360.360 36 and 0020.020 02 stemsm2 for fremont cotton- stromberg et al 1991 1993 flows should wood goodding willow and salt cedar respec- peak prior to fremont cottonwood and good- tively as of october 1995 compared to ooi0010010.010 01 ding willow spring seed dispersal periods from stems per species in november 1993 many of about february to april depending on eleva- the 1995 sprouts originated from flood pros tion river stage should decline during the tratedgrated 1993 seedlings in 1995 and 1993 seed short month long periods of cottonwood and 204 GREAT BASIN naturalist volume 57

dry soil surface dry soil surface

JU50 POFR 40 SAGO ct E 1 WE TACH 0g 30 I i S 1 1 to 20 1 1 0Q Q 1 10 11 11 1 ea v as ll1 0 ll1 1 HR SM DC HR SM DC

age 1 yr age 2 yr

saturated soils saturatedsaturatedaurated soils

50 i

r 40

30 Cg0 20 I1 & I1 10 L sikgli iasgiasarask i 0 T i i HR SM DC HR SM DC

age 1 yr age 2 yr

fig 4 stem density of 1993 cohorts of populus fremonfremontiifrefremontiafremontzibremontzitiitil POFR salix gooddinggooddmgugooddingiiii SAGO and tamant chinenchinensischtnenszssis TACH in dry surface and saturated soil zones after I1 and 2 growing seasons HR hassayampaHassayampa river SM santa maria river DC date creek values shown are means and standard error bars asterisk indicates that a species had significantly different density from the other species at P 0050oos05

200 fig 5 shoot heights of woody seedlings established 180 POFR afterafretarretaeter the 1993 flood POFR populusfremontiipopulus fremontzz SAGO 160 salix gooddingitii TACH tamanttamarixtamariz chinensissis BASA SAG chinen baccharisBacchans salicisahcifoliasalicifolia TESE tessadatessariatessanaTesrcssariasarla sericea 140 folia TACH 120 BASA loo100

80 0 0

20 B

0- 1 1 0 3 6 4 1 2 1 5 1 8 21 24 months past jan 119931 1993 199719971 NATIVE TREE survivorship AFTER FLOODING 205

dry soil surface dry soil surface

200koocouZUU j PCPOFR 175 EM 3fr EM sSAGOGO E 150 u 19 TACHT CH 125 0 c5u 100 E 1 75 B I1I c A i W 50 1 i 1 6 25 i 1 1 6 1 I1I1 0 T T T T i i HR SMc DC HRR SM DC

ageage 1 yr ageage 2 yr

saturatedsatu raitedcited soilssoisol S saturatedsatursatunatedabed soils 200uu 175 E 150 i 125 100 E 75 1 n3 50 1 1 1 i 25 i 1 1 1 n 11 0 T HR SM DC HR SM DC

age 1 yr age 2 yr

fig 6 stem height of 1993 cohorts of populus fremonfremontiifremontiatii POFR salix gooddinggooddingiigooddinguii SAGO and Tainfamarintainarixfamarivarix chinenchinensischinen&fsis JACHTACH in dry surface and saturated soil zones aftelafterahneranner 1 and 2 gigrowingowing seasons HR ilassayampahassayampaHassayampa rivetriverriven SM santa mariamarlamana river DC date creek values shown aiealeare means and standardstandaiddard enoienorerror baisbalsbars asterisk indicates that a species hadllad significantly different stem height hornfrombormbomm the other species at P 005oos0 05

willow seed viability to expose moist germina- natingbating native pioneer species such ascis seep wil- tion sites mahoney and rood 1993 braatnebraathe low however if initial flood scour is not too et al 1997 the rate of stage decline and of great this species will rapidly regenerate asex- alluvial groundwater decline during early seed- ually stromberg et al 1991 ling growth should not exceed rates of root floods that occur after seed germination also growth of fremont cottonwood and goodding may influence relative abundance of salt cedar willow seedlings these rates are about 1 3 cm vs cottoncottonwoodswoods and willows in this study the per day and vary with soil texture mahoney 1993 seedling cohorts of fremont cottonwood and rood 1992 segelquist et al 1993 to gooddmggoodding willow and salt cedar all had high reduce salt cedar establishment river stage mortality from the 1995 flood and survivor-survivor should stabilize prior to the onset of salt cedar ship did not differ significantly between the 3 germination in late spring this also could species however some studies suggest that salt reduce germination rates for summer germi cedar may be less tolerant of the scouring effects 206 GREAT BASIN naturalist volume 57

100 tiai responses to soil moisture and soil aeration 1993 cohort goodding willow shoot height was greatest in 1995 cohort conditions of saturated soils or standing water 80 whereas fremont cottonwood shoot height was greatest in areas with dry surface soils but shal- E low water tables salt cedar were equally short 60 in both zones comparison between years suggests that salt cedar is as tall as the native tree E species under conditions of reduced water 40 availability yearlingsYearyearningslings of all 3 tree species were somewhat shorter in 1995 than in 1993 most likely due to lower river stage and deeper water 20 tables however the difference between years was greater for native trees other studies dem-

0 onstrateonstrate that salt cedar is more drought toler- POFR SAGOSA 0 TATACHH ant than native cottoncottonwoodswoods and willows and species thus may have a competitive advantage on drier floodplainsfloodplains with deeper groundwater levels fig 7 stem height of 1993 and 1995 cohorts of populus stevens 1987 busch and smith 1995 there ftemontiijrernontii POFR salix gooddingii SAGO and tamarix is a need for additional controlled studies of chinen abterabher 1 chinensischmensissis oachJACHTACH afterarrel I growing season at the has seedling growth siscoe 1993 shafroth et al sayampa river preservePieseiveserve values shown are means and stanstandarddarddaid enerroror bars asterisk indicates signficantsignficautsignificant differ- 1995 to quantify soil moisture ranges over ence P 0050 05 between coholcohortsts which native species have a competitive edge once such ranges are described activities that reduce floodplainfloodplain water availability such as sur- of floods but more tolerant of prolonged inun- face water diversion and groundwater pump- dation than cottonwood and willow trees waring could be managed to produce soil mois- ren and turner 1975 irvine and west 1979 ture levels and groundwater depths that favor everitt 1980 stevens and waring 1985 salt growth and survivorship of native species dur- cedar seedlings may have low survivorship of ing establishment periods floods because of their distribution in the flood GRAZING although this study was not explain because most salt cedar seeds germinate plicitly designed to determine effects of live- later than fremont cottonwood and goodding stock on riparian tree seedlings there were willow seeds salt cedar seedlings sometimes differences in cattle browsing between sites are more abundant on sites close to the chan- which resulted in significant differences in nel where risks of mortality from subsequent seedling heights at the 2 legally ungrazed floods are high stromberg et al 1991 also river sites fremont cottonwood had a large there may be physiological differences between height advantage over salt cedar in certain areas species in tolerance for flood scour or burial of the floodplainfloodplain ie dry surface soil zone at by sediments for example low rates of shoot date creek which is grazedbrowsedgrazed browsed by cattle growth may increase the probability of com- from november to march selective browsing plete burial of salt cedar seedlings existence on fremont cottonwood seedlings caused them of differential survivorship thresholds for fac- to lose their height advantage over salt cedar tors such as shear stress and sediment deposi- between species differences in seedling shoot tion should be experimentally tested if there height ultimately may determine the composi- are differences release of occasional large tion of the stand dominant given that shorter scouring floods in alluvial perennial rivers in plants may have greater mortality due to light the southwest may be a management strategy limitation all 3 species including salt cedar to increase the mortality of salt cedar seedlings appear to be shade intolerant additionally re- and saplings relative to that of native pioneer duction in shoot height may reduce a seed- tree species ling s ability to survive deposition of sediment SOIL MOISTUREM 01 STU RE differences in tree seed- during flood events to favor native tree species ling shoot height between soil moisture zones in on cattle grazed rivers recruitment zones this study were most likely a result of differeddifferen should be protected year round from livestock 199719971 NATIVE TREE survivorship AFTER FLOODING 207 during at least 2 growing seasons to allow systems of the southwestern USU S ecological mono- seedlings to grow above browse height graphsgraphsephs 6534765 347 370 EVERITT BLB L 1980 ecology of saltcedarsalt cedar a pleapledpiedpiea foifolforoor re- PLANT competition local- herbaceous in search environmental geology 3773 77 84 ized areas of the hassayampaHassayampa river floodplainfloodplain 1995 hydrologic factors in regeneration of fre- the exotic herbaceous species yellow and white mont cottonwood along the fremont rivelriverriven utah sweet clover preempted establishment of salt natural and anthropogenic influences in fluvial geo- cedar by germinating in spring and rapidly morphology geophysical monographs 8919789 197 208 GRIFFIN GF DM SMITH SR MORTON GE ALLENALLLN ground surface salt covering the prior to cedar AND KAK A MASTERS 1989 status and implications of germination fremont cottonwood seeds in the invasion of tamarisk tamarix aphyllyaphylla on the contrast germinated prior to sweet clover finkepinke river northern territory australiaaustrallaAusti aha journal of fremont cottonwood seedlings were over environmental management 2929729 297 315 topped by sweet clover during early summer HORTON JSJ S 1957 inflorescence developmentdeveloumentlopment in tamarix penpentandrapentandriatandra pallas tamaricaceae southwestern nat- but they were exposed to full sunlight after uralist 21352 135 139 the midsummer death of sweet clover and had 1977 the development and pelpeiperpetuationpetuationpetuation of the high survivorship this suggests a possible permanent tamarisk type in the phreatophyte zone management strategy for salt cedar at least of the southwest USDA forest serviceselvlee general technical report RM 4312443 124 127 within small areas if native annual species HOUSE PKRK 1995 Hydrohydroclimatologicalclimatological and paleohydiopaleohydro were identified that germinated after or simul- logical context of extreme winter flooding in ari- taneously with native trees but prior to salt zona 1993 arizona geological survey open file cedar it might be feasible to disperse seeds of report 9512959512112112 1 25 huckleberry G 1994 channel to the annual herb into the floodfloodplainplain at an appro- contrasting response floods on the middle gila river arizonaanzona geology priate time after large floods however care 22108392108322 1083 1086 should be taken to insure that such an effort IRVINE JRJ R AND NEN E WEST 1979 riparian tree species would reduce abundance only of salt cedar and distribution and succession along the lower esca- not of desirable summer germinating species lante river utah southwesternSouthwestein naturalist 24 331 346 such as seep willow MAHONEY JMJ M AND SBS B ROOD 1992 response of a hybrid poplar to water table decline in different sub- acknowledgments strates forest ecology and management 5414154 141 156 1993 A model for assessing the effects of altered we thank the arizona nature conservancy rivernver flows on the recruitment of riparian cotton game woods pages 228 232 in B teilmantellmantollman HJ Coitcortnergoitgoltgortnerner date creek ranch and arizona and MGM G wallace LEL F debano and RHR H hamre tech- fish department for allowing access to study nical coordinators riparian management common sites and anonymous reviewers for their help- threads and shared interests USDA forest service ful comments joelle don de ville and leigh general technical report RM 226 assisted with field sampling OHMART RDR D BWB W ANDERSON AND WC HUNTER 1988 hedrick the ecology of the lower colorado river from davis dam to the mexico united states intelinternationalnational literature CITED boundary a community profile united states fish and wildlife service biological report 85719857857.1919 1 BRAATNE JHJ H SBS B ROOD AND PE HEILMAN 1997 life 296 history ecology and conservation ofriparianof riparian cotton POFF NLN L ET AL 1997 the natural flow regime a para- woods in north america pages 57 85 in restetrespetR F stet- digm for river conservation and restoration bio- tler HD bradshaw jr PE heilman and TM science in press hinckley editors biology of populus and its implica- segelquistSECseg ELQUIST CA ML SCOITSCOTT AND CTGT aubleaume 1993 tions for management and conservation national establishment of populus deltdeltoidesoides under simulated research council ottawa canada alluvial groundwater declines american midland BROCK JHJ H 1994 tamarix sppapp salt cedar an invasiveinvasivcinvasivesivc naturalist 130274130 274 285 exotic woody plant in and and semisemiaridarid riparian SHAFROTH PB JMJ M FRIEDMAN AND LSL S ISCHINGER habitats of wewesternt n USA pages 27 44 in LCL C de 1995 effects of salinity on establishment of populus waal LEL E chichildi PM wade and JHJ H brock edi- fremonbremonfremontnfremontiifremontiatiitiltn cottonwood and taniarixtamanatamanxTani arix ramosissima tors ecology and management of invasive riverside saltcedarsaltcedar in southwestern united states great basin plants john wiley and sons ltd west sussex eng- naturalist 555855 58 65 land SISCOE RJ 1993 A comparison of an invasive riparian BROWN DED E 1982 biotic communities of the american shrub species tamarix pentandrapenpentandriatandra with a native southwest united states and mexico desert plants species salix exigiaexigua unpublished master s thesis 414 1 342 new mexico state university las cruces BUSCH DED E AND SDS D SMITH 1995 mechanisms alsocassocassoci-i STANFORD JAJ A JXJ V WARD WJ liss CAC A FRISSELLFRISSLLL RNR N ated with decline of woody species in riparian eco WILLIAMS JAJ A lichatowich AND CCC C couram 208 GREAT BASIN naturalist volume 57

1996 A general protocol foiformoimol restoration of regulated STROMBERG JCJ C DTD T PATTEN AND BDB D RICHTER 1991 rivers regulated rivers research and management flood flows and dynamics of sonoran riparian forests 1239112 391 413 rivers 22212 221 235 STEVENsS revensFEVENS LEL E 1987 the status of ecological research on STROMBERG JCJ C BDB D RICHTER DTD T PATTEN AND LGL G tamarisk tamaricaceae tamarix ramosissimaramosissimd in ari- WOLDEN 1993 response of a sonoran riparian for- zona pages 99 107 in MRM R kunzmann R johnson est to a loyear10 year return flood great basin naturalist and PS bennett technical coordinators tamarisk 5311853 118 130 control in southwestern united states special report WARREN DKD K AND RMR M TURNER 975I19751 salt cedar tamarix 9 cooperative national park resources studies unit chinenchinensissis seed production seedling establishment sievensslevensSILVENSSTEVENs LEL E AND GLG L WARING 1985 the effects of pro- and response to inundation journal of the arizona longed flooding on the riparian plant community in academy of science 10 135 144 grand canyon USDA forest service general technical report RM 438143 81 86 received 6 august 1996 STROMBERGSTHOMBLRG JCJ C J FRY AND D PATTEN 1997 marsh accepted 24 march 1997 development after large floods in an alluvial and land river wetlands 1729217 292 300 great basin naturalist 573 0 1997 appp 209 219 zoogeographic affinities OF THE stonefliesSTONE FLIES plecoptera OF THE RAFT RIVER MOUNTAINS UTAH

richard A Houseman 1 and richard W Baumann2

ABSTRACT we examined faunal affinities of the raft river mountains using stonestonefliesflies plecoptera as indicators this islandlikeisland like mountain range is isolated from other major mountain ranges in the intermountain west by low elevation and regions thirty seven species were recorded from collections made from 19 sites in the raft river mountains cluster analysis demonstrated the raft river mountain stonstoneflyefly assemblage to be most similar to faunalfaunas of the sawtooth and wasatch mountains and quite different from that of the sierra nevada an analysis of the distribution patterns of each species on a family by family basis showed that the raft river mountains fauna consists mostly of species widespread in western north america most families were represented by at least 1 species whose distribution supports faunal affinities with regions to the north and west logistic regression of 6 long distance dispersal factors against stonstoneflyefly presence absence data did not support long distance dispersal as a viable means of colonization for the raft river mountains this suggests that stonstoneflyefly distribution pattpatternseins may be attributed to expansion and subsequent vicariancevicarivicanancevicaanceaneenancenanee of suitable stonstoneflyefly habitats during pleistocene climatic oscillations

key words plecoptera stonestoneliesstostonefliesneliessliesflies zoogeography affinity pleistocene great basin raft river mountains utah

biogeographic properties of islandlikeisland like amoenus alienallenailen yellow pine chipmunk and mountain ranges in the intermountain west citellus belbeldingidingi hall belding ground squirrel have been a topic of much research macarthur both found in the raft river mountains but and wilson 1963 brown 1971 johnson 1975 nowhere else in utah in addition the broader behle 1978 harper et al 1978 wells 1983 distributions of these species extend north and these mountainmountainaln islands are appealing as oppor- west into regions associated with the colum- tunitiestunities to study the distribution diversity bia plateau durrant 1952 and evolution of organisms inhabiting them the damselfly calopteryx aequabilis say the raft river mountains in extreme north- and rugose stag beetle sinodendronSino dendron rugosum western utah are such a mountain range and mann are insects in the raft river mountains various studies have recognized that they are with interesting faunal affinities calopteryx distinct from nearby regions durrant 1952 aequabilis is known to occur throughout most behle 1958 mcmahon and wiebolt 1978 of canada the north central and northeastern mcmahon and wiebolt 1978 used a classi- united states and in isolated pockets in the ficationfication system developed by holdridge 1947 western united states including the columbia and holdridge et al 1971 to divide utah into river drainage provonsha 1975 sinodendronSinodendron life zones based on mean annual tempera- rugosum has an overall distribution outside ture mean annual precipitation and potential utah that includes idaho washington ore- evapotranspiration according to their classifi- gon california and british columbia essig cation the raft river mountains are an eco- 1929 hatch 1971 logically isolated island of subtropical mon- behle 1958 lists 172 bird species and sub- tane moist forest surrounded by regions of species from the raft river mountains and montane steppe and desert mcmahon and concludes that the bird assemblage in the raft wiebolt 1978 river mountains more closely resembles that durrant 1952 concluded that the raft river of the great basin than the nearby wasatch mountains have mammal faunal affinities with mountains the columbia plateau his conclusions were the most desirable organisms to use when based on 2 unique mammal species tamias studying patterns of faunal affinities are those

idepartmentepaitment of zoology brigham young university provo UT 84602 plesentpresent address department of entomology texas a&maam university college station TX 77843 2departmentapaiepai tmentament of zoology and monte L bean life science museum brigham young university provo UT 84602

209 210 GREAT BASBASININ naturalist volume 57 restricted to definable habitats sargent et al an elevation of 2900 m the range has an area 1991 Stonestonefliesflies plecoptera are such a model of approximately 990 km2 and lies in an east group nelson 1994 many stonstoneflyefly species west orientation as part of the northern edge have limited ranges and distinctive differences of the great basin occur between the stonstoneflyefly faunalfaunas of the rocky considerable differences in vegetation and mountains the coast and cascade mountains precipitation exist between the north and south and the sierra nevada jewett 1959 slopes of these mountains due to their east west Stonestonefliesflies depend on a water connection orientation behle 1958 northern slope streams with very specific habitat requirements surdick are unique in utah since they flow into the and gaufin 1978 baumann 1979 to expand snake river drainage system southern drain- their distribution they are greatly influenced ages that once flowed into pluvial lake bonne- by water temperatures baumann 1979 pollu- ville stokes 1987 now drain into the bonne- tion levels surdick and gaufin 1978 and dis- ville basin solved oxygen concentrations gaufin et al the insular nature of this mountain range 1966 rocky streamstreambedsbeds or rocky lakeshoreslake shores makes it excellent for testing biogeographical often are required for nymphs to complete their hypotheses immediately surrounding the raft development gaufin et al 1966 hynes 1976 river mountains are andaridarld lowlands great basin and the riparian environment is important for bonneville basin and snake river plain adult stonestonefliesflies to survive and successfully re- which effectively isolate them from other moun- produce once they emerge from the final nym- tain ranges in the region fig 1 the bonne- phal instar ville basin currently isolates the raft river adult stonestonefliesflies normally fly only short dis- mountains from the wasatch mountains to the tances marden and kramer 1994 and many southeast the snake river plain is a barrier are short winged which further reduces flying for stonstoneflyefly dispersal from the sawtooth moun- ability additionally phenology of adult emer- tains in central idaho nebeker and gaufin gence may have an effect on the distribution 1967 and the great basin isolates the raft of many species for example species in the river mountains from the sierra nevada family capniidae emerge only during winter nebeker and gaufin 1967 brown 1971 john- through breaks in the ice frison 1929 species son 1975 for purposes of this study we con- in this family are generally more limited in sider these 3 mountain ranges as hypothesized their distributions than those of other families source pools for stonstoneflyefly dispersal to the raft nebeker and gaufin 1967 that emerge dur- river mountains ing seasons when habitats and temperatures are more conducive to dispersal METHODS in this study we examine species composi- we visited most major drainages in the tion faunal affinities and long distance dis- raft river mountains at least once during each sea- persal potential of stonestonefliesflies in the raft river son for 2 yr we collected data during 1994 95 we composition mountains compare species and also included previous collection records from 3 regional ranges with that other r mountain from 1977 to 1980 to determine overall similarity between their Stonstoneflyefly nymphs were collected using an stonstoneflyefly faunalfaunas and possible modes of coloniza- aquatic kick net adults were collected with a tion the sawtooth mountains in central idaho beating sheet from riparian vegetation and with wasatch mountains in northern utah and sierra ultraviolet light traps near streams at night nevada in california are regions with which exuviae were also collected the raft river mountains stonstoneflyefly fauna may nymphal and adult stonestonefliesflies were preserved have affinities in 70 ethyl alcohol and identified to the low- est possible taxon using current identification STUDY AREA keys baumann et al 1977 nelson and baumann 1987 1989 stewart and stark 1988 and the raft river mountains near the borders stanger and baumann 1993 of utah idaho and nevada in extreme north- species lists from the sawtooth mountains western utah resemble a gently sloping pla- wasatch mountains and sierra nevada were teau that rises from surrounding andaridarld basins to compiled directly from the literature jewett 199719971 stonefliesSTONE FLIES OF THE RAFT RIVER MOUNTAINS 211

sawtooth mountains

ift river mo5iiiains

fig 1 mountain ranges considered in this study and intervening barriers for stonstoneflyefly dispersal

1960 nebeker and gaufin 1965 gaufin et al hypothesized source pool mountain ranges were 1966 logan and smith 1966 nebeker and determined with cluster analysis using NTSYS gaufin 1966 sheldon and jewett 1967 bau- pc 1701.70 rohlf 1992 stoneflyStonefly presence absence mann and gaufin 1969 newell and minshall data were entered into a matrix from which a 1976 baumann et al 1977 nelson and bau- distance matrix was constructed using jaccard s mann 1987 1989 kondratieff and baumann coefficient the distance matrix which quanti- 1988 stanger and baumann 1993 and stark fies the similarity between a pair of areas as a and nelson 1994 we annotated counties with decimal value was SAHN sequential agglom- any portion of these mountain ranges within erativeerative hierarchical nested clustered using their boundaries and assumed that records from UPGMA unweighted pairs group method with these counties indicated the presence of a arithmetic averaging single link and complete species in that particular mountain range link methods were also employed all cluster- we investigated faunal affinities of each ing methods resulted in dendrograms that species collected in the raft river mountains demonstrate similarities of faunal composition by examining its distribution in north america between mountain ranges each species was classified according to the since SAHN will produce clusters whether region of faunal affinity it supports or not natural groups are present in the data similarities between stonstoneflyefly species com- rohlf 1992 a cophenecophenetictic value matrix was position of the raft river mountains and the 3 computed from the dendrogram matrix to 212 GREAT BASIN naturalist volume 57 analyze goodness of fit between the clustereluster ence absence data in the raft river moun- dendrogramclench ogi am matrix and distance matrix the tains significant correlation would provide cophenecophenetictic matrix was compared element by evidence for long distance dispersal as a mode element with the original distance matrix of colonization for those stonstoneflyefly species found according to a test developed by mantel 1967 in these mountains this comparison produces a product moment WING LENGTH because macropterous correlationcon elation r which measures the degree of long winged stonestonefliesflies are better fliers than relationship between the distance matrix and brachypterous short winged micropterous ogo dcndiogiamdendrogram matrix values 0900900.900 90 indicate that minutely winged or apterous without wings the tree accurately represents natural group- marden and kramer 1994 they were assumed ingsggsmgs plesentpresent in the data values 0700700.700 70 indicate to be better adapted for longiong distance dispersal that natural groups may not be plesentpresent rohlf both sexes were examined for wing morphol- 1992 ogy and scored according to wing length dispersal abilities weiewelewere quantified foifolfor all apterous or micropterous species of eithereithelelthelelther sex stoneflystonefly species recorded flom the hypothe- were given a score of 1 species with bra- sized source pool mountain langesfangesranges scores for chypterous members of either sex were scored dispersal ability weiewelewere based on published data 2 and species where both sexes were macrop- toiloifolfor 6 factors that influence stonstoneflyefly distribu- terous received a score of 3 tion and long distance dispersal logan et al ecological TOLETOLERANCERA NCE species that sur- 1966 nebeker et al 1966 baumann et al 1977 vive in a broader range of ecological condi- surdicksinsun clickcilek and gaufin 1978 baumann 1979 nel- tions were assumed to be better adapted for son and baumann 1987 1989 kondratieffkondiatieffandand long distance dispersal because a broad toler- baumann 1988 stangelstanger and baumann 1993 ance allows a species to survive in a greater season of emergence length of emergence dis- number of post dispersal environments bau- tributiontribution within the souleesource mountain range mann 1979 identifies 3 stonstoneflyefly environments relative distance fornhornfiemflemfrombomm the raft rivelriver mountains cold lotic warm lotic and cold lentic and environmental tolerance and wing length table calls them ecological groupings species that 1 ifcolonizationifcolomation of the raft rivelriver mountains aiealeare limited to only 1 of these ecological group- by stonestonefliesflies occurred viavla landomrandom long dis ings received a score of 1 species in 2 of these tance dispersaldispel salsai one would expect the raft river groupingsgioupmgs were scored 2 and those species mountains fauna to be composed of species capable of living in all 3 ecological groupings with high quantified dispersal ability coleman were given a score of 3 etalet al 1982 SEASON OF EMERGENCE dispersal is moiemolemore we used logistic regression analysis P probable during warmer seasons nebeker and 005 to test toifoifollorfor correlationcon elation between high gaufogaufmgaufin 1967 fall- or winter emerging species scores foifolfor these 6 factors and stonstoneflyefly pres received a score of 1 spring emerging species

TABLE gor falliTAISI i 1 characteristics important foiforfol stoneflystonefly long distance dispersal each characteristic is divided into 3 categories and scored according to assumed influence on long distance dispersal to the raft rivelriver mountains 1 I1 low 2 medimedlmediuminniuntun 3 high

category and score characteristic 1 2 3 season of emergence nrailfallralitat1 or01 winter springspong summetsummer length of emergence 1 3 months 4 5 months 6 12 months ecologicalideological tolerance I1 grouping 2 gigroupingsonpmgs 3 groupingsgi oupmgs wing length apterousaptelouseions 01or brachypterous macropterous micropterous relative distance from raft riverrivel mountains farthestfaifal thestchest 13 middle 13 closest 13 distributiondistiibntion within source mountain i lingetingerange 1 of 3 legionsregions 2 adjacent 2 nonadjacent or01 regions 3 regionslegions 199719971 stonefliesSTONEFLIES OF THE RAFT RIVER MOUNTAINS 213 were scored 2 and summer emerging species are listed inm table 2 by collection site and were scored 3 september through february presence inm each source mountain range of was considered fall and winter spring months these 37 species 5 are unique records for were considered april and may and june utah and have distributions extending outside through august was considered summer the state to the northwest malenka tina LENGTH orOF EMERGENCE we assumed dis- ricker taeniopteryxtaemopteryx nivaksnivalisnavalis fitch capniaacapniacapma persal to be more probable for species whose petila jewett capnura intermontana nelson adults are present for longer periods of time and baumann and doroneuriaDoroneuria sp in addition by using emergence and collection records a potentially undescribed species in the genus we determined how many months during the kogotus was collected in several of the raft year adults are present for each source pool river mountain drainages mountain range species whose adults were literature records contain 82 species in 41 present up to 3 months were given a score of genera and 9 families for the sierra nevada 1 presence during 4 5 months of the year was jewett 1960 nebeker and gaufin 1965 shel- scored 2 and species with adults present 5 don and jewett 1967 nelson and baumann months weiewelewere scored 3 1987 1989 kondratieff and baumann 1988 distribution WITHIN hypothesized SOURCE stanger and baumann 1993 and staikstark and MOUNTAIN RANGERANGESS long distance dispersal nelson 1994 there are 62 species in 32 gen- was assumed to be more probable for species era and 8 families in the wasatch mountains with widespread distributions in the hypothe- nebeker and gaufogatifingaufm 1965 gaufogaufmgatifin et al 1966 sized source mountain ranges to assess how baumann and gaufogaufmgatifin 1969 baumann et al widespread each species was within a source 1977 nelson and baumann 1987 1989 kon- pool mountain range sierra nevada sawtooth dratieffdratieff and baumann 1988 and stanger and wasatch we subdivided each mountain range baumann 1993 the sawtooth mountains have along its length into 3 equally sized regions if 70 species in 33 genera and 9 families nebe- a species was recorded from only I1 of these ker and gaufogaufmgatifin 1965 1966 logan and smith regions we scored it as 1 if a species distri- 1966 baumann et al 1977 nelson and bau- bution covered 2 adjacent regions it was scored mann 1987 1989 kondratieff and baumann 2 presence inm all 3 regions was scored 3 if a 1988 stanger and baumann 1993 and stark species was present inm the regions at opposite and nelson 1994 ends of the mountain range we assumed it UPGMAUPCMA clustering suggested that stonstoneflyefly also was present in the middle and scored it 3 species composition of the raft rivelriver moun- RELATIVE DISTANCE FROM THE rapfraptRAFTRAFF RIVER tains was most similar to the sawtooth moun- MOUNTAINS dispersal is more likely between tains species composition is also similar to areas nearer to each other maemacarthurMacAithur and the wasatch mountains and different flomfrom the wilson 1963 we measured straight line dis- sierra nevada fig 2 A matrix correlation of tances from the raft river mountains to the r 0940.940 94 demonstrated a good fit between the single nearest and most distant counties near- dendrogiamdendrogram data and taxonomic distance est was cache county in the wasatch moun- matrix these data support previous findings tains and farthest was kern county in the by nelson 1994 harper et al 1978 and sierra nevada once the nearest and most dis- wells 1983 tant counties were located we calculated the single and complete link methods producedpi oduced difference in distance between them and divi- the same tree topology as UPGMALJPGMA and simi- ded it into thirds species whose nearest liter- lar cluster and matrix correlation values weiewelewere ature record was from a county in the most obtained single link clustering produced simsim- distant third were scored 1 if the closest ilarity values of 04310.4310 431 04070.4070 407 and 02720.2720 272 with a record was from a county in the middle thudthird matrix correlation r 0940.940 94 complete link it was scored 2 species from the nearest third cluster values were 04310.4310 431 03750.3750 375 and 02720.2720 272 were scored 3 with a matrix correlation value r 0940.940 94 clear creek yielded the highest number of RESULTS species howeverHowe veivel this is probably a result of the number of different times the site was we collected 37 species inm 25 geneiagenelagenera and 8 sampled the number of species per site and families from the raft river mountains they the number of times a site was sampled had a 214 GREAT BASIN naturalist volume 57

TABITABLE L 2 stoneflyStonefly species records and collection sites in the raft river mountains and records for these species in the sawtooth mountains sienaslenasierra nevada or wasatch mountains

collectionbolle3olleetloin sitessltessite s by numberilumber

pourcisourcet e raft river northNorth slope raft rivelriverr southscluths lopeslopesiope i angeranges s

species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19

pteronarcys princeps X x x x X x x x x x pteronarcella badia X X x x X x x x x x doddsia occidentoccioccidentalsoccidentaleoccidentalisdentalsalisails X X x x x x XY x taenioneinataemmema pallipallidumdum X x x x x x x x theniopteryxtaemopteryx aivalismvalisnivalis X x x Podpodmostapodmostkamosta decepta X x x x x Podpodmostapodmostkamosta delicatuladelicatula x x x x Prprostoiaostoia besametsabesainetsabesametsa X x x x X x x x x x capadazapada cinctipescinctipes X x x x x x x x x x x x capadazapada caysihaysi x x x x x x malenka califorcaliformcanicamca X x x x x x x malenka tinaunanna X x x x x paraleuctra ververshinavepshinashina X x x x x x x x x capniaacapnia gracigraciletriagi ucilanaletria X x x x x x x x x capniaacapniacapnw petila x x capniaacapnia vernalis X x x x capnwacapnura intermontana X x eucapnopsis brebredibredtbrevibrevicaudacauda x x x x x acapniautacapniaUt lemomanalemonianalemiemlemo nianaomana X x x x x x doroneuriaDoroneuria sp X x x x hesperoperla pacifica X x x x x x x x x x x x ciuradiura knowltoniknowltoni X x x x x x x x x isoperlqfulvaisoperla fulvabulva X x x x isoperla pinta X x x x isoperla quinquepunctatequinquepunctata x X x x x x x isoperla sobriabobriasobi id x x x x kogotus sp A X x x x x x Megarmegarcyscys sigsimsignatanata X x x x x x x x sawalaskwala americanaamenamencanacana X x x x x sitsliSuSiiwallia lineosafineosa X x x x x x x Suwallia pallidulapallidula X x x x x sweltsa borealis x x x x x sweltsaswelt&a coloradensis X x x x x x x x x x x sweltsa lamba X x x x x x x x triznakaTriznaka pintado x X x x x x riznakainznaka signata X x x x x x x utaperlaUtaperla sopladorasopladora X x x x x x x x species richness 23 22 17 23 1 3 8 1 1 4 2 6 2 14 21 10 34 22 29 collection visits 13 13 11 12 1 1 11 1 2 8 1 4 1 5 9 6

1 IL C clear11 creekgreekrcckreck CaupC unpgioiinclcaupgroundcampgroundground ll11 twin cleekcreek 2 clearC lc n creeki c c k lowerlowel budgebridge 12 big hollow leckdeckcreekmeek 3 onone mile ruckruekcuck 13 lettleftlertleht forkfoikfolknoik pine cleekcreek 4 gcorge creekC icekteek 14 rockroeknock cleekcreek 55 wildwildtwilde itt creekcu d 15 fisher cleekcreek 6 johnson leckleekcreekC 16 dunn leckdeckcreekmeek 7 raftrahthittrivliri upper nirnarrowslows 17 sawtooth mountains 8 rittkivuraftrahtRitt riverKivubiver lower idahoid1dahoihithi 18 sielsierri a nadaN ada 9 junction creekC ink 19 wasatch mountains 10 dovidove creek correlation coefficient of 0870.87 the upper nar- occur bodPodpodmostapodthostapodmostkamosta delicatuladelideficatuladeficatula claassen doron rows of the raft river displayed the greatest euria sp and Suwallia pallipalliduladula banks were amount of local endemism this is the only site collected only from clear creek and capniaacapnia in the raft river mountains where taeniopteryx petila was found only in fisher creek aivalisnivalis fitch capniaacapnia vernalis newport cap A distribution analysis of the 37 species nura termontanaintermontanaintermontanein nelson and baumann iso based on their occurrence in either northern perlafulva claassen and isoperla pinta frison or southern drainages showed that 36 of 37 199719971 stonefliesSTONEFLIES OF THE RAFT RIVER MOUNTAINS 215

024 028 032 036 040 oaboa4 048 052 pteronarcyidae pteronarcys princeps banks is abundant throughout the pacific raft river northwest and california but rare in the sawtooth rocky mountains jewett 1959 baumann et 0391 al 1977 its presence in the study area sup- W tch 0277 ports a faunal relationship with regions to the northwest pteronarcella badia hagen is sierra nevada common in the rocky mountains extends into the pacific northwest baumann et al 1977 fig 2 UPGMA dendrogram of similarity between the but is absent in the sierra nevada raft river mountains sawtooth mountains sierra pteronarcys californicacalifornica newport was absent nevada and wasatch mountains stonstoneflyefly species assemblages from the raft river mountains even though it is commonly found throughout the west baumann et al 1977 its absence is due probably to lack of high quality large rivers in the study species 97 were collected in northern drain- area ages capniaacapnia petila being the exception only taeniopterygidae taeniopteryx nivalis 28 of 37 76 species were collected in south- commonly found in eastern north america is ern drainages of the 9 species not collected the west being limited to from southern drainages 5 from the upper rare in western canada and the pacific northwest ricker narrows of the raft river and 3 mClearfromclearfrofrom clear creek both northern drainages were not found 1964 kondratieff and baumann 1988 ricker 1964 this distribution elsewhere in the study area these 2 sites alone attributed to a postglacial interglacial accounted for 32 of 37 84 total species col- or dispersal across canada lected in the raft river mountains differences and down the rocky mountains with subse- in northern versus southern drainage diversity quent extinction in central canada taeniopteryxtheniopteryx are almost completely explained by the unique- aivalisnivalis reaches the southernmost part of its ness of these 2 sites range in the raft river mountains where its logistic regression analysis of the 6 factors presence indicates a faunal relationship between affecting stonstoneflyefly long distance dispersal found these mountains and areas to the northwest that distance was the only factor significantly doddsia occidentoccidentalisoccidentalistalis banks and tainiotaeniothenio correlated with stonstoneflyefly presence absence data nema pallidumpallidum banks were both found in the in the raft river mountains every species raft river mountains and are widely distrib- collected in the raft river mountains was also uted in western north america baumann et found in the closest distance category those al 1977 they are not useful species for de- species in the middle or farthest distances were terminingter faunal affinities with any particular present in the raft river mountains only if region they were also present in the nearest distance taenionemathenionemaTaenionema pacificuspacificumpacificum banks is widespread an additional analysis of the remaining 5 fac- in western north america from alaska to new tors for only those species within the nearest mexico stanger and baumann 1993 but the distance category also demonstrated that none absence of sufficiently large rivers in the raft are significantly correlated with stonstoneflyefly pres river mountains may explain why it was not ence absence in the raft river mountains A collected in this study long distance dispersal model does not explain nemouridaeNEMOUR IDAE malenka califcalifomicacalifornicacalicallcailfornicaomica claas- presence absence data for the raft river sen Podpodmostapodmostkamosta delicatuladelicatula Podpodmostapodmostkamosta decepta mountains frison frostoiaPrFrprostoiaostoia besametsabesametsa ricker zapadacapada cinctcinctipesipes banks and capadazapada hansibansi ricker discussion are all widely distributed in western north america baumann et al 1977 and occur in faunal affinities the raft river mountains giving no clear indi- distributions of each species in the raft cation of faunal affinities river mountains on a family by family basis the presence of malenka tinaunanna supports fau- showed some interesting patterns of faunal nal relationships between the raft river moun- affinity not revealed by analyses of overall tains and regions to the northwest jewett similarity 1959 216 GREAT BASIN naturalist volume 57

CAPNIIDAECAPNIIDAL species in this family typi- hesperoperla pacifica banks has a broad cally have limited distributions nebekerNebekeikel and distribution in western north america and gaufin 1967 only 1 species eucapnopsis does not give any indication of affinity brevicaudabrevicauda claassen was collected in all 3 chloroperlidaeCHLORO PERLIDAE Susawalhasuwalhawallia pallidulapallidula hypothesishypothesizedhypothesiseded regions offaunalof faunal affinity banks sweltsa borealis banks and tnznakatriznakaTriznaka capniaacapniacupma gracilariagracilana claassen capniaacapnia ver- pintadapintadopinctada ricker were shared by all mountain nalis and Utacapnia lemonianalemoniana nebeker and ranges considered in this study Suwallia line gaufin were shared by the raft river saw- osa banks sweltsa coloradensis banks tooth and wasatch mountains capniaacapnia graci- sweltsa lamba needham and claassen and laria is also found in the pacific northwest but tnznakatriznakahrizTriznaka signata banks were shared between does not support faunal affinities with that the raft river sawtooth and wasatch moun- legionregion since its overall distribution is much tains only since these species all have such moiemolemore widespread widespread distributions they indicate little capniaacapnia petila and capnura intermontana about faunal affinities with any single region weiewelewere shared only by the sawtooth mountains utaperlaUtaperla sopladorasop ladora is primarily a northern the presence of C petila in the study area rocky mountain species and its presence in representsi epresents the southernmost collection of this the raft river mountains indicates faunal species in western north america it supports affinities with regions to the north extensive a faunal relationship with noinolnoithernnorthernthern regions of collecting has failed to find this species inm the the rocky mountains but not with the pacific wasatch mountains baumann et al 1977 or northwest capnura intermontana is limited the pacific northwest jewett 1959 to diadladrainagesmages in the northern great basin and peltoperlidaePELTOPERLIDAE no species inm this family tnbutanestributatributariesriesrles of the snake river nelson and were collected fromfi om the raft river or wasatch baumann 1989 nelson 1994 it indicates fau- mountains but it was represented by yora nal relationships between the raft river moun- perla bremsbrevis banks in the sawtooth mountains and regions to the north and west tains and yoraperlaYoraperla nigrisornamgnsomanigrisorna banks in the pipe111101didaeHI hnidaronidar eight species from this fam- sierra nevada stark and nelson 1994 the ily weiewelewere collected in the raft river mountains pacific northwest also has several peltopeltoperlidperlid seven of these ciuradiura knowltonknowltomknowltoniknowltonitom ansonfnsonfrison iso species jewett 1959 and their absence in the perla fulda claassen isoperla pinta fnsonfrisonanson raft river mountains implies a lack of strong soperlauoperlaisoperla quinquepunctatequinquepunctata banks isoperla faunal relationships with regions to the north- sobnasobriabobria hagen Megarmegarcyscys signata hagen and west sawalaskwala americanaamenamencanacana frison are relatively wide- in summary faunal affinities of those species spread in western north america baumann and families collected in the raft river moun- etalet al 1977 tains indicate the plecoptera fauna is composed an interesting species kogotus sp A with of 2 dominant groups table 3 the largest in achypterouspachypterousbrachypterous wings was collected in the raft group 68 of 37 species consists of species rivelriver mountains and further west in the jar widely distributed in western north america bridge mountains of north central nevada species associated with regions north and two other species of kogotus occur in western west of the raft river mountains constitute northnoi th america kogotus nonus needham and the and2nd group and represent 22 of the raft claassen in the coast and cascade ranges river mountains fauna most families in the and kogotus modestus banks in the rocky study are represented by at least I1 species from mountains jewett 1959 kogotus sp A in the this and2nd group similarities revealed by clus- raft rivelriver mountains may be a potential new ter analysis are the result of repeated faunal species with distribution between K nonus relationships within each family and not the and K modestuskmodestus result of any single family influencing overall PERLIDAEperiperlPLRI loalloaeIDAL only nymphs of dotodoroneuriadoroneunaDorodotonDoroneuriaeurlaneuna similarity patterns sp weiewelewere collected and since identification at colonization the species level is possible only with adult specimens we could not positively identify the raft river mountains provide a good this species as D theodora needham and model of an island habitat however geologic claassen 01or D baumanmbaurnanni stark and gaufin history indicates they have not been completely 199711997 stonefliesSTONEFLIES OF THE RAFT RIVER MOUNTAINS 217

TABLE 3 Stonstoneflyefly species found in the raft river mountains listed by family and region of faunal affinity each supportssupp oitsorts region of faunal affinity family widespread northwestern otherothel pteronarcyidae pteronarcys princeps pteronarcella badia taeniopterygidae doddsia occidentoccidentalisoccidentalistalis theniopteryxtaeniopteryx aivalisnivalis taenionemataemonemaTaeniTaeonemamonema palhdumpallidumpallidum

Nemournemoundaenemouridaeidae malenka californicacalifornica malenka tinaunannanne Podpodmostapodmostkamosta fehdehdelicatuladehcatuladelideilcotulacatula Podpodmostapodmostkamosta decepta Prprostoiaostoia besametsabesametsa capadazapada cinctcinctipesipes zapala caysihaysi leuctndaeleuctridae paraleuctra ververshinavepshinashina capniidaecapnndae eucapnopsis brevicaudabrevicauda capniaacapnia petila capniaacapnia vernalis capnio gracilariagracgraoilanaolana capnura intermontana Utacapnia letnonianalemonianalemoniana perlodidae ciuradiura knowltonknowltomknowltoniknowltonitom kogotuskogotiisKogotiis sp A fsoperlafulvaisoperla fuluajulua isoperla pinta isoperla quinquepunctataquinquepunctateqmnquepunctata isoperla sobnasobriabobria Megarmegarcyscys signata sawalaskwala americanaamencanacano perlidae hesperoperla pacifica dotodoroneunadoroneuriaDorodotonDoroneuriaeurlaneuna sp

Chlorochloroperlidaeperlidae Suwallia pallidulapallidula utaperlaUtaperla sopsopladoraladora Suwallia lineosa sweltsa botborealisealis sweltsa coloradensis sweltsa lamba tnznakatriznakaTriznaka pintadapintadopinctada tnznakatriznakahrizTriznaka signata isolated from these hypothesized colonization croizat et al 1974 the later retreat of glaci- sources in the past petersen et al 1980 cli- ers moved stonstoneflyefly habitats northward and matic changes caused by pleistocene glacial into higher elevations islandlikeisland like habitats were and interglacial cycles had a profound effect isolated by intervening lowlands and dry lake on biotic distributions and connectedness of beds this occurred most recently within the habitats within the great basin region axel- last 10000 yr stokes 1987 rod 1981 grayson 1993 these climatic oscil- during glacial intervals corridors were lations provided the mechanism for a vicari opened between the raft river and other ance model of stonstoneflyefly colonization in the raft mountain langesranges to the north and west species river mountains in regions to the north and west were able to pleistocene climatic changes directly affected reach their southernmost distributions in the the location of stonstoneflyefly habitats in north amer- raft river mountains before vicanancevicavieavicariancevicariancenancenanee isoiso- ica cooler climates moved south from polar lated them these same corridors of distribu- regions and pushed stonstoneflyefly habitats further tion did not exist between the raft river moun- south and into the lower elevations of the tains and sierra nevada due to the presence intermountain west sargent et al 1991 in- of extremely low elevation valleys in western creased precipitation and subsequent runoff nevada wells 1983 this explains why all from glaciers caused an overall expansion of species shared between the raft river moun- pluvial environments Stonstoneflyefly habitats ex- tains and sierra nevada are also present in the panded into generalized tracks of distribution rocky mountains 218 GREAT BASIN naturalist volume 57

the vicariancevicari ance argument is strengthened literature CITED by repeated patterns of affinity for each of the stonstoneflyefly families present in the raft river AXELROD DID I1 1981 holocene climate changes in relation disjunction similar for mammals to vegetation and speciation american mountains patterns exist naturalist 117847117 847 870 durrant 1952 brown 1971 and plants billings BAUMANN RWR W 1979 nearctic stoneflystonefly genera as indica- 1978 harper et al 1978 in the great basin tors of ecological parameters plecoptera insecta similarities in distribution patterns for differ- great basin naturalist 3924139 241 244 ent taxonomic groups are undoubtedly the BAUMANN RW AND ARA R GAUFIN 1969 the stonestonefhesstoneflicsfliesflics plecoptera of the wasatch mountains utah pro- result of a vicariantvicariant event splitting all biotasbigtas ceceedings of the utah academy of sciences arts and rather than multiple long distance dispersal letters 4610646 106 113 events stoneflyStonefly distribution patterns in the BAUMANN RW AR GAUFIN AND RF SURDICK 1977 southwestern united states stewart et al 1974 the stonestonefhesstonefliesflies plecoptera of the rocky mountains memoirs of the american entomological society al 1991 and northern mexico sargent et have 31131 1 260 also been attributed to vicariancevicariance events dur- BEHLE WH 1958 the birds of the raft river moun- ing the pleistocene tains northwestern utah university of utah biological series 111iiiill11 1 40 1978 avian biogeography of the great basin and conclusions intermountain region great basin naturalist memoirs 2552 55 80 the stoneflystonefly fauna of the raft river moun- BILLINGS WD 1978 alpine phytogeography across the tains consists of 37 species in 25 genera and 8 great basin great basin naturalist memoirs 2 families five are unique records for the state 105 117 BROWN JHJ H 1971 mammals on mountainmountaintopsmountamtopstops nonequi- of utah and 1 is a potentially undescribed librium insular biogeography american naturalist species in the genus kogotusKogotus 105467105 467 478 most species collected in the raft river COLEMAN BDB D MAM A MARES MRM R WILLIG AND YHY H mountains are those with widespread distri- HSIEH 1982 randomness area and species richness ecology 63112163 1121 1133 butions in western north america repeated CROIZAT L G NELSON AND DED E ROSEN 1974 centers patterns of faunal affinity for most stoneflystonefly of origin and related concepts systematic zoology families show strong faunal affinities with 232652396523 265965ggs 287 stonstoneflyefly assemblages in the rocky mountains DURRANT SDS D 1952 mammals of utah taxonomy and dis- tributiontrib university of kansas publications museum to the north the ranges exam- ution of mountain of natural history 616 1 549 ined the raft river mountains stonstoneflyefly fauna ESSIG EOE 0 1929 insects of western north america most closely resembles that of the sawtooth macmillan new york mountains in central idaho frisonPRISON TH 1929 fall and winter stonestonefhesstonefliesflies or plecoptera regression analysis demonstrated of illinois bulletin of the illinois natural history logistic survey 1834518 345 409 that a long distance dispersal model of stone GAIIINGAUFIN ARA R AVA V NEBEKER AND J SESSIONS 1966 the fly colonization cannot explain patterns of stonestonefhesstonefliesflies plecoptera of utah university of utah presence absence in the raft river mountains biological series 19 1 93 Stonstoneflyefly distributions in the moun- GRAYSON DKD K 1993 the desertdesentdesenn s past a natural prehistory raft river of the great basin smithsonian institution press tains appear to be the result of expanded stone washington DC fly distributions and subsequent vicariancevicari ance HARPER KT DC FREEMAN WK OSTLEROSTLEB AND LG caused by pleistocene climatic oscillation KLIKOFF 1978 the flora of great basin mountain ranges diversity sources and dispersal ecology great basin naturalist memoirs 2812 81 103 acknowledgments HATCH MHM H 1971 the beetles of the pacific northwest part V rhipiceroidearbipiceroidea sternoxiStern oxi phytophaga rhyn- we thank RR tolman MC belk JD chophorachophora and lamellicornia volume 16 university brotherson and DK shiozawa for help with of washingtonofwashington pipressess seattle and london experimental design analysis and use of HOLDRIDGE LRL R 1947 determination ofofwoildworld plant for- mations from simple climatic data science 105 equipment and facilities the following indi- 367 368 viduals assisted in collecting insect specimens HOLDRIDGE LRL R WC GRENKE WH HATHAWAY T LIANG from the raft river mountains C campora AND JAJ A TOSI 1971 forest environments in tropical SM clark K dobry BO huntsman DKDX life zones a pilot study pergamon new york HYNES H B N 1976 biology of plecoptera annual re- shiozawa stanger leavitt and GM webb HBN JA view of entomology 2113521 135 153 we especially thank our families for their sup- JEWETT SGS G JR 1959 the stonestonefhesstonefliesflies plecoptera of the port throughout this project pacific northwest oregon state monographs 313 1 95 199719971 stonefliesSTONEFLIES OF THE RAFT RIVER MOUNTAINS 219

1960 stonestonefliesflies the stonefhes plecoptera of california PETERSEN MSM S JKJ RIGBY AND LEL F HINTZE 1980 his- bulletin of the california insect survey 61256 125 177 torical geology of north america winwm C brown JOHNSON NKN 1975 controls of number of bird species company publishers dubuque IA on montane islands in the cleatcreatgleatgreat basin evolution PROVONSHA ANA N 1975 the zygoptera odonata of utah 2954529 545 567 with notes on their biology great basin naturalist kondratieff BC AND RWR W BAUMANN 1988 taeniopteryxtaemopteryx 3537935 379 390 of western north america plecoptera taeniopterytacnioptery RICKER WE 1964 distribution of canadian stonestonefliesflies pan pacific gidaigidae entomologist 6438164 381 390 Gegelwassergewasserwasser und abwasserAbwasser 3435503435 50 71 LOGAN ERE R AND SDS D SMITH 1966 new distributional rohlkROHLFROHLE ejFJ 1992 NTSYS pc numerical taxonomy and records of intermountain stonefhesstonefliesstoneflies plecoptera multivariate analysis system applied biostatistics occasional papers of the biological society of nevada inc setauket NY 91 3 SARGENT BJJ RWR W BAUMANN AND BC kondratiefrkondratieffkondratiefe MACARTHUR RHR H AND EOE 0 WILSON 1963 an equilibrium 1991 zoogeographic affinities of the nearctic stone theory of insular zoogeography evolution 17373 387 fly plecoptera fauna of mexico southwestern nat- MANTEL NA 1967 the detdetectionechon of disease clustering and uralist 3632336 323 331 a generalized regression approach cancer research SHELDON ALA L AND SGS G JEWETT 1967 stoneflySton efly emergence 2720927 209 220 in a sierra nevada stream plecoptera pan pacific MARDEN JH AND MG KRAMER 1994 surface skim entomologist 43143 1 8 stonefhesstonefliesflies ming stone a possible intermediateintel mediate stage in STANGER JAJ A AND RWR W BAUMANN 1993 A revision of the insect flight evolution science 266427266 427 430 stonstoneflyefly genus tahthenionematohtaemonemaTaemonema plecoptera taeniopterytaemoptery mcmahon JA AND TF WIEBOLDT 1978 applying bio- gidaigidae transactions of the amerleanamerican entomological geographic principles to resource management a society 119171119 171 229 case study evaluating holdridge s life zone model STARK BPP AND CRR NELSON 1994 systematics phy- cleatgleatgreat basin naturalist memoirs 22452 245 257 logeny and zoogeography of the genus yoraperiayoraperlaYorayotaperlaperia NEBEKER AXA V AND ARA R GAUFIN 1965 the capniaacapnia plecoptera peltoperlidaePelto perlidae Entomologicentomologicaentomologicala scandi columbiana complex of north americaamenea capniidaecapnndae navicagavica 2524125 241 273 plecoptera transactions of the american entomol- STEWART KW AND BP STARK 1988 nymphs of northnoi th ogical society 9146791 467 487 americanamerlean stonstoneflyefly genera plecoptera entomologi- 1966 new stonestonefhesstonefliesflies from idaho plecoptera cal society of america thomas say foundation 12 entomological news 273627 36 42 1 460 1967 geographic and seasonal distribution of the STEWART KW RW BAUMANN AND BP STARK 1974 family capniidaecapnndae of western north america pieple- distribution and past dispersal of southwestern copteracoptera journal of the kansas entomological soci- united states plecoptera transactions of the ameri- ety 4041540 415 421 can entomological society 9950799 507 546 NELSON CRR 1994 insects of the colorado plateau and STOKES WL 1987 geology of utah occasional papers of great pages basin 211 237 in KT harpelharper LLL L st the utah museum of natural history 616 1 280 clair KHH thorne and WM hess editors natural SURDICK RER F AND ARA R GAUFIN 1978 environmental history of the colorado plateau and great basin requirements and pollution tolerance of plecoptera press university of coloradoofcolorado boulder USU S environmental protection agency report EPAE PA NELSON CRR AND RWR W BAUMANN 1987 the winter 6004780626004 78 062 environmental monitoring and sup- stonstoneflyefly genus capnura plecoptera capniidaecapnndae in port laboratory office of research and develop- north america systematics phylogeny and zoo- ment cincinnati OH geogiageographyphy transactions of the american entomol- WELLS PV 1983 paleobiogeography of montane islands ogical society 1131113 1 28 in the great basin since the last glaciopluvial eco- 1989 systematics and distribution of the winter logical monographs 5334153 341 382 stonstoneflyefly genus capniaacapniacapnw plecoptera capniidaecapnndae in north america great basin naturalist 4928949 289 364 received 16 august 1996 NEWELL RL AND GW MINSHALL 1976 an annotated accepted 2 april 1997 list of the aquatic insects of southeastern idaho part I1 plecoptera great basin naturalist 3650136 501 504 creatgreat basin naturalist 573 D 1997 appp 220 230

BROOD REARING HABITAT USE BY RIO GRANDE WILD TURKEYS IN OREGON

thomas W keeganl2Keegankeegan1212 and john A CrawforCrawfordgrawforddl1

aiisibacralisallsails 1 liaiiaita 1 wild tinturkeytun key meleagrisMele agi is gallopavogallopavo broodin ooclkocl i earingrearing sites have beenbeed described for portions of their range butbutbioodbrood icaimgreariiig habitat use and charactcrischaiacteiisticsties of brood rearing sites used by rio glandegrande wild turkeys M g intermeintermew dia iriirlin the pacific northwest weicwelcwelewere unknown we described cover types at 362 bloodbrood rearing sites and meameasuredsuied habitat cliaiactcnsticse1ea iaracteris ticsbics at 64 of these sites used by a lecentrecentlyly established rio grande wild turkey population in southwesternsouthwestein oregon diningduring may septemberSeptembeibel 1989 and 1990 hens with bloodsbroods used 9 of 10 available covelcover types meadows mixed haihal dwoodconifeihardwoodcoriifer woodlands and savannassavatmasarmas weiewere used moiemolemore often than expected 47 of observations P 5 005 broods used mature mixed conifer and dense saplingpolesapling pole mixed conifer cover types less than expected and did not use brush fields many brood rearing sites weiewere characterized by a parklikepaikpalklikeilke appearance understory vegetation averagedaveiaver aged 20 cm tall and occupied 44 52 of broodbi ood rearingi eaiediedt ing sites whereas bare ground accounted foiroirolforhorbor 35 55 brood i earingrearingfearing sites were on southeast slopes moiemolemore often than expected and north slopes less than expected P 005oos0 05 we suggest that land managers maintain mixed hardwoodhaihat dwoodconifeihardwoodconiferconifer woodland and savanna covelcover types adjacent to meadows on south slopes to provide bloodbrood icanngrearirig habitat foifor rio giandegrandegeande wild turkeystiutin keys in southwestern oregon

kenkeykelkei wordswoun rio crandegrandegrandc wild tineinturkeyTiu ketkei brood rearing habitat use site characteristics oregon radio telemetry meleagrisMeleagns gallopavogallo pavo interinterinediamedlamediainedialuedia

rio glandegrande wild turkeys alediedleare native to the habitat use by introduced rio grande wild south central great plains of north america turkey populations received little attention or but translocation programs established popu- was investigated in simple systems eg 2 3 lations in 9 westeinwestern states wunz 1992 vari- cover types because brood rearing habitat ability in wild turkey movements and home is important in maintaining wild turkey popu- langesranges among geographic regions and sub- lations we examined habitat use by rio species was attributed primarily to variation in grande wild turkey hens with boultspoults in south- lesouiceresource availability brown 1980 wild turkeys western oregon and quantified brood rearing h equentlysequentlyfrequently demonstrated a high level of adapt- site characteristics ability by using a variety of cover types but selected specific vegetative characteristics with- STUDY AREA in covelcover types holbrook et al 1987 environmentalenanenvn onmental factorss that affect brood rear the 675 km2 study area is in the upper ing habitats aiealeare critical to population mainte- south umpqua river basin douglas county nance eveietteveletteverett ctet al 1980 suggested that poult oregon elevation ranges from 310 to 1525 m suisulsurvivalvivalvivai is directly i elatedrelated to the suitability of and the area is dissected by steep east west bloodbrood ieanngjeanngrearing habitat vegetative composition ridges franklin and dyrness 19731301973.130 attrib- and structure in brood langeiangerange influence protec- uted the heterogeneous association of plant tion fromhiomhrom predatorspiedapledatoistols poult mobility arthropod cover types in this area to diverse edaphic and abundance and exposure to dew which can geologic conditions overstoriesOverstories are dominated induce hypothermia bloodbrood rearing habitats by douglas fir pseudotsuga menziemenziesiimenziesiasii and have been described as paikpalkparklikeilielikeilke with moder- other conifers or oregon white oak quercus ate understory vegetation and nearby escape garryanagarryana and pacific ladronemadrone arbutus men covelcover porter 1992 habitat use by adult rio ziesiizieziesigsii deciduous midstory tree species include gigrandeande turkeys in their native i fangeangerange was stud- oregon ash fraxinus latifolialatifolia and big leaf ied extensively ege g logan 1974 baker et al maple acer macrophyllum common shrubs 1980 but quantitative descriptionsdescibesci iptionsoptions of brood include poison oak rhus diversilobadiversiloba oregon rearing habitats arearc lacking bloodbrood rearing grape berberis sppapp ceanothus ceanothus

I1 lpiiliiiiiloljnpartnct d fklwrilislidns urlduland Wildwikllikwildliftwildlifelift mh I1 laiifall 104 0vgocichoncicgon statst ite uniunelUnciuncisitysitssity cooioivillisvillisalli 01197331OH 97331 3803 11vsmticsciil addasaddvsaddIKSSvs OKKOIIoregon dpartnintlputiiiitofof fish tidiidildand xildlificvvildiitc p0papobx5ybox 59 portland 01197207OH 97207

220 199719971 WILD TURKEY BROOD REARING 221 sppapp and manzanita arctostaphylos sppapp plant were obtained in 3 approximately equal time nomenclature follows hitchcock and cloncronquistCionquist periods morning 05050.50 5 h before sunrise to 4 h 1973 the oregon department of fish and after sunrise midday 4 h after sunrise to 4 h wildlife ODFW released 58 rio grande wild before sunset and evening 4 h before sunset turkeys from texas and kansas on the study to 050.50os 5 h after sunset we located each hen 1 area in 1982 and 1983 RRR R denney ODFW time during each daytime period in every 2 unpublished data wk interval to the extent possible hens were monitored daily to identify mortality and METHODS movements accuracy of telemetry procedures was tested capture and radio telemetry by taking bearings on transmitters at 5 loca- we used rocket nets to capture turkeys dur- tions from 3 distances encompassing the range ing january 1989 and from december 1989 of tracking situations differences between esti- through february 1990 hen age yearling or mated and actual azimuths were used to calcu- adult was assigned from characteristics of pri- late error within and among distances and mary feathers larson and taber 1980 and locations variances of error angle estimates we equipped hens with 90 to 110 g radio were not homogeneous among observers and transmitters attached with modified backpack distances therefore standard deviations of harnesses kenward 19871031987 103 transmitters error angles were pooled when appropriate were equipped with motionmotlon sensitive switches and assigned to each angulationtriangulationtn based on expected transmitter life ranged from I1 to 3 yr observer and estimated distance from transmit- during 2 trapping seasons we captured 181 ter mean difference between estimated and wild turkeys in 1988 89 we equipped 26 true azimuths for all tested observer distance adult and 19 yearling hens with transmitters combinations was I1 s 04 we entered fifteen adults and 15 yearyearlingsyearningslings considered azimuths and receiver locations into program adults during the and2nd yr survived to 1990 in XYLOG dodge and steinerstemer 1986 to process the 1989 90 trapping season we equipped 10 angulationtriangulationtn data habitat availability was de- additional adults and 21 yearyearlingsyearningslings with trans- fined by a minimum convex polygon mohr mittersmitters bringing the total sample to 36 adult 1947 for all hen locations except for 2 hens and 40 yearling hens all radio marked hens that were excluded because of movements survived 2 wk after release we monitored 30 km 46 hens for 1 nesting season and 17 for 2 sea- habitat mapping and quantification sons we did not monitor 26 hens during nesting seasons because of deaths or transmitter we identified 10 cover types from aerial failures 19 adult 7 yearling therefore the photographs taken during summer 1989 and approximate maximum potential sample was ground reconnaissance sites n 56 for quan- 80 broods during the 2 yr period this was an tifying physiographic and vegetative variables approximation because some hens nestedrenestedrerenester were randomly located in all cover types and after brood loss producing 2 broods in 1 yr consisted of 3 points located 30 m apart cover keegan and crawford 1993 types were delineated and descriptions were hens with broods were monitored 2 times developed by sampling physiographic charac- wk from may through september during 1989 teristicste and 4 vegetative strata overstory and 1990 we considered young birds boultspoults woody plants 3 m tall midstory woody until 12 wk of age we verified brood survival plants 3 m tall but beneath canopy shrub by audio or visual evidence weekly until all woody plants 1 3 m tall and understory boultspoults disappeared or until broods were inte- woody and herbaceous plants I1 m tall grated into autumn flocks we quantified physiographic and overstory we ascertained direction to radio signals by and midstory vegetative variables at each site the peak signal method springer 1979 pre- percent slope aspect elevation percent non- liminary bearings and signal strength were forested habitat within 030.30 3 km species compo- used to move within 050.5os0 5 km of birds subse- sition density basal area percent cover and quent bearings provided tnangulationtriangulation data canopy height we estimated slope with a cli- hens were located by angulationtriangulationtn from 3 nometerno aspect with a compass and elevation locations or by visual observation locations from topographic maps percent nonforested 222 GREAT BASIN naturalist volume 57 habitat 10 tree cover within 03 km of white fir abies concolor and incense cedar each brood rearing site was estimated from calocedrusCalocedrus decurrentdecurrensde currens dense largematurelarge mature habitat maps with an overlay of 50 randomly mixed conifer DMC was the most common distributed points marcum and loftsgaarden cover type with overstory trees 50 cm dahdbh 1980 we recorded species distance to sam- and 110 yr old open largematurelarge mature mixed ple point and diameter at breast height dahdbh conifer OMC often developed from natural of the nearest tree in each quarter for mid- or management related thinning in dense story and overstory strata to calculate density stands some OMC stands may have devel- and basal area cottam and curtis 1956 per- oped following sparse regeneration in areas cent cover of overstory and midstory strata under even age management the second most combined was estimated with a sighting tube prevalent cover type was dense medium james and shugart 1970 by presence or young mixed conifer DYC open medium absence at 2 m intervals along four 10lom m tran- young mixed conifer stands were rare 02 020.2 sects originating at sample points we mea- and structurally similar to open saplingpolesapling pole sured heights of 5 randomly selected trees in conifer stands therefore we combined open each stratum with a clinometer to estimate mediumyoungmedium young mixed conifer stands with canopy heights open saplingpolesapling pole stands open saplingpolesapling pole we quantified the following shrub and young mixed conifer OSPQOS PC stands likely understory characteristics at all sample sites developed as a result of sparse regeneration or tall shrub cover understory vegetation height precommercial thinning dense saplingpolesapling pole understory groundcoverground cover and horizontal mixed conifer DSPC stands resulted from screening we estimated tall shrub cover along normal tree growth after even age regenera- each of four 10lom m transects with the line tion harvest or catastrophic disturbance intercept method canfield 1941 understory brood rearing site quantification vegetation was sampled in five 1 m2ma circular plots 1 at the central sample point and 4 at the steep terrain and dense vegetation of randomly selected points within 4 in we mea- the study area hampered direct observation of sured understory vegetation height at 4 ran- undisturbed broods therefore telemetry loca- dom locations in each im2ima1 m2ma plot and estimated tions and visual observations of undisturbed percent cover of grasses and grasslike plants broods defined sample sites we considered forbs low shrubs 1 I1 in bare ground and hen poult flocks undisturbed when turkeys woody debris A vegetation profile board 03030.30 3 were apparently unaware of observer presence x 121.21 2 in nudds 1977 was placed at sample or did not alter their activity eg feeding or points and observed from 4 locations at a dis- loafing tance of 10 in at 0750.750 75 in above ground level we quantified 2 randomly selected brood to provide an index of horizontal cover we rearing sites each week during 1989 and 1990 estimated percent horizontal screening for brood rearing seasons mid may through mid each 030 3 m interval on profile boards september with the provision that each brood three nonforested cover types covered 12 hen was included I1 timeseasontime season each hen of the study area recent clearcut 10 yr since with a brood contributed an average of 272.7 harvest brushfield and meadowpasturemeadow pasture table measured sites range 1 7 although a single 1 savannas were the rarest habitat typified hen was randomly selected as a focal hen by scattered trees or clumps of trees that usu- gang brooding was a common phenomenon ally had not been managed for timber produc- and brood rearing sites were occupied by up tion in contrast timber management likely to 4 radio marked hens as well as unmarked influenced stand development in hardwood hens with and without broods conifer woodlands HCW remaining cover vegetative characteristics of brood rearing types were seraiseral or management stages of habitats were sampled at 3 points I1 at the forested mixed conifer stands douglas fir observation or triangulation point and 2 located was a prominent component of most stands 30 in from the site at random compass bear- but several other conifer species frequently ings brood rearing sites were quantified with occurred as co dommantsdominants ponderosa pine the same methods used to develop general finusrinuspinus ponderosa sugar pine P lambetlamberlambertilambertianalambertiandandondtiana cover type descriptions 199719971 WILD TURKEY BROOD REARING 223

TABLE 1 descriptions of habitats available to rio grande wild turkeys in douglas county oregon 1989 90

MIXED hardwoodconiferHARDWOOD CONIFER at least 30 hardwoods in canopy layer usually dominated by oregon white oak and pacific ladronemadrone with scattered conifers all tree size classes present understory dominated by bare ground with approximatelyappi oxioxlmately equal proportions of grasses forbs low shrubs and debris poison oak was a common low shrub generally at lower elevations 750 m on southerly aspects included relatively rare riparian zones dominated by oregon ash and big leaf maple relatively sparse tall shrub cover 1 woodland stand canopy closure 40 occupied 7 of the area 2 savanna overall stand canopy closure 10 40 rarest cover type 2

MIXED CONIFER less than 30 hardwoods in canopy layer most stands were dominated by douglas fir but often contained 1 1 co dominants 1 dense largematurelarge mature overall stand canopy closure 70 average dbhdah ofoverstoryof overstory conifers 50 cm trees of this size were mature usually liollo1 l11010 yr old disturbance if any was related to fire wind or selectiveselectivesalvagesalvage logging characterized by sparse grass cover large amounts of bare ground low shrubs and slash most common covelcover type 49 2 open largematurelarge mature same size classes as in I1 with canopy closure between 10 and 70 stands generally resulted from shelterwood regeneration harvest commercial thinning or sparse regeneration understory dominated by bare ground and slash few shrubs grasses or forbs covered 4 of the area 3 dense mediumyoungmedium young canopy closure 27077070 average dbhdah of overstory conifers was 23 50 cm these diameters corresponded to approximate ages of 30 110 yr typically understory vegetation was 10 cm with little grass cover and much bare ground occurred on 14 of the area virtually all stands in this size class were classified as dense 4 dense saplingpolesapling pole canopy closure 70 average conifer dahdbh was 23 cm trees were usually 10 30 yr old grass was scarce whereas woody plants were dominant in the understory these stands generally resulted from even age management such as clearcut or shelterwood regeneration harvests found on 8 of the area 5 open saplingsaplingpolepole young canopy closure was between 10 and 70 in most stands overstory dahdbh was 23 cm open stands generally resulted from precommercial thinning or sparse regeneration there was much variation among stands but tall understory vegetation and high horizontal screening values were characteristic features most forbrichforbhorb nchrich cover type in some stands well developed shrub layers contributed to horizontal screening because of structural similarities open mediumyoungmedium young stands were combined with open saplingpolesapling pole stands occupied 3 of the area brushfield tree canopy was 10 and hardwood shrub cover was 15 seraiseralsera or climax communities dominated by a diverse association of tall shrubs including ceanothus manzanita and poison oak dense shrub growth provided high horizontal screening sparse grass cover and large amounts of bare ground occurred in understonesunderstoriesunderstoriesstones commonly occurred on areas that were previously clearcut or burned particularly where regeneration failed included rocky areas with scattered shrubs found on 4 of the area

meadowpastureMEADOW PASTURE natural or management induced openings with 10 tree canopy dominated almost entirely by low grasslike plants mosses and bare ground shrubs were rare and horizontal screening was low coverage of 3 included small pastures and hayfields

CLEARCUT areas where overstory was harvested within 10 yr and generally with 10 tree canopy often included seed tree regeneration and shelterwood regeneration after residuals were removed bare ground was the most common understory component relatively tall underunderstorystoly vegetation dominated by grasses with similar amounts of forbs slash and low shrubs including conifer seedlings occupied 6 of the area

statistical analyses were variable we detected few distinct outliers and inclusion of those observations did we analyzed data sets with a series of uni- results several variables displayed variate and multivariate procedures we com- not alter bined all brood rearing sites within cover nonnonformalnonnormalnormal distributions however transfor- types based on year hen age poult age and mations did not improve normality nor alter brood fate because of small sample sizes all results so original values were retained for all data sets were examined to assess outliers analyses when 2 variables were highly cor- normality multicolinearitymulticollinearity and homogeneity related r 070.7 we selected those variables of variance covariance matrices although data with the greatest ecological relevance or 224 GREAT BASIN naturalist volume 57 potential for management application that not used and open and dense MC and dense contributed to the most parsimonious descrip- SPC stands were used less than expected FP tion of relationships 0050.05 use of clearcutsclearcuts and OSP conifer stands we used analysis of variance to identify did not differ from availability variables that differed between groups eg habitat characteristics random and brood rearing sites and to reduce the number of variables entered in subsequent most characteristics of randomly located multivariate procedures stepwise discrimi- sites differed among cover types available to nant analysis SAS 1989 was employed to wild turkeys tables 3 5 similarly several select variable sets to distinguish between differences were apparent among cover types groups of observations we then included vari- used for brood rearing tables 6 7 able sets selected in stepwise procedures in brood rearing sites were structurally simple canonical analyses of discriminance SAS 1989 mean understory vegetation height at sites was to identify correlations between discriminating 20 cm in all cover types table 7 horizontal variables and canonical functions numbers of screening from ground level to 30 cm ranged variables included in these analyses were re- from 43 in meadows to 80 in OSP conifer strictstricteded according to sample size considerations stands whereas screening in strata above 30 for each group cm rarely exceeded 50 tall shrub cover was chi square analysis was used to test the null sparse 77 cm10 m in any cover type except hypothesis that cover types were used in pro- in meadows understory composition at brood portion to availability neu et al 1974 byers rearing sites was dominated by bare ground et al 1984 when the null hypothesis was re- 36 57 we observed considerable variability jected we calculated simultaneous confidence among proportions of other understory com- intervals to identify which cover types con- ponents grass cover ranged from 7 in OM tritributedbuted to differences in use and whether use conifer to 42 at meadowpasturemeadow pasture sites forbs was greater or less than expected preliminary low shrubs and woody debris each accounted analyses indicated that habitat use did not dif- for 7 28 ofbrood site understory cover among fer with year or hen age for any comparisons 8 cover types however total understory vege- consequently observations were pooled for tative cover grass forb and low shrub at examination of habitat use brood rearing sites was consistent and ranged from 44 to 52 among all cover types except RESULTS meadowpasturemeadow pasture brood habitat use rearing sites in meadows and pastures n 7 were distinguished from random hens produced 47 broods and we identi- locations by the amount of bare ground based fied a 1 brood site for 33 of those broods we on discriminant analysis we ascertained that did not locate 14 broods because of radio fail- group differences explained 70 of variation ure death of hens or brood disappearance in the canonical function P 00030.003 random we identified cover type at 362 locations of sites were characterized by twice as much hens with broods 12 wk old xY 11711.7 loca bare ground as brood sites tionshentionshen range 1 49 and 64 of these sites two variables overstory tree height and were subjected to detailed vegetation sam- density in the OSP conifer cover type pro- pling gang brooding occurred and sites vided discrimination between brood sites n sometimes were occupied by 1 I1 brood hens 7 and random locations P 00090.009 however used 9 cover types for brood rearing table 2 only 54 of the function variation was attrib- and habitat use differed from availability P uted to group differences random locations 00050.005 nearly 50 of brood locations were in tended to have fewer shorter overstory trees the 3 types used more often than expected compared with brood rearing sites HC woodland 27 meadowpasturemeadow pasture 12 within the DM conifer cover type midstory and HC savanna 8 collectively these cover tree height woody debris and horizontal types represented only 11 of available habi- screening 60 90 cm discriminated between tat furthermore when 90 confidence inter- brood sites n 11 and random locations vals were applied to brood habitat data use of the canonical function with these variables DYC exceeded availability brushfieldsBrushfields were accounted for 81 of the variation between 199719971 WILD TURKEY BROOD REARING 225

TABLE 2 habitats used for brood rearing by rio grande wild turkey hens douglas county oregon 1989 90 n 362 locations associated with 33 broods

Bonbonferroniferroni 95 pelpercentcent confidence intervalinteival cover type sites n available used lower upper seleeSelecselectionlselectionaselectionaltionatlona clearcut 17 61 47 08 86 0 meadowpastureMeadow pasture 44 25 122 71 172 brushfield 0 38 0 19 19 open saplingpolesapling pole mixed coniconlconiferferb 8 34 22 06 50 0 dense saplingpolesapling pole mixed conifer 2 80 06 24 35 dense young mixed conifer 76 144 210 140 279 0 dense mature mixed conifer 84 489 232 152 312 open matuiematulemature mixed conifer 5 43 14 13 40 mixed hardwood conifer woodlanwoodlandwoodlandswoodland0woodlandcdc0 97 68 268 198 338 mixed hardwood conifer savanna 29 20 80 38 123

where 0 presentsrepresentsle use in proportion to availability represents greatelgreater use of a habitat thinthiuthluthan expected indand represents less use of a habitat thinibinthanthun expected P 0050oos05 11nlinin conifer covelcover types open defined as canopy closure 70 woodlandcwoodland defined as canopy closure 24040 savanna canopy closure was 40

TABLE 3 overstory and midstory characteristics of available habitats in rio grande wild turkey study area douglas county oregon 1989 90 overstory midstory basal area basal area height m dahdbh cmem maham2ha treeshatreeshyTreesha height m dbh cm maham2haiwvha trepshaTreeshaesba cover dah treeshatreesba y X y y s y y type n x STs x sys x sT x x sT x s V X s T X7 s CC 3 4 2 6 3 01 01 19 10 MP 3 B 1 7 5 12 356 OSPC 8 14 1 23 2 55 13 126 36 5 1 7 1 2 05 282 77 DSPC 2 17 3 21 1 116 35 291 69 7 2 6 02 3 002 749 38 DYC 11 22 1 30 1 308 55 396 70 8 1 11 1 10 25 1005 250 DMC 15 50 2 82 4 524 53 100 12 15 1 16 1 16 242 4 545 82 OMC 2 43 11 86 25 144 68 23 1 10 2 16 2 2 04 82 1 HCW 7 16 1 25 3 100 11 213 31 7 1 9 1 4 10 701 153 HCS 4 20 5 33 10 65 14 130 66 8 1 14 6 2 03 294 117

CC clclearetcleaitutclearedcleaiclearearettut MP meadowpastuiemead pasture B brushfield OSPC open saplingsiplmgpolesaplingpolepole mixed conifer DSPC dense hiplmgpoltsiplingsiplingpolepole mixed conifeiconifer DYC auseansedense young mixed conifer DMC dense mature mixed conifer OMC open matuiemature mixed conifer HCW hardwoodh irdwoodcomteihardwoodconiferconifer woodland indand HCS hardwoodhardwoodconiferhaidwoodconiferconifer savanna

brood and random sites P 000010.0001 random 67 of between group differences FP sites had taller midstory trees more woody 000070.0007 random sites had more bare ground debris and greater horizontal screening than and forb cover and taller midstory trees than brood rearing sites brood rearing sites twelve brood sites were measured in HC midstory tree height and grass cover dis- woodlands bare ground forb cover and mid- criminated between random and brood rear story tree height discriminated between brood ing sites n 6 in HC savannas encompass- sites and random locations accounting for ing 89 of variation in the canonical function lo GREAT BASIN volumeZ M 57 22610 creatgrest naturalist 0 5

S IH ti t mixed i mature gil 15 2 CO 4 0 4 8oocicimci2 2 2 4 siCO 1 4f 5mcnmormanmom4 2 3 2bincooinco2 3 cmg a c debrisn ajl mature open u understory & Q chump 12 13 16 17 19 101 HX 23coocotdlaqcq 0 i i biloti oiloti 25cqco31 11 eciociocl height rir caco ii a 5 open li i U ci c 111 12 M 22 11I1 21 25moc16 15mio 1 OMC X falfnlct mocciomocmio9 5 t 5 clmlclelciot 0

caccaoOMC 10 I1 conifer einenn 3 CO 2miz0100 5 111 3coconco03 2 3 6 8 shrub1 S conifer

mixed zoklow IMX ioosmomcci17 0 39 22 30 15 26 5inoriimori10 11 gm oo 1 t 1 s COOICOINCOOICO IN il td mixed sxfsx7 DO 48 141 161 11 21 24 1 i 6 0 1 r i n 0 cl mature shrubmalum5 cm10 6 mature VI a y dense IH sy 101 ta 10 ntncooooto1 tall t3 a CO OPi 4 4 2 5 3 8 6 1 cover corcoe 11 dense F cover 13 73 69 49 43 34 31 IM coocclosoi0 t odim154 141coco odin bare 0 1 0 0 oj u DMC groundcover73 M Q IMX lococooocor25 63 33 28 43 51lctiooorl39 50 58 41 Ground3 2 ncdcocotiincommf DMC conifer3 u 090 conifer

S 198990 mixed 0 a i i 198900 SYCO mreboemo col co rs co i s CT 5 1 inn5imnimo2 4 2 1 coi3 1 3 2 3coidol2211 2oicocsicoi13 2 2 1 1 1 e mixed forb g young 1i v 0 i ajobjo young oregon IMX 16 15 26 11 16cdciooco012 8 13 16 slope S u 2 denseg 90 ea e5 13 1 00oo IMX 11 32rne17 13 15 19 15 14 0 dense 8 cotnotintx8 11 198990s 1989 DYC county t CO I1 Q f cSY 10O 4 3 1 2cli1linncts5 2 9 0 DYC g U of conifer grass oregonbo 3 2 S Js

1 conifer

1 1 l 3 douglas5 0 11i IHX 25 36 141 7 i 21 0 lotdltfolcocmlc1ca CO 5 i 2 3 2 iciice n0389 56toqotct68 57 59 74ptdoo36 78 mixed c m 602202ooomuo cotcor S Q huobuo000 mtoioirai ammom a c1ca mixed county 0 R pole rt 0 0 areamrea U 1saplingpole CO elevation sapling 1 03 saphugpole cmg SYco OTOomo 13 D 00oo ci a study 9 8 9 5 4 4 4 i omoo gae 684 p u oooiiraco douglas tooomoo888 945 928 7100 808 701 tb 937teotoodeo910 1072 iel K coil ooolrooola ioioao D zdy 1 120wot 5 1 005 000soi 0001I I Mlot0 dense1 Q w melot90120 0 CO dense turkey 03 nj 11 IMX 090 37 22 40 30 mooa 05 83 60 86 51 areaS 3.3 31COiocootdi lf c100citeCN fcopoo E oousooloco 3 eii zoloDSPC crtormomm Q T DSPC studytat3 wildkamb 5 1 conifer 1 i 11 1 i ll SY 1510 12 naznax 0 11 nbcmocermocnrMOc0 2Nr2 1 4 1 1 0 conifer turkey grandecamama g osto 11 10oco1 3 cmS S tcoT 3 9 6 4 5ioiracoira4 5 11 i nonforests 03 1mixed 0 o S savardasavarmaS bab0 mixed 0 E g 0 C 13 s 90OT0 mac0 6090meme pole riomlo S sladwild a screening within S ID 1 CO rt pole savarma IM IOCOOO30 38 28 savarda f ji 60 37CO 3 81 65lciot089 50 41 saplingpole iiicti69 1 181 11 151 conifer s Y 1555 C oooooiocoim48 0 5 6 i 1 3 ina V i il c saplingpole sapling 0 S sapling grande 5 hardwoodconifer S g habitats 1 a open hardwood g 5atlctl open horizontal0 os oo i 00 knect 14 bardwoodconifer 1 rtiarti N cma 9 8 7 6czicoicot4 5 3 7 mao S riorao ii C i U 0 0 g ffiafi 60 in OSPC eeme available 3060 coloOSPC t HCSxummus S 1 mt cootii booti lOO cOCI 40 01 si 0oo0 5 7cootie4 3 9 7 7 30CO0 IH 60ooti9 85laoloococi76 93 62 56 40 47 ol 00 1 D ooloot cttcldu30i0 habitats 03 andnadmam 1 y3ya c t zosHCS canopy brushfield of 2 0 brushfield Q S 505 0 co cover c U 5 0 andacm woodland

8 1 K SY ooooom10 14 0001 i i 36 31 61 0 CO 8 1 5 2 4 5 3 7 available M 37 tdoscoi79 83 91 il 46 mcimcoi X 00010 QOCTS COID cme 3 bbrcocotCOCOI coto Bm t r rt characteristics 0 B woodlndwoodlmd ca c5 l i 0 li conifer CTJ 300 030 t M pasture c i pasture tm 5polpoo CS of0 oi IM 25 96 85 98 73 72 51 60 59 0 81imtoirsooconi i oodira hardwoodconifer boriosoorios boott meadowpasture ooott meadowpasture 1 0 conifer hardwood meadowti meadow

s hardwoodconifer U rt s characteristics t it5a coco iai2 111 151 understory 1 1 7 l y en 3 3 1 8 2 2 4 hardwood 11 OO 11 u 0020COCO 00 111 15 en 3 3 1ii 8 2 1 llomli i 2 7 4 ft fe tat3 lloil MP a MP shodhowHCW u D Z imnyma earcutearcus ira tupeatypea tupeatypea clearcut 4 5 zobzowHCW conifer M M 3 1

1- c TABLE TABLE 0 1 cover cobozoboOSPC zoboDSPC apoDMC modHCW 1 cover OSPC amboDSPC HCW 0mcomc zod conifer colo DMC omc DYC acsaosHCS i 0mc 1mixed DYC HCSmoo h moo up moo 0 u MP esusssus 0 laclocMP cc 0 ccuswoqqqoaffiB 55ssus u 1 U uspqoqqqoffiffiB 199719971 WILD TURKEY BROOD REARING 227

TABLE 6 overstory and midstory habitat characteristics at rio grande wild turkey brood rearing sites douglas county oregonoregon19891989 90 clearcut meadow open sacaspcaS PC dense YC dense MC open MC HCW HCS n 2 n 7 n 7 n 15 n 11 n 3 n 12 n 6

11 y S S SF varlaVariaVariablevariablebvariablesvariable11bleb xy sy ST x ST f ST SF overstory height m 30 12 27 3 21 3 26 1 37 1 40 13 19 1 16 1 dahdbhdbhcmcm 44 17 54 4 31 4 38 2 60 4 80 5 29 3 27 2 babasalsalareaaredarea ahauhanha2ha 9 6 5 2 10 3 20 3 29 4 17 9 12 1 4 1 density noha 37 1 17 4 175 95 166 22 96 14 28 13 189 36 61 12 midstory height m 10 4 9 1 7 1 9 05 10 05 9 2 6 03 6 04 dbhcmabbdbb cm 13 7 17 2 9 2 11 1 12 1 12 3 9 1 9 1 basal area ahamaha2ha 3 3 1 04 3 1 7 2 8 1 1 05 5 1 1 03 density noha 101 28 44 21 268 123 493 95 580 99 59 7 594 162 156 28 canopy cover 25 20 32 9 45 7 71 4 69 5 23 12 67 3 30 5 elevation m 570 5 658 35 658 106 603 30 524 20 653 56 580 20 586 39 slope 11 3 15 2 16 2 13 2 15 2 18 5 12 1 11 3 nonforest within 03 kmkra 9 1 14 3 6 2 6 1 3 1 3 2 5 1 6 2 asicaspcSPC saplingsaplingpolepole conifer YC young conifer MC mature conifer HCW hardwoodhardwoodconiferhardwoodconifeiconifer woodland HCS hardwoodconiferhaidwoodconiferhardwood conifer savanna variablebviriable means followed by an asterisk were selected by stepwise discriminantbiscidisci imitantiminant analysis when compared to randomly located sites within the same cover type

TABLE 7 undeiunderstorydundei story habitat characteristics at rio grande wild turkey brood rearing sites douglas county oregon 1989 90

clearcut meadow openspcopenOpenS sacaspcaPC dense YC dense MC open MC HCW HCS n 2 n 7 n 7 n 15 n 11 n 3 n 12 n 6

15 S sy varlaVariaVariablebleb x S x sf S 7 V tall shrub cover colomcmlomcm10mcm lOm 9 9 13 7 67 15 48 11 77 13 12 3 50 12 12 3 horizontal screening 0 30 cm 44 9 43 5 80 6 67 5 71 3 65 11 57 5 63 5 30 60 cm 23 3 22 5 69 6 50 5 53 4 47 11 43 5 36 6 60 90 cmern 11 6 12 4 54 6 36 4 38 4 26 5 34 5 22 4 90 120 cm 13 10 12 5 51 7 36 4 36 4 23 6 38 5 21 3 understory height cm 5 2 15 2 20 3 15 2 15 1 12 4 13 1 18 3 undeiunderstorydundei story cover crassgrass 8 6 42 7 8 2 10 3 12 3 7 2 17 3 37 8 forb 12 1 18 4 28 2 15 2 16 3 22 6 10 2 12 1 baiebalebare 57 15 28 5 37 5 40 4 36 3 43 15 48 4 38 8 low shrub 8 6 10 2 18 4 23 4 27 2 11 3 18 3 11 3 debris 19 14 7 3 15 4 17 2 13 2 22 8 9 2 7 2

SPC saplingpolesapling pole conifer YC young conifer MC matuiematulemature conifer HCW hardwoodconiferhaidwoodconiferhardwood conifer woodland HCS hardwoodharhai dwoodcomfeihardwoodconiferconifer savannsavanna i tvariablebvariable means followed by an asterisk were selected by stepwise discriminant analysis when compared to randomly located sites within the same cover type 228 GREAT BASIN naturalist volume 57

P 0 0004000040.0004 midstory tree height followed pretedprated observations of hens with broods inm a the same trend noted in HC woodlands higher number of habitats as evidence that hens made valuesvalnesvaines at landomrandom sites grass cover was greater use of a range of conditions available in sev- at landomrandom sites compared with sites used by eral cover types however we recognized that broodshi oods we could not ascertain specific activities asso- bloodsbroods weiewelewere most often found on southeast ciated with some locations and measured sites slopes and those aspects were used more than may have represented a blend of areas used expected FP ooi0010.010 01 all north facing slopes for several reasons ege g food rest or escape less than for brood rear- weiewelewere used expected habitat characteristics ing P ooi00100.0101 southwest slopes were used min proportionpi opol tion to availability descriptions of specific brood site characteristics for rio grande turkeys in native range discussion were lacking however brood rearing habitats used by other subspecies were described as habitat use parklike porter 1992 our observations were comparisonsComp ansons of brood habitat use among consistent with previous observations through- rio glandegrande wild turkey populations were dif- out many parts of the country but tended to ficult because of the scarcity of quantitative represent the low end of the range of vegeta- evaluations in other regions or the relative sim- tive cover values the relatively open charac- plicity of vegetation where other populations ter of sites occupied by brood hens probably existed foifolfor example schmutz et al 1990 allowed for greater poult mobility and forag- delineated only 3 habitats for an introduced ing opportunity and reduced contact with wet rio grande turkey population in northeastern vegetation conversely sites typified by heavy colorado our results were similar to those of slash concentrations and dense understory mackey 1986 for merrlMerrimerriarrsmernamsarrsairs wild turkey M vegetation eg 80 horizontal screening g mernamimerriamimerrimerfimernamiami broods in washington where oak from 0 to 121.212min probably were used little and oak pine habitats weiewelewere used more than because poult mobility and hen vision were expected but only 2 othelother covelcover types were restricted mackey 1986 speculated that dense available our results also supported findings vegetation conditions limited use of some sites of rumble and anderson 1993 for merrlMerriaernammerriarrsmernamarrsairsaris s in washington turkeys in south dakota brood hens in both structural characteristics of sites used by populations used meadows more than expected rio grande hens with broods were different and dense conifer stands less than expected from those described by mackey 1986 for we hypothesize that rio clandecrandeglandegrande brood use of berriansmerriansMermemamrianss turkeys and by schmutz et al 1990 cover types was influenced by habitat patchi- for rio grande turkeys min colorado berriansmerriansMermemamrianss ness particularly in dense mature conifer and rio grande brood sites outside oregon stands for example less debris less canopy weiewelewere composed of taller understory vegetation covelcover and more grass cover at brood rearing 44 cm than we found in southwestern ore- sites in DMC stands pioploprobablybablybabiy indicated use of gon 15 cm we also observed different trends small openings 01or parklike areas inm otherwise in total understory cover and horizontal screen- dense stands use of small openings 1 ha ing compared with othelother areas Mermegamrianss within larger forest tracts was noted for mer brood sites mackey 1986 had greater screen- riansnam s turkeys in washington mackey 1986 ing than random sites average horizontal rumble and anderson 1993 reported that screening was moderate 33 64 at rio grande hens with boultspoults 12 wk old rarely moved brood rearing sites in oregon but little differ- 10 in into meadows similarly we saw few ent from random locations by contrast total broods near centers of lugelargehuge openings until understory vegetation cover at washington boultspoults were 3 months old brood sites 34 was less than at random sites ouioulour findings coincided with patterns of habi- mackey 1986 but total cover was higher at tat use summarized by porter 1992 savannas brood rearing sites 52 than at random sites and open woodlands provided brood rearing in oregon habitat and these cover types were used more schmutz et al 1990 found more grass cover than expected by rio grande wild turkey hens at some rio grande brood sites similar to that with broodsbi oods in southwestern oregon we inter in southwestern oregon but day et al 1991 199719971 WILD TURKEY BROOD REARING 229 noted less grass cover at some brood sites than woodland and savanna complexes would at random locations however available grass ensure availability of brood rearing habitat cover in study areas outside oregon was much turkeys would also benefit from conservation higher than that in southwestern oregon like and enhancement of openings and parklike schmutz et al 1990 we did not observe dif- areas in conifer cover types particularly in ferencesferences in sites used by different age broods areas with low hardwoodconiferhardwood conifer stand abun- nor did we record consistent differences be- dance or distribution tween brood rearing sites and random locations because dense mature conifer was used less rather rio grande brood rearing sites inm ore- than expected for brood rearing rio grande gon encompassed a variety of plant associa-assoriaassocia turkeys may be able to utilize landscapes tions and structures but were within ranges dominated by relatively young forests 30 110 observed in other parts of wild turkey range yr old and 23 50 cm dahdbh however dense southeast slopes provided boultspoults with the best mature conifer received heavy use for most opportunities for feeding and easy travel components of turkey life history eg ranked because these slopes dried quickly each day second for brood habitat use and first for hen and supported cover types used most by poult boostsroosts keegan 1996 therefore we do brood hens not recommend extensive harvest of mature timber as a means of increasing wild turkey management implications numbers AND recommendations acknowledgments our research indicates that grande wild lesearch rio this research was supported by the ODFW turkey broods the southern oregon in cascades US forest service lagrande forestry and use a variety of cover types and a range of con- range science laboratory national wild tur- ditions within cover types because they use key federation inc and oregon state uni- available most cover types for brood rearing versityversity we thank SR denney RA zalunardo rio crandegrande turkeys would probably thrive and other personnel of the umpqua national under a variety of habitat conditions includ- forest and ODFW for assistance and support ing some not considered typical wild turkey we appreciate the fieldwork of elPI burns habitat however before selecting sites for NE golly and BC quick CE braun B translocation managers should consider the healy ECEG pelren and RJ steidl reviewed types of habitat available and likely future drafts of the manuscript this is technical land management scenariosscenanossceseenanos esthetic and eco- paper 10858 of the oregon agricultural exper- nomic returns of translocation programspioprograms will iment station we followed wild bird research be enhanced by selection and pnontizationprioritization of guidelines described by oring et al 1988 sites best suited to wild turkey brood rearing habitat needs literature CITED although rio grande turkeys use a variety of habitats successfully several management BAKER BW SL BEASOM AND NJ SILVY 1980 turkey pioductivityproductivity and habitat use on south texas range- practices would enhance brood habitat only lands proceedings of the national wild turkey sym- dense stands resulting from recent perturba- posium 4145 158 tions conifer or tall shrub stands that devel- BROWN EKE K 1980 home range and movements of wild oped after even age management practices or turkeys a review proceedings of the national wild turkeytulkey symposium 42514 251 261 disturbance received little use com- when BYERS CRC R RKR K steinhorst AND PR KRAUSSMAN 1984 patible with other objectives we hypothesize clarification of a technique for analysis of utilization that prescribed burning eg in brushbrushfieldsfields availability data journal of wildlife management or patch thinning eg in dense saplingpolesapling pole 48105048 1050 1053 CANFIELD R H 19419411 application of the line stands to excessively interception reduce dense vegetation method in sampling range vegetation journal of to 25 low shrub cover and 25 cm vege- forestry 3938839 388 394 tation height would likely increase wild turkey COTTAM G AND JXJ T currisgurrisCURTIS 1956 the use of distance use particularly in areas where bloodbrood habitat measures in phytosociological sampling ecology 3745137 451 460 is limited maintaining areas of mixed hard datDAY KS LD FLAKIFLAKE AND WL tuckerTUCKLR 1991 move- woodconiferwood conifer covelcover types particularly oak ments and habitat use by wild tinturkeykey lienshensilens with 230 GREAT BASIN naturalist volume 57

broodsbloods in a gigrassass woodland mosaic in the northern southeastern association of game and fish commis- plains prairie naturalist 237323 73 83 sioners 277427 74 91 dodgeDODGL WE AND AJ STEINER 1986 XYLOG a com- MACKEY DLD L 1986 brood habitat of merrlMerriaernammerriarrsmernamarrsairs s turkeys puter program for field processing locations of radio in south central washington northwest science tagged wildlife USU S fish and wildlife service fish 6010860 ios108 112 and wildlife teciteelTecltechnicalmicalmicai report 4 22 appp MARCUM CLL AND DOD 0 loftsgaarden 1980 A non evererrevercrrevenEVERErrcrr DD DW SPEAKE AND WK MADDOX 1980 mappingmappmg technique for studying habitat preferences natality and mortality of a north alabama wild journal of wildlife management 4496344 963 968 tulkeyturkey population proceedings of the national wild MOHR CO 1947 table of equivalent populations of north turkeytinolnoun key symposium 41174 117 126 american small mammals american midland natu- FRANKLINFHANKLIN JEJ F AND CT DYRNESS 1973 natural vegeta- ralist 3722337 223 249 tion of oregon and washington pacific northwest NEUNEUCWCRCW CR byersandjmBYERS AND JM PEEK 1974 A technique forest and range experiment station USU S forest for analysis of utilization availability data journal of selvleeservice general technical report PNW 8 portland wildlife management 3854138 541 545 OR 417 appp NUDDsNUDDSTDTD 1977 quantifying the vegetative structure of HIICIKOCKHITCHCOCK CLL AND A CRONQUIST 1973 flora of the wildlife cover wildlife society bulletin 51135 113 117 pacific northwest university of washington press ORING LWL W ET AL 1988 guidelines for use of wild birds seattle 730 appp in research auk iosilosiloso1051 supplement 1 3311 HOLBROOK TH MRM R VAUGHAN AND PT BROMLEY 1987 PORTER WFWE 1992 habitat requirements pages 202 213 wild turkey habitat preferences and recruitment in in JGJ G dickson editor the wild turkey biology and intensively managed piedmont forests journal of management stackpole books Harnsharrisburgburg PA wildlife management 5118251 182 187 RUMBLE MAM A AND SHS H ANDERSON 1993 habitat selection JAMESJAMLS FC AND HHH H SHUGART JR 1970 A quantitative of merrlMerrimerriaasmemammegamaass turkey meleagrisMele agns gallopavogallopavo mesamimetrimerrimemamimerriamiami method of habitat description audubon field notes hens with boultspoults in the black hills south dakota 2472724 727 736 great basin naturalist 5313153 131 136 KEEGANKELGAN TW 1996 habitat use and productivity of rio SAS INSTITUTE INC 1989 SASSTAT user s guide version grande wild turkey hens in southwestern oregon 6 ath4th edition volume 1 SAS institute inc carsgarscarygary NC doctoral dissertation oregon state university cor- 943 appp vallis 135 appp SCHMUTZ JA CEE BRAUN AND WFWE ANDELT 1990 broodblood KEEGANKLLGAN TW AND JA CRAWFORD 1993 renestingRenesting by habitat use of rio grande wild turkeys prairie nat- rio crandegrandegi ande wild turkeys after broodbi ood loss journaljoul nalnai of uralist 2217722 177 184 wildlife management 5780157 801 804 SPRINGER JTJ 1979 some sources of bias and sampling KENWARDKLNWARD RER E 1987 wildlife radio tagging equipment error in radio angulationtriangulationtn journal of wildlifeofwildlife man- field techniques and data analysis academic press agement 4392643 926 935 london 215 appp WUNZ GAG A 1992 wild turkeys outside their historic range LARSON JSJ S AND RDR D TABERTABCR 1980 criteria of sex and pages 361 384 in JG dickson editor the wild age pages 143 202 in SDS D schemnitz editor wild- turkey biology and management stackpole books life management techniques manual ath4th edition the harrisburgharosHarnsburg PA wildlife society washington DC LOGAN TH 1974 seasonal behavior of rio grande wild received 22 february 1996 ruksysrukeysturkeys in westeinwestern oklahoma proceedings of the accepted 28 april 19919977 great basin naturalist 573 C 1997 appp 231 237

distribution OF THE MILLIPED TYLOBOLUS UTAHENSIS chamberlin WITH REMARKS ON T fredericksoniFREDERICKSONI CAUSEY spirobolidaSPIRO BOLIDA spirobolidae

rowland A sheiShelShelleyleyl1 and selena B bauerl

ABSTRACT tylobolusTylobolus utahenstsutahensis chamberlin the only presentativerepresentativele of the genus occurring in the southwestern deserts ranges from central inyo county california to the western periphery of kane county utah this distribution roughly corresponds to the northern limit of the mojave desert ecosystem and is also shown by the millipedmillimillipedeped bihPiepiedoluspiedoltisboh dolusdoltis utlisatusutus Chamberchamberlinlm Spirospirobolidabolida atopetholidaeatopethohdae and the centipede theatopsTheatops posticusposposticumticus say scolopendromorpha cryptopidae enttylobolustylobolusfredericksoninylTylobolus fredencksoni causey ostensibly from douglas county kansas is designated a nomen dubium and disregarded pending collection of fresh material darceusnarceus gorgordanusgorganusdanus chamberlin is deleted from south carolina and tennessee tylobolusTylobolus unciuncigeruswicigemsgerus wood occurs north of the columbia river in klickitatKlickitat county washington and Hilhiltoniustonius thebanusthebanus chambeilinchamberlinChambellinbeilin is referrable to Onyonycheluschelus cook in the family atopetholidae

key words tylobolusTylobolus T utahensis T frederickfrederiekfredencksonifredericksonisoni darceusnarceus americamericanusamericanosamencanusanus washington county utah inyo county california hiltoniusHiltonius

the subfamily tylobolinae of the diplopod questionable reports of an additional species family spirobolidae comprises 2 genera tylo T fredericksonifredericksoni causey in douglas county bolus cook and hiltoniusHiltonius chamberlin keeton kansas on the eastern periphery of the cen- 1960 revised both taxa recognizing 3 species tral plains we report 37 additional samples of of hiltoniusHiltonius in southwestern california and T utahensis that expand its range some 300 mi adjacent baja california norte and 2 species 480 km westward to central inyo county in mainland mexico that range northward into california east of the sierras and provide data santa cruz county arizona keeton 1960 on segment numbers lengths and widths we shelley 1995 plus unreported samples exam- also designate T fredericksonifredericksoni as a nomen ined by the ist author the ath6th species H the dubium because there is no recent indis- banus chamberlin occurring at theba mari putable evidence that a representative of this copa county arizona is errablerefreferriblereferrable to Onyonycheluschelus genus occurs east of the area occupied by T cook in the family atopetholidae as the ist utahensis the distribution of the tylobolinae author has learned from examining the female in the united states and the adjacent periph- holotype at NMNH see acronyms below in ery of mexico is shown in figure 1 acronyms a supplemental paper on the california repre- of sources of preserved study material are as sentativessentatives of tylobolusTylobolus keeton 1966 cited 6 follows species west of the crest of the sierra nevada BYU monte L bean life science museum I1 of which T unciuncigerusgerus wood ranged north- brigham young university provo UT ward to the columbia river at portland ore- CAS california academy of sciences san gon we extend its distribution into the adja- francisco cent periphery of washington based on a male CDFA california department of food and and female from klickitatKlickitat klickitatKlickitat county at agriculture sacramento FSCA tylobolusTylobolus extends southward into the DC life sciences division dixie college adjacent fringe of baja california norte boll- st george UT man 1888 keeton 1960 loomis 1968 and like FSCA florida state collection of arthro- hiltoniusHiltonius occupies the western interior as T pods gainesville utahensis chamberlin inhabits zion national LACMNH los angeles county museum park washington county utah there are also of natural history los angeles CA

northmorthorth carolina state musmuseumm of natiNattnaturaliraltrat sciences box 29555 raleigh NC 27626055527626 0555

231 232 GREAT BASIN naturalist volume 57

1924 at an unknown location in zion national park washington county utah DIAGNOSIS Metmetazonalmetaconalazonal striae terminating ventrolateralventrolateradventrolaterad well below level of ozoporesozopores anterior gonapodgonopod telopoditetelopodite apically blunt and rounded not uncinate posterior gonopodgonapod tibiotarsus forming right angle with distal pro- A jection figs 2 4 VARIATION new material agrees closely with the holotype the posterior gonopodgonapod tibiotarsus is slightly broader and blunter and the denticlesdenticles which causey 1955 and keeton 1960 termed bricklesprpricklesickles cover larger areas on both the anterior and posterior gonogonopodspods in juveniles the distal projection of the posterior gonapodgonopod is less angular extending submediadsubmediansubmediad A rather than downward or dorsad we reexam- A ined the holotype and found the dentidenticlescles to be stronger than shown by keeton 1960 figs 262 264 there is also a pronounced line or ridge of these along the ventral surface of the posterior gonopodgonapod telopoditetelopodite that he did not B show fig 5 in his supplemental work on california species keeton 1966 tabulated meristic and fig 1 distribution of the tylobolinae in the united morphometricmorphometric data we present such data for states and adjacent periphery of mexico solid lines A T utahensis table 1 to provide parallel accounts Tilltyloholusobolus dashed lines B hiltoniushiltomusholhoitomus tillobolusaillaili Hiltonius for all species of tylobolusTylobolus on the average females are slightly longer and broader than males adults equivalent length MCZ museum of comparative zoology are in throughout the range but those on the harvard university cambridge MA east are nar- NCSM north carolina state museum of rower the adult segment number varies from natural sciences raleigh 51 to 57 as all individuals with 52 segments NMNH national museum of natural have no legless segments except the epiproctepiproct history smithsonian institution wash- three females and 2 males have 51 segments ington DC none legless while 3 males and 1 female with SEM snow entomological museum uni- this count have I1 or 3 legless segments all versity of kansas lawrence individuals with S 50 segments have at least I1 UCD bohart entomological museum without appendages university of california at davis distribution occurring from the inyo UGA university of georgia museum of mountains on the eastern side of owens val- natural history athens ley inyo county california to the eastern edge of zion national park in western kane tylobolusTylobolus utahensis chamberlin county utah a distance of approximately 300 figs 2 5 mi 480 km figs 1 6 the millipedmillimillipedeped should also be expected along the virgin in the tylobolusTylobolus utahensis chamberlin 192560 61 river corner of mohave county keeton 19601311960.131 132 figs 262 264 northwestern ari- along this and other of califcalifornibolusCaliforniorniornibolusbolus utahensis chamberlin 1949 zona river in parts 166 chamberlin and hoffman 1958162 northern clark county nevada and throughout most of lincoln county nevada particu- TYPE SPECIMENS male holotype female larly in canyons and gorges east of caliente allotype and 2 male and 1 female paraparatypestypes panacea and piocheploche specimens in inyo MCZ collected by RXRV chamberlin in may county were taken in pitfall traps at elevations 199719971 distribution OF TYLOBOLUS UTAHENSIS 233

figs 2 5 tylobolusTylobolus utahensis 2 anterior gonogonopodspods of a male from saline valley inyo county california anterior view 3 left posterior gonopodgonapod of the same anterior view 4 the same caudal view 5 right posterior gonopodgonapod of holotype anterior view scale line 1001.00loo1 00 mm for all figures of 2296 6560 ft in addition to the types spec- 40 km S saline valley 2mam 29 april 1975 imens were examined as follows ARA R hardy CDFA and death valley nat pk california inyo co inyo mts lead can- panamint mts johnson gyn F 2 june 1961 yon ayncyn F 9 march 13 august 1981 D R waner NMNH giuliani CAS inyo mts hunter cynayn M F NEVADA nye co nuclear test site ranierramer 9 june 1980 26 may 1981 D giuliani CAS mesa 2mam juvguv date unknown DBD B thomas inyo mts willow cr F 16 september 1976 LACMNH NCSM collector unknown LACMNH saline valley UTAH washington co 12 mi 19219219.219 2 km NW 11 different stations but exact locations un- st george nr baker dam 3mam af2f 16 april known lom 17f juvguv 5 april 1959 14 june 1971 BKB K carrell DC pine valley F 26 may 1960 B banta CAS saline valley 10 stations 1971 prothero DC snow cynayn st pk M 16 along grapevine gyn rd but exact locations april 1982 RWR W baumann BYU pintura F 8 unknown 16m af6f 15 august 1959 7 may march 1941 J & W ivie NMNH motaquaMotaqua 1960 B banta CAS 21 and 25 mi 33633633.633 6 and M 17 april 1933 M zuiezulezue NMNH oak grove 234 GREAT BASIN naturalist volume 57

TABLE 1 meristic and morphometncmorphomorphometricmetric data on tylobolustylobolmhylhyiTylnyloTylobolmboimobolus utahensisutahenszs individuals are listed in descending order according to segment counts which include the epiepiproctproct the number of legless segments in addition to the epiepiproctproct if any is shown in parentheses measurements are in mm averages are for specimens with no additional legless segments males females segments length max width segments length max width CA hunter canyon silsiislisioslo511 417 40 491 358 39 CA lead canyon 502 274 31

CA willow creek 513 361 34

CA saline valley 57 703 51 55 867 83 55 738 56 55 488 56 55 727 70 54 764 74 55 672 72 54 714 66 55 628 68 54 655 66 54 626 68 54 649 72 54 593 60 54 623 66 11 11 53 651 68 54 612 76 53 542 55 53 742 82 53 488 56 53 710 61 521 462 51 53 608 67 52 648 65 51 471 53 51 461 55 464 226 28

averagesAveiaver ages 638 67 644 67

CA grapevine canyon road 55 603 59 56 652 77 55 596 62 56 506 52 11 55 591 57 54 720 76 55 480 52 54 638 67 54 604 61 53 668 79 54 561 60 53 646 69 54 550 62 52 563 59 53 778 67 53 564 61 52 689 71 52 583 70 52 578 63 52 576 60 averagesavelAvei ages 596 62 628 68

NV ranier mesa 53 592 57

UT zion national paikpalkpark 53 621 53 53 548 47 52 675 55 52 769 67 11 52 668 55 52 715 67 52 653 56 52 488 41 52 488 41 51 643 62 51 675 58 51 491 45 513 365 35 513 312 31 averagesavelAvei ages 610 52 633 57 199711997 distribution OF TYLOBOLUS UTAHENSIS 235

blechrostriatus shelley and bauer julida pae romopodidae occurring in saline valley and along the eastern slope of the sierra nevada shelley and bauer 1997 and idrionaria dineh shelley known only from washington county shelley 1996

enitylobolustylobolusfredericksoniTylobolus fredericksonifredericksoni causey Spirspirobolussporobolusobolus unciuncigerusgerus wood cragin 1885145 kenyon 189316 tylobolusTylobolus unciuncigerusgerus gunthorp 1913164 192188 californibolusCalifornibolus fredericksonifredericksoni causey 195578 80 figs lc 4 5 chamberlin and hoffman 7 1958161 tylobolusfredericksonitylobolusTylobolus fredericksonifredericksoni keeton 1960132 133 fig 6 distribution of tylobolusTylobolus utahensisutahensts for over a century records have existed of a mysterious tylobolinetyloboline on the eastern periphery of the central plains confusion began cpgd FE 2 june 1988 LL smithee BYU and the with zion nat pk exact site unknown MM FF aithyith cragin 1885 who reported Spirspirobolussporobolusobolus may 1924 VM tanner NCSM 3f3e 1927 unciuncigerusgerus wood from topeka shawnee af county that there AMU NMNH M july 1931 WJ gertsegertsch kansas stating were important differences with gerus NMNH and behind visitor s center F 30 unciuncigerus but the speci- april 1991 CS crawford NCSM kane co men was closer to this species than to any other zion nat pk east entrance M 3 march 1967 kenyon 1895 repeated craminscragins citation in- GE knowlton NMNH cluding the parenthetical question mark for an REMARKS hyiTylenloeniotylobolustyloboltisTyloboltisobolus utahensis demonstrates individual from weeping water cass county 2 important distribution patterns among north nebraska he also reported the abundant east american myriamyriapodsmyriapodapods the occurrences of the nearctic spirobolid darceusnarceus anwilcanusamericanosamericanusamericanus beau- genus and subfamily in coastal california and vois cited as Spirspirobolussporobolusobolus margimarginatusnatus say southwestern utah are reiterated by the tribe from adjacent sarpy county thus kenyons tynommatini and subfamily tynommatinae in record of unciuncigerusgerus is plausibly a misidentifi- the callipodoidcallipodoid family schizopetalidae as ty cation of this common species gunthorp 1913 nomma loomis occurs around san francisco 1921 questioned cragingragin s record as being geo- san pablo and monterey bays and idrionaria graphically improbable and deleted unciuncigerusgerus shelley inhabits washington county utah shel- from the kansas fauna he suggested that ley 1996 secondly 2 millipedsmillipedes and I1 cen- mutilated specimens of N americanosamericanusamericanus from tipede are now known to occur between cen- topeka cited as arctobolusArctobolus marginatusmarginatus say tral inyo county california and washington may have been misinterpreted thirty four years county a range that corresponds roughly to later causey 1955 proposed californibolusCalifornibolus the northern limit of the mojave desert eco- frederickfredericksonisoni for 2 males ostensibly collected system the other millipedmillimillipedeped is Piepiedolusdolus atusutus in 1949 in douglas county kansas and an chamberlin Spirospirobolidabolida atopetholidae which immature female taken in 1950 from monroe was previously known only from st george county iowa her illustrations of the antero- washington county hoffman and orcutt 1960 ventral corner of the mandible and both pairs but can now be recorded from inyo county of gonogonopodspods resemble the conditions in un based on 2 males from panamint springs UCD sigeruscigeruscigerus and all other kansas spirobolids that similarly the scolopendromorph centipede she examined were N americanusamericanosameric anus cited as N theatopsTheatops posticusposposticumticus say cryptopidae occurs oklahomaeoklahoman chamberlin keeton 1960 exam- in saline valley the nuclear testing site nye ined the holotype transferred C fredericksonifrederick soni county nevada and st george utah shel- into tylobolusTylobolus and noted that the vial con- ley 1990 1997 two additional millipedsmillipedes that tained fragments of 2 specimens the anterior may demonstrate this pattern are californiulusCaliforniulus end of a small male and the caudal end of a 236 GREAT BASIN naturalist volume 57 larger individual As the gonopodsgonopods appeared keeton 1959 1960 deyrup and franz 1994 to be from an immature specimen keeton there is no recent credible evidence that withheld commentary on the status of T fred another form exists in the eastern two thirds erickericksoniericksonesoni pending discovery of an adult male of the continent and the locality of T freder in our reexamination of the holotype we found icksoni may represent a labeling error the dis- what appeared to be 1 highly fragmented tributiontribution pattern of species along the pacific specimen with no legless segments at the cau- coast and in the western interior and another dal end its gonogonopodspods are accurately figured by some 1000 mi 1600 km to the east in the causey and keeton and are open to different eastern central plains is not demonstrated by interpretations keeton 1960 thought they another diplopod genus and thus seems im- were incompletely developed but gave no ex- plausible for tylobolusTylobolus we therefore designate planation for this opinion they seem mature T fredericksonifrederick soni as a nomen dubium and rec- to us and while similar to those of T uncigerusuncigerus ommend disregarding the species until its and plausibly representing a variant they also presence in the plains is confirmed by a show enough differences to be reasonably freshly collected male with unimpeachable interpreted as representing a distinct species locality data or until the millipedmillimillipedeped is discovered today 47 yr after its description T freder in the principal generic area probably in cali- icksoni is still questionable and no potential fornia and an accurate locality can be reported tylobolines have been collected from kansas nebraska or iowa during this time its osten- acknowledgments sible occurrence in the central plains continues to generate confusion as enghoff 1995 we thank RW brooks and JS ashe SEM family clade no 13 recorded tylobolusTylobolus from for loaning the holotype of C fredericksonifredericksoni the east and west nearctic based on this and L leibenspergerLeiben sperger MCZ for loaning the species to our knowledge the only definite types of T utahensis the following curators spirobolid records from kansas are of N2vav amer collection managers and university faculty icarusicanus cited as N oklahomaeoklahoman chamberlin from loaned specimens from the indicated reposito- douglas county causey 1955 and N annu ries BYU RW baumann CAS CE gris- laris rafinesqueRafinesque from osage county keeton wold CDFA AR hardy DC AH barnum 1960 we have seen I1 authentic kansas spiro LACMNH the late CL hogue NMNH JA bolidboled a juvenile from lawrence douglas coddington UCD the late RO schuster and county UGA that is clearly errablereferriblerefreferrable to UGA CL smith narchus aside from south texas which is occu- pocock pied by anelus richardrichardsonirichardsonbsoni allocoallopo literature CITED cockiidae and 1 or more representatives of the atopetholidae hoffman and orcutt 1960 BOLLMAN C 1888 notes on north americanamerlean julidae shelley and hoffman 1995 only 4 spiro annals of the new york academy of science 4 boloids definitely occur in the united states 25 44 CAUSEY NBN B 1955 spirobohdaespirobolidae Spirospirobolidabolida diplopoda east of the rocky mountains N americanosamericanusameric anus east of the rocky mountains journal of the kansas which is widespread east of the central plains entomological society 2869886928 69 80 and extends northward into quebec shelley 1957 flondobolusfloridobolusFlondFloridobolus a new millipedmillimillipedeped genus spiro 1988 N gorgordanusgorganusdanus chamberlin in peninsular bolboibohdaebolidaebohdamidae proceedings of the biological society of washington 7020570 205 208 Florida2 chicobolusChicobolus spinigerousgerus wood spinispinigerus spiro chamberlinCAMBERLINCHAMB EKLIN RVR V 1925 notes on some centipedscentipedes and bolidaebolmolidaeidae ranging from southern south car- millipedsmillipedes from utah pan pacific entomologist 2 olina to the south florida keys keeton 1960 55 63 and floridobolusFloridobolus panneripenneri causey floridoboli 1949 on some western millipedsmillipedes of the order dae endemic to the wales ridge Spirospirobolidabolida journal of the washington academy of lake high- science 3916339 163 169 lands polk causey and counties florida 1957 chamberlin RVR V AND RLR L horrmanHOFFMAN 1958 checklist of the millipedsmillipedes of north america united states national museum bulletin 2121212 1 236 CRAGIN FWEW 1885 first contribution to a knowledge of 2ktzkxton 1960r10 lleleeILL idldidadorad fcnni&lcinalls ofnoanof N nordogordaffiidaiuii from chtcutcarteicartel county the Mynomynopodamyxopodamyriopodapoda of kansas bulletin of the washburn liniussktennesetennessliniustennesvTenne svsK andwidmid chalicstollchalks on countscounty south caioluiaCaiog1101imtiulaluiaihla but tbtilctile ist autlioiauti101 hasliasilas college laboratoryLaboi atory of natural history 41434 143 145 loundfound kiiloiilonly N aiiihliinamm leannis ililiiin111 thsctlicscthac acas authentic samples of N palawspalaisgoirfnnitslamsiamslaws pa DrYdeydeyrupDLYRUPrueRup M AND R FRANZ 1994 rare and endangered ritliwitliith adnltnwhxliilt inah s liiviha banan takntmaknk n inin peniptiiliisnliiniar floridaI1 londloud 1 fronhonihonl aladiaaladmaalaelliiii iudandindtud st raie joh anthonth d conC onsttjikiitlqueotly we daadahdeletedeleto N norxorgorgordanitsoidnmidanilsdanits fronilnlioinleoin lenleuien biota of florida volume IV invertebratesinveiinmei tebrates univer- lussaiussa11111.11111 al1allailalaand south C arheaarhoano sity presspi ess of florida gainesvilleGames ville 798 appp 199719971 distribution OF TYLOBOLUS UTAHENSIS 237

ENGHOFF H 1995 historical biogeography of the hol- SHELLEY RMR M 1988 the millipedsmillipedes of eastern canada arctic area relationships ancestral areas and disper- arthropoda diplopoda canadian journal of zool- sal of nonmarinenoiinorinorlnon marine animals cladistics 1122311 223 263 ogy 66163866 1638 1663 GUNTHORP H 1913 annotated list of the diplopoda and 1990 the centipede theatopsTheatops posticusposticumpos ticus say in chilopoda with a key to the myriapodaMynapoda of kansas the southwestern united states and mexico scolo kansas university science bulletin 71617 igi161 182 pendromorpha cryptopidae canadian journal of 1921 craminscraginscragids collection of kansas myriapodaMynapoda zoology 68263768 2637 2644 canadian entomologist 538753 87 91 1995 the millimillipedmillipedeped family hnudisomatidaehirudisomatidae in the HOFFMAN R L AND BSB S ORCUTT 1960 A synopsis of the new world polyzonndapolyzoniida brimieBrimlebrimleyanabnmleyanayana 2310323 103 143 atopetholidae a family of spiroboloid millipedsmillipedes pro- 1996 the millipedmillimillipedeped order callipodidaCallipodida in western ceceedings oftheodtheof the united states national museum illiiiililii111 north america schizopetalidae tynommatinaetynommatmae and 95 166 a summary of the new world fauna Entomologic a KEETON WT 19195959 A new family for the diplopod genus Scandinscandinavicascandmavicaavica 272527 25 64 Floridfloridobolusflortdobolusobolus spirobolidaSpirobolida spirobolidea bulletin of 1997 the holarctic centipede subfamily plutoniplutonic the brooklyn entomological society 54154 1 7 uminaeumidaenumidae chilopoda scolopendromorpha cryptopi 1960 A taxonomic study of the millipedmillimillipedeped family dae brimieBrimlebrimleyanabnmleyanayana 245124 51 113 spirobohdaespirobolidae diplopoda Spirospirobolidabolida memoirs of SHELLEY RMR M AND SBS B BAUER 1997 new records and the american entomological society 17117 1 146 species and taxonomic alterations min the millipedmillimillipedeped 1966 the species of the millipedmillimillipedeped genus tylobolusTylobolus family paeromopodidae julida entomological news diplopoda spirobolidaSpirobolida A examinationreexaminationre transac- 1081108 1 14 tions of the american entomological society 921792 17 28 SHELLEY RMR M AND R L holholfmanHOIFMANHOIHOFFMANFMAN 1995 anelus richard KENYON FC 1893 A preliminary list of the myriapodaMynapoda of sonisom pocock a gulf coastal millipedmillimillipedeped of the united nebraska with descriptions of new species proceed- states and mexico spnobolidaSpirospirobolidabolida allopocockiidaeallopocockndae ings of the nebraska academy of science 3143 14 18 mynapodologicamyriapodologica 31073 107 114 LOOMIS MEH F 1968 A checklist of the millipedsmillipedes of mexico and central america united states national museum received 24 february 1997 bulletin 266 137 appp accepted 17 march 19919977 great basin naturalist 573 0 1997 appp 238 244 contrasting MOVEMENT AND ACTIVITY OF LARGE BROWN TROUT AND RAINBOW TROUT IN SILVER CREEK IDAHO

michael K younglyoung1youngbl richard A Wilkisonwilkison2wilkisonnwilkison23232 3 jmJ M phelps iiiiiii2j112 and JSJ S Griffithgriffith2griffitha2

ABSABSTRACTRAur1 recent radioradiotelemetrytelemetry studies demonstrated that stream dwelling trout are mobile but few have compared sympatric species we used radiotelemetryradiotelemetry to simultaneously monitor positions of 20 brown trout and 21 rainbow trout from may or june 1994 to february 1995 in silver creek a small spring fed stream in south central idaho our biweekly observations fromblom may to septemberSeptembeibel indicated that rainbow trout had larger home ranges medians 606 in v 131 in and moved greater distances medians 1109 in v 208 in than brown trout furthermore rainbow trout used moiemolemore positions than brown trout means 7 v 3 over this interval hourly diel monitoring revealed no significant difference in 24 h home ranges of rainbow trout and brown trout means 77 in v 105 in however activity patterns of the 2 species differed rainbow trout activity was usually highest during the day whereas brown trout activity tended to peak at night differences in foraging strategies and response to disturbance may be responsible for differences in mobility

key words diel activity kornehorne range movement brown trout salmo truttarainbowtrutta rainbow trout oncorhynchus mykiss

until recently stream dwelling trout often vary between day and night due to changes in were considered relatively sedentary with home light intensity prey availability and water ranges 50 in gerking 1959 northcote 1992 temperature other behaviors change during rainbow trout oncorhynchus mykiss and the diel cycle campbell and neuner 1985 brown trout salmo trutta were thought to and hill and grossman 1993 noted that rain- exemplify this pattern klein 1974 solomon bow trout move inshore and become less and templeton 1976 hesthavenHesthesthagenhestbagenhagenbagen 1988 in part active at night and feeding by rainbow trout because of the advent of radiotelemetryradiotelemetry stream apparently declines at night angradi and trout mobility has received greater notice and griffith 1990 brown trout were reported to seasonal movement may be more prevalent feed primarily in the evening elliott 1973 or than previously believed gowan et al 1994 during the day bachman 1984 clapp et al for example mean summersummerfallsummersallfall home range of 1990 noted that large 430 mm brown very large 435 mm brown trout exceeded trout tend to be more active at night but pat- 494.9 km in the ausableabusableau sablesabie river in michigan terns fluctuated monthly clapp et al 1990 median summer home there are few comparisons of diel and sea- range of large 340 mm brown trout was sonal mobility of sympatric salmonidssalmonids bjornn 400 in in north platte river tributariestributaries in and mallet 1964 evaluated movements of wyoming young 1994 and median home rainbow trout westwestslopeslope cutthroat trout 0 c range of small 240 mm colorado river cut- lekisilewisilewisi and bull trout salvelinus confluentconfluentusus throat trout 0 clarki pleuriticpleuriticusus was 233 in in in the middle fork salmon river idaho but a small wyoming stream young 1996 no results were based on angler recoveries of seasonal radiotelemetryradio telemetry study of rainbow trout tagged fish over several years matthews et al in streams has been reported 1994 used radiotelemetryradiotelemetry to monitor diel most movement studies have focused on changes in rainbow trout and brown trout hab- long temporal scales ie movement over weeks itat use but monitoring lasted only a month seasons or years miller 1957 mense 1975 and only I1 brown trout was tagged riley et al 1992 but distances moved within our objectives were to examine rainbow a diel period have been largely overlooked but trout and brown trout position changes from see clapp et al 1990 trout movement may late may to early february and to compare

1 rockyirocky mountain foiestfoyestroyest and range experiment station 222 S 22nd st laramieLararniearnle WY 82070 2jcpaltolentepalapaiepai tinenttenent of biological sciences idahoidalio state university pocatello ID 83209 3plcsenticscnt address idahoidalio poweipowelpower box 70 boise ID 83707

238 199711997 TROUT MOVEMENT AND ACTIVITY 239 their movement and activity over several diel intervals on a line parallel to the thalwegthalweg fish periods in a small stream with naturalized initially were located from a canoe then from populations the stream bank where we measured distance to the nearest stake fish were generally located STUDY AREA once or twice every 2 wk may september once or twice each month in october november and silver creek is a spring fed tributary of lit- december and once on 9 or 10 february 1995 tle wood river on the northern edge of the diel observations of fish activity and move- snake river plain south central idaho mean ment began on 22 june 12 july 17 august monthly discharge for 1994 ranged from 17 and 23 september groups of up to 7 fish up ma 1 1 m3 s in september to 515.1 m3ma s in march to 4 fish per species were monitored for 24 h discharge increased in autumnautuimi after irrigation all positions were identified from the bank of farmlands ended stream gradient of the and observers did not disturb fish we attempted 08os loio 1 edth study area is 080.8 101.0 in km and stream wwidthidth to monitor each fish every hour for at least I1 15 is 30 in aquatic macrophytes especially min and longer if a fish was active number of chara vulgaris and potamogeton sppapp are abun- fluctuations per minute in transmitter signal dant in summer and silt is the predominant strength was used as an index of activity clapp substrate with areas of gravel and marl much et al 1990 fluctuations resulted from changes of the riparian habitat consists of dense over- in transmitter antenna orientation caused by hanging stands of willow salix sppapp and birch fish movement relative to the receiver antenna fish brook trout betula sppapp other species are sunset and sunrise were 2121 h MDT and salvelinus whitefish pro- salvelinusfontinalisfontinalis mountain 0558 h MDT on the ist observation and 1933 h sopium williamsoniwilliamwilliamsonm bridgelipbridgelip sucker catosto- soni MDT and 0727 h MDT on the last observation mus columbianuscolumbianus redsidebedside shiner richardso we used telemetry to determine home range nius balbalteatusteatus longnoselongnose dace rhinichthys cata difference between furthest up and down- rachaeracractaetae speckled dace rhinichthys esculusososculus culus stream points and total distance moved sumsurn paiute sculpin cottus belbeldingidingi and wood river of all observed movements from the loca- sculpin cottus leioleiopomuspomus wilkison 1996 ist tion after implanting to the end of september we investigated trout movement in 2 non- summer and to early february overall and contiguous reaches of silver creek upper the during each 24 h cycle for these calculations reach is 414.1 kinkm long and largely on a nature we disregarded initial capture position because conservancy preserve where angling is per- some fish have been displaced during mitted but no harvest allowed the lower reach may electrofishingelectrofishing we also excluded fish that were is 51sisl5.1 km long andharvestand harvest is state regulated 2 followed for fewer than 50 d n 5 positions fish 305 minmiumm or 406 minmm can be kept an inin- 10 considered different terveningtervening 38 km reach was periodically visited in apart were to determine presence offishof fishhishbish with transmitters biweekly movement data were nonnonformalnonnormalnormal kolmogorov smirnov one sample test P METHODS AND MATERIALS 00010.001 n 257 observations to determine whether there were differences in movement we collected rainbow trout mean total among 2 wk intervals and between species length TL 419 mm range 357 475 mm n from may to september 1994 we used 2 way 21 and brown trout mean TL 494 mm range kruskal wallis tests home ranges and dis- 342 622 mm n 20 by angling or electro tances moved were analyzed by season sum- fishing and implanted transmitters on 13 15 mer may september n 36 and overall may and 12 14 june 1994 we surgically im- may february n 23 because most data planted sealed coiled antenna transmitters in also were nonnonformalnonnormalnormal differences in seasonal anesthetized fish in the body cavity immedi- home ranges and distances moved between ately anterior to the pelvic girdle and released species were compared using mann whitney fish at or near the point of capture after recov- tests number of positions occupied in sum- ery see young 1995 for details we monitored mer was normally distributed and analyzed fish by radiotelemetryradiotelemetry until early february using t tests 1995 only healthy fish were implanted water temperature was measured on a ryan to define the longitudinal position of each thermograph in the middle of the upper reach telemetry location we staked the bank at 50som m we used daily maxima to calculate a mean 240 GREAT BASIN naturalist volume 57 maximum temperature for each 14 d period brown trout we used rank correlation to relate mean biweekly maximum water temperature to median 250260 laibowraibow trout biweekly movement temperature we compared percentage of fish active 200 mean signal fluctuations and mean 16 per minute E E distance moved in each 2 h period among 3 8 150 diel intervals night crepuscular periods inter- 14e vals containing sunset and sunrise and day re- iai5 loo100 maining light hours we used an arcsin transformation on the percentage of active fish to 0 correct for heteroscedasticity we used a square 12 root transformation on mean signal fluctua- 0 tions per minute and mean distance moved 5265126 69 623 77 721 84 8188118 91 915 we examined differences between species and summer 1994 among times of day using 2 way analysis of variance and we used tukey s HSD test to fig I1 median biweekly movement of brown trout n compare activity at different times of day for 18 and rainbow trout ra 18 from late may to late september 1994 silver creek ID labels on the horizontal each species data on diel home ranges and axis represent 2 wk midpoints distances moved by each species were normal and were compared using t tests we 1 20 pimentel used biostatbiossat I version 202.0 and relatively active throughout the summer with smith 1990 to perform the kruskal wallis peak movement 14 27 july median biweekly tests and the parametricnonparametricnon tukey s HSD tests movement of rainbow trout was positively cor- for multiple comparisons and SPSSPC related with mean biweekly maximum water version solsoi5015.01 SPSS 1992 for all other analy- temperature r 0800.80oso FP 00010.001 though ses throughout we considered FP 005oos0.05 as this relationship may represent a threshold indicating significance response to high temperature more than a consistent trend at all temperatures water RESULTS temperature peaked on 21 22 and 25 july at 195c median biweekly movement of brown biweekly observations demonstrated that trout was not related to mean biweekly maxi- brown moved less trout than rainbow trout mum water temperature r 0260.26 P 0490.49 from may to september in summer brown directional trends in movement were not trout had smaller home ranges medians 131 evident nine of 36 trout followed in summer m v 606 m P 00460.046 and moved shorter dis- had moved up and 7 had moved downstream tances medians 208 m v 1109 m FP ools00180.018 100 m eight of 22 trout followed until than rainbow trout brown trout also used december or february had moved up and 6 fewer positions than rainbow trout means 3 v had moved downstream 100 m species dif- 7 P 00010.001 there were no significant differ- fered however in fidelity to the site where ences P 0890.89 in the overall home range first located by september and by late winter medians 1158 ra v 941 m and distance 60 of brown trout were found within 100 moved medians 1971 m v 1687 m between m of their first location by september 40 brown trout and rainbow trout which we of rainbow trout were within 100 m of such attributed to greater mobility of brown trout sites and by late winter only 3 of 14 fish were from october to february possibly associated such patterns suggest that brown trout use the with spawning nonetheless rainbow trout same sites during the day throughout much of had the largest summer 3865 m v 2530 m the year whereas rainbow trout are less likely and overall home ranges 10390 m v 3452 m to use the same position consistently biweekly movement of brown trout was less there were differences in diel activity be- than rainbow trout from may to september tween species throughout the diel cycle rain- medians 6 m v 46 m P 00010.001 fig 1 brown bow trout had more signal fluctuations per min- trout movement peaked in late may and then ute than brown trout means 5 v 4 P 00030.003 declined rapidly whereas rainbow trout were and a greater percentage of fish were active 199719971 TROUT MOVEMENT AND ACTIVITY 241 during 2 h observations means 87 v 51 P creek was smaller than that of brown trout in 00010.001 furthermore we observed contrasting other radiotelemetryradiotelemetry studies 29 km hudson behavior between species at different times 1993 34 km meyers et al 1992 96 km young table 1 fig 2 for brown trout the peaks in 1994 unlike those studies we did not exam- percentage of fish active and signal fluctua- ine all downstream portions of silver creek tions per minute were at night for rainbow nor did we follow fish throughout an annual trout these values were highest during the day cycle either factor could explain home range and crepuscular periods whereas brown trout differences nevertheless variables such as a moved most during crepuscular periods and greater food supply or environmental stability night rainbow trout failed to demonstrate a eg reduced discharge variation or water tem- significant diel pattern for this variable there peraperatureture may have rendered movement less were no significant species differences P advantageous than in other systems 00130.1313 in diel home range means 77 in v 105 in feeding strategy and fish size may also con- or diel distance moved means 192 in v 274 inm tribute to differences in movement based on maximum diel home range for brown trout daytime bank observations in a pennsylvania was 238 in and 352 in for rainbow trout stream bachman 1984 contended that brown diel movements of brown trout were more trout had small summer home ranges ca 4 in predictable than those of rainbow trout brown long and were active during the day but brown trout left daytime locations each evening and trout in our study moved extensively and were 8 of 10 returned before 0800 h the next day in largely nocturnal also see regal 1992 hud- contrast rainbow trout patrolled diel home son 1993 the largest fish in bachmaibachmalbachmanbaehman s 1984 ranges irregularly often visiting the same posi- study 330 minmm was smaller than the smallest tions throughout the 24 h cycle brown trout in our study though both streams are productive spring creeks brown trout 400 discussion minmm in silver creek forage primarily on large invertebrates and fish wilkison 1996 whereas home ranges of brown trout in silver creek adult brown trout in the pennsylvania stream were larger than in many other studies mense appeared to feed largely on drift piscivorous 1975 bachman 1984 hesthavenHesthesthagenhagen 1988 but brown trout may forage more successfully at differences may be attributed to differences in night and move to new habitats when prey methods ie tagging and mark recapture tech- become locally depleted or have fled to cover niques ignore behavior of marked fish that are clapp et al 1990 drift feeding juvenile brown not recaptured and tend to produce smaller trout in a new zealand stream however were home range estimates than radiotelemetryradiotelemetry most active at night perhaps in response to a young 1994 gowan and fausch 1996 yet the nocturnal increase in maeromacromacr6invertebratemacroinvertebrateinvertebrate drift maximum home range of brown trout in silver mclntoshmcintoshmeintosh and townsend 1995

TABLE 1 activity patterns of brown trout and rainbow trouttront in silver creek ID may 1994 to february 1995 during the day n 48 crepuscular periods n 16 and night n 32 using means standard deviations in parentheses of untransformed variables an asterisk indicates a significant difference for the species main effect for comparisons between different times of day for individual species values followed by the same letter are not significantly different time of day species day crepuscular night mean percentage offish active brown trout 2732 60436043lb6043ahlbah 83321 rainbow trout 9810- 92159215abab 693569351 mean fluctuations minmm I1 brown trout iia11 22a22 871 rainbow trout W52a 63163 1 3313311 mean distance moved in 2 h 1 brown trout 612giggiga 5673b 46581 rainbow trout 1513a 3038l3038a30381 126126l12611201 242 GREAT BASBASININ naturalist volume 57

that colorado cutthroat trout isolated 00oo 0 river above a barrier moved less than fish in a larger 80- connected stream segment below the barrier 60 differentDifferent patterns of biweekly movement C puzzling 10 by each species were we considered it unlikely that the early peak in brown trout 20 bo tout R at7atat activity was related to disturbance associated 0 with electroelectrofishingfishing and surgery only 11 of 20 13 15 1 7 1 2 1 3 1 brown trout were implanted in mid may we B 0 implanted the remainder in mid and the 10 june biweekly movement of this group in late june was less median 16 m than that of brown Z trout implanted in may median 34 m we 4 speculate that brown trout movement in late 2 2 may was associated with migration possibly

0 from outside the study area we captured only 13 is 17 19 21 23 1 3 5 7 1 7 brown trout suitable for implanting during electroelectrofishingfishing of the lower reach of the study C area in mid may but by mid june large numbers of brown trout were observed and cap- 0 E 60- tured perhaps the fish captured in may were migrating to suitable summer positions whereas 40 brown trout captured in june had already 20 selected such positions see brideutbridcut and giller

0 1993 for a similarly timed peak in brown 3 15 17 19 21 3 7 trout emigration year round tracking over a time of day larger area would be necessary to test this fig 2 diel patterns of the A mean percentage ofoffishfishhishbish assertion the late july peak in rainbow trout active B mean signal fluctuations per minmm and C mean movement may be attributed to fish moving to distance moved per 2 h by species in silveisilversliver creek ID on lower water temperatures but we did not ob- 12 17 august and 23 september 1994 n 22 june july serve this species concentrating in particular 264 observations closed and open circles lepresentrepresent 11 average times of sunset and sunrise respectively note areas clapp et al 1990 and meyers et al the change in the scale of the horizontalhoihol izontal axis in C labels 1992 attributed large scale movements of on the horizontalhoihol izontal axis represent 2 h midpoints brown trout to changes in water temperature and nielsen et al 1994 noted that juvenile steelhead move to colder habitats as water rainbow trout in streams are assumed to temperatures increase rainbow trout also may feed primarily on drift elliott 1973 tippets have moved in response to decreases in dis- and moyle 1978 cada et al 1987 and thus solved oxygen summer fish kills attributable are unlikely to locally overoverexploitexploit their prey to hypoxiahypoxis have been noted in several reaches stefanichStefamch 1952 also concluded that rainbow of silver creek in previous years paul todd trout seemed more mobile than biownblownbrown trout in the nature conservancy personal communi- a montana stream and unlike brown trout in cation nonetheless we cannot discount that this study and others ege g clapp et al 1990 variables such as food availability or macro rainbow trout showed little fidelity to daytime phytepayte growth could have contributed to rain- positions relative differences in movement in bow trout movement different streams may be related to stream size adult rainbow trout tended to be most for example movements of rainbow trout in active during the day which may reflect forag- silver creek exceeded those of rainbow trout ing preferences although rainbow trout can in a smaller minnesota stream cargill 1980 feed on drift at night jenkins 1969 evidence but were less than those of rainbow trout in suggests this behavior is uncommon in sum- the larger middle fork salmon river bjornn mer edmundson et al 1968 campbell and and mallet 1964 similarly young 1996 noted neuner 1985 angradi and griffith 1990 but 199719971 TROUT MOVEMENT AND ACTIVITY 243 see matthews et al 1994 warner and quinn acknowledgments 1995 noted that lentic rainbow trout moved less at night and remained inactive for long we thank K meyer E partridge R king periods another drift feeding species col- and PE todd for reviewing the manuscript and orado riverrivercutthroatcutthroat trout was consistently K meyer and E watters for assistance in the active only before dusk and after dawn young field this study was funded in part by grants et al in press from the idaho field office of the nature in part differences in the activity of brown conservancy idaho department of fish and game trout and rainbow trout may have led to differ- and idaho state university ences in summer home range and biweekly movement disturbed rainbow trout tended to literature CITED move up- or downstream but did not seek angradiangraditrandjsTR AND JS GRIFFITH 1990 diel feeding chron- cover during electroelectrofishingfishing we observed ology and diet selection of rainbow trout oncorhyn- schools of rainbow trout fleeing downstream chus mykiss in the henry s fork of the snake river and we often chased them for 100 m idaho canadian journal of fisheries and aquatic sciences 4719947 199 209 because rainbow trout active during the were BACHMAN RAR A 1984 foraging behavior of free ranging day we believe that anglers frequently dis- wild and hatchery brown trout in a stream transac- placed these fish but because brown trout tions of the american fisheries society 1131113 1 32 often were concealed in cover during the day BJORNN TC AND J MALLET 1964 movements of planted and wild trout an idaho system transactions see also clapp al 1990 young 1995 they in river et of the american fisheries society 937093 70 76 were less likely to be disturbed by anglers BRIDCUT EEE E AND PS GILLER 1993 movement and site and those we monitored typically sought near- fidelity in young brown trout salmo trutta popula- by cover when displaced lack of significant tions in a southern irish stream journal of fish biology 4388943 889 899 diel trends in rainbow trout movement despite CADA GFGE JM LOAR AND DK cox 1987 food and the customary trend in angler presence sug- feeding preferences of rainbow and brown trout in gests that angler disturbance explains only southern appalachian streams american midland part of the difference between the 2 species naturalist 117374117 374 385 CAMPBELL RFR F AND J H NEUNER 1985 seasonal and such as site specific variability JH factors in mac diurnal shifts in habitat utilized by resident rainbow roinvertebrate drift inherent behavioral dif- trout in western washington cascade mountain ferenferencesces or competitive displacement by brown streams pages 39 48 in FWEW olsen RGR G white and trout gatz et al 1987 also contribute to the RHR H hamre editors symposium on small hydro power and fisheries westeinwestern division american greater movement by rainbow trout in summer fisheries society bethesda MD because of the growing use of radioteleme CARGILL ASA S 1980 lack of rainbow trout movement in a try clapp et al 1990 intensive electrofishingelectrofishing small stream transactions of the american fisheries decker and erman 1992 and 2 way fish traps society 109484109log 484 490 CLAPP DFD F RDR D CLARK JR AND J S DIANA 1990 range al 1992 the prevalence of move- JS riley et activity and habitat of large free ranging brown trout ment in stream dwelling trout has begun to in a michigan stream transactions of the american receive greater recognition use of radioteleme fisheries society 1191022ilg119 1022 1034 try has enhanced the estimation of home range DECKER LML M AND DCD C ERMAN 1992 short term seasonal changes and abundance of fish and the distance that fish in composition size move but inad- in sagehen creek california transactions of the equate temporal sampling may still overlook american fisheries society 121297121 297997 306 certain fish movements for example a 576 mm EDMUNDSON E FEEE EVEREST AND DWD W CHAPMAN 1968 permanence brown trout was observed in the same daytime of station in juvenile chinook salmon and steelhead trout journal of the fisheries research 12 position from june 1994 to 10 february board of canada 25145325 1453 1464 1995 thus it had an overall daytime home EELLIOTTlliottlliotr J A 1973 the food of brown and rainbow trout range of zero but on 3 occasions it had diel salmo trutta and S gairdnerigairdneri in relation to the home ranges of 31 m 121 m and 141 m re- abundance of drifting invertebrates in a mountain stream oecologia 1232912 329 347 spectspectivelyively furthermore median summer and GATZ AJA J JR MJM J SALE AND JMJ M LOAR 1987 habitat mean diel home ranges 131 m v 105 m and shifts in rainbow trout competitive influences of distances moved 208 m v 274 m for brown brown trout oecologia 74774 7 19 trout were similar these results highlight the GERKING SDS D 1959 the restricted movement offish populations biological of diel review 3422134 221 242 importance monitoring in evaluations GOWAN C AND KDK D FAUSCH 1996 mobile brook trout of movement by stream fishes in two high elevation colorado streams evaluatingrereevaluating 244 GREAT BASIN naturalist volume 57

the concept of i estnctedrestricted movement canadian jour- PIMENTEL RA AND JD SMITH 1990 biostatbiossat I1 a uni-um nal of fisheries and aquatic sciences 53137053 1370 1381 banatevariatevanate statistical toolbox version 20 sigma soft GOWAN C MKM K YOUNG KDK D FAUSCH AND SCS C RILEY placentia CA 1994 the restricted movement of stream resident REGAL G E 1992 range of movement and daily activity salmonidssalmonids a paradigmpaipal adigm lost canadian journal of fish- of wild brown trout in the south branch auan sable erieserles aquatic sciences 51262651 2626 2637 river michigan fisheries researchReseaichsealch report 1988 hlsiiiagenhestfiagen T 1988 movements of brown trout salmo michigan department of natural resources ann trutta and juvenile atlantic salmon salmo salar in a arbor 46 appp coastal stream in northern norway journal of fish RILEY SCS C KDK D FAUSCH AND C GOWAN 1992 movement biology 3263932 639 653 of brook trout salvelinus fontinalisfontjontmalis in four small HILL J AND GDG D GROSSMAN 1993 an energetic model subalpine streams in northern colorado ecology of of micromiciomicrohabitatmiciohabitathabitat use foiroiforbolhorbor rainbow trout and rosyrosysideside freshwater fish 11121 ilg112 122 dace ecology 7468574 685 698 SOLOMON DJD J AND RGR G TEMPLETONTEMFLETON 1976 movements HUDSON JPJ P 1993 seasonal and daily movements of large of brown trout salmo trutta L in a chalk stream blownbrown trout in the mainstream au sable river journal of fish biology 94119 411 423 michigan fisheries researchReseaichsealch report 1998 michigan SPSS 1992 SPSSPCS PS spcbasebase system version 50155.0101 SPSS departmentdepaitment of natural resources ann arbor 65 appp inc chicago IL JINKINSTMJEINKINs TM JR 1969 night feeding of brown and rain-rain STEFANICH FA 1952 the population and movement of bow trout in an experimental stream channel jour- fish in prickly pear creek montana transactions of nal of the fisheriesfisherpisher lesies research board of canada 26 the american fisheries society 8126081 260 274 3275 3278 TIPPETS WE AND PB MOYLE 1978 epibenthic feeding KLEINKLFIN WD 1974 special regulations and elimination of by rainbow trout salmo gairdnergairdnengairdgairdnerinetinerinerlnen in the mccloud stocking influence on fishermen and the trout popu- river california journal of animal ecology lation at the cache la poudre river colorado state 4754947 549 559 of colorado division of wildlife technical publica- WARNER EJE J AND TPTIP QUINN 1995 horizontal and verti- tion 30 WRTW R T 30 57 appp cal movements of telemeteredtelemetered rainbow trout oncor- matthewsMAITHLWSMATThewsnEws KRK R NHN II11 BERGBLRG DLD L AZUMA AND TR lam- hynchus mykiss in lake washington canadian jour- imablin 1994 cool watelwater formationfoiroibol mationmatlon and trout habitat nal of zoology 7314673 146 153 use in a deep pool in the sierra nevada california WILKISON RAR A 1996 fish community structure and transactions of the americanamerlean fisheries society brown trout predation in upper silver creek idaho 123549123 549349 564 unpublished master s thesis idaho state university mclniosilmcintosfl ARA R AND CRC R TOWNSEND 1995 contrasting pocatello 128 appp predationpiedation risks presented by introduced blownbiownbrown trout YOUNG MKM K 1994 brown trout mobility in south central and native common tiverriverilverliver galaxias in new zealand wyoming streams canadian journal of zoology 72 stistreamsearnsearms canadian journal of fisheriesFishenes and aquatic 2078 2083 sciences 52182152 1821 1833 1995 telemetry determined diurnal positions of MENSEml nseNSL JBJ B 1975 relation of density to brown trout brown trout salmo trutta in two south central movement in a michigan stream transactionstiansactions of the wyoming streams american midland naturalist 133 americanamerlean fisheries society 104688104 688 695 264 273 MEYERSMLYLKS LS TE1 F THUEMLER AND GWG W KORNELY 1992 1996 summer movements and habitat use by seasonal movements of blownbiownbrown trout in northeast colorado river cutthroat trout Oncooncorhyoncorhynchusrhynchusachus clarki wisconsin noinorthth american journal of fisheries pleuriticpihunpleunpleunticuspleuriticusticusficusus in a small montane stream canadian management 1243312 433 441 journal of fisheries and aquatic sciences 53 millekMILLLKMILLER RBR B 1957 permanence and size ofhomeof home territory 1403 1408 in stream dwelling cutthroat trout journal of the YOUNG MKM K RBR B RADER AND TA BELISH in press fisheriesFishenes research board of canada 1468714 687 691 influence ofmacroinvertebrateofmacromveitebiate drift and light on the NILLSLNJLNIELSEN JL FETE LISLELISLL AND V OZAKI 1994 thermally activity and movement of colorado river cutthroat stratified pools and thentheir use by steelhead in northern trout oncorhynchus clarki pleuriticpleunpleunticuspleuriticusticusus transactions california streams transactions of the americanamerlean oftheofodthethe american fisheries society fisheriesFishenes society 123613123 613 626 northcote TG 1992 migration and residency in stream received 25 november 1996 salmonsaimonsalmondssalmomdssalmonidsids some ecological considerations and evo- accepted I1 may 1997 hitionarylutionallutiutionallonallonai y consequences nordic journal of freshwater research 67567 5 17 great basin naturalist 573 C 1997 appp 245 252

SHOREBIRD PREDATION ON BENTHIC macroinvertebratesMACROMACROIN vertebratesINVERTEBRATES IN AN irrigation RESERVOIR

janet R mihucl2mihuc12 charles H Trostltrostletrostl3trost133 and timothy B mihucl4mihuc14

ABSTRACT american falls reservoir in southeastern idaho is an irrigation reservoir used as an inland feeding stopover by many shorebird species six exclosure experiments weiewelewere conducted during the 1990 drawdown period to investigate shorebird predation impact on benthic macroinvertebratemacromaeromaciommaciog vertebrateinvertebrate populations the study sites differeddifdlffeiedhered in sediment composition sediment slope invertebrate densities and shorebirdshoiesholebird abundance shorebird predation significantly affected invertebrate densities in only I1 of 6 experiments aberdeen mouth this site had higher sediment slope and slower water recedence than other study sites resulting in concentration of shorebird predation on a smallersmailer aleaarea of newly exposed sediment shorebird predation had the greatest impact on medium size class chironomid larvae at aberdeen mouth our results suggest that inland sites such as american falls reservoirReservon repierepresentsent viable shorebird habitat and may be managed to insure consistent pleyprey availability drawdown rate sediment slope invertebrate densities and shorebird abundance are all important factors influencing shorebirdshoiesholebird predation monitoring shorebird abundance and predation impact on invertebrate densities may help in manipulating drawdown latetaterate to provide adequate shorebird pleyprey management of inland sites for shorebird use may become moiemolemore important in the future as human encroachment in coastal areas continues

key words benthic macromacroinvertebratesmacromvertebratesinvertebrates exclosuresenclosures idaho management predation shorebirdsshore birds

reliable and food rich staging areas are quammen 1981 schneider and harrington essential for migrating shorebirdsshorebirds senner and 1981 raffaeli and milne 1987 in this study howe 1984 myers et al 1987 paulson 1993 exclosures were used to assess experimentally although coastal staging areas support the the impact of shorebird predation on benthic largest numbers of migrating shorebirdsshorebirds many macroinvertebratesmacro invertebrates at several american falls inland staging areas exist and may become reservoir sites sites differed in sediment com- more important as human encroachment upon position sediment slope and rate of sediment coastal areas continues skagen and knopf exposure investigating the impact of shorebird 1993 knowledge concerning shorebird inland predation at different sites may help identify use is limited compared to coastal staging areas physical factors that influence predation the focus of this study was to quantify shorebird use of food resources at a freshwater in- STUDY AREA land staging area american falls reservoir idaho the annual presence of 30000 indi- american falls reservoir is an east west viduals and 30 species of shorebirdsshore birds during oriented shallow depth impoundment located fall migration has been documented at this on the snake river southeastern idaho the reservoir taylor et al 1992 common probing reservoir is part of the bureau of reclama- or benthic feeding species using the reservoir tion s Miniminidokadoka project that provides irrigation include baird s sandpiper calidrisCalidris bairdi water to thousands of hectares of land in western sandpiper calidrisCalidris mauri long southern and eastern idaho the snake river billed dowitcher limnodromus scolopaceusscolopaceous enters the reservoir in the springfield bottoms lesser yellowlegs tringa flavflavipesipes and mar- at the northeastern end and exits through a bled godwit limosafedoa dam at the southwestern end located at an ele- closuresexclosuresenclosuresEx are commonly used to assess vation of 1328 inm the reservoir at full capacity shorebird predation in marine and estuarine is 35435.4 km long has a surface area of 23503 ha environments schneider 1978 bloom 1980 and has 161 kinkm of shoreline annual drawdown

idepartmentepaitment of0 biological sciencesciences idaho state university pocatello ID 83209800783209 8007 present21res t addiesaddlesaddress 4718 north alby street godfrey IL 62035 hauthor3authorauthoiautholauchoi to whom con e&pondencecorrespondence should be sent 4presedtniesentplesentPiepiesentsent addlessaddress illinoisIlli1111 noinol natural history survey creatgreat riverbiverriversbivers field station LTRMP 26 4134 alby street alton IL 62002

245 246 GREAT BASIN naturalist volume 57 typically begins in april and continues through 1328 september fig 1 and is most rapid between 1326 june and late august when irrigation water 1324 demand is greatest several kilometers of mud flats consisting mainly of sand and silt are ex- 1322 posed at this time 1320 six experiments were conducted at the reser- 1318 voir in summer 1990 table 1 study sites dif- 1316 fer in sediment composition and sediment slope and were selected because of observed 1314- shorebird foraging activity in the area the 1312 1 slope of the sediment tan depth of place- 1310 ment m distance from shoreline m ranges JFJ F M A M J J A S 0 N D from 0350.35 to 2692.69 table 2 the bronco site month is on the east side of the reservoir two experiments were conducted at aberdeen bay on the fig 1 daily average water level at american falls dam west side of the reservoir one at back bay at from 1 january to 31 december 1990 the back of the bay and the other at aberdeen mouth at the mouth of the bay willow and silo sites are at the southern end of the reser- bance between treatments and the shoreline voir within 5 km of the dam two experiments the 6 treatment sets were placed in a row par- were conducted at the willow site allel to the shoreline in the same water depth fig 2 arrangement of the exclosure and 2 METHODS controls in each treatment set was random placement depth for treatment sets in each for each experiment at each site we placed experiment ranged from 19 to 40 cm table 1 treatment sets in water depth inaccessible to reservoir drawdown was constant and aver- probing shorebirdsshorebirds and conducted benthic sam- aged 14 3 s cadaycmday during the study period pling the day after water had receded beyond july to mid august 1990 the rapid drawdown in the sets being sub- the treatment sets A treatment set included I1 resulted treatment merged 1 4 d each 1 exclosure I1 open control and I1 exclosure con- at site table invertebrates trol quammen 1981 fig 2 six treatment sets benthic macromacroinvertebrates were sampled the after the had completely were used during each experiment the open day water receded beyond all treatment sets because treatment control was marked by 4 wooden stakes the sets were placed parallel to the shoreline all exclosure control which consisted of a top and became exposed at the same time and were I1 side to allow shorebird access was used to sampled simultaneously according to the fol- account for any influence the exclosure itself lowing procedures we took randomly spaced might have on shorebird predation exclosuresenclosuresEx closures cores of sediment with a 5 cm diameter 10 cm sides consisted of 4 and a top closuresExexclosuresenclosures and long plastic core tube two cores were taken exclosure controls were constructed of hard- from each exclosure and control and then com- 1 cm2 ware cloth mesh stapled to wooden bined to represent a single sample generating stakes 65 cm long the area within each ex- 6 samples for each treatment type exclosure closure or control was 0250.25 m2ma exclosure control open control during each we carried treatment sets from the water s experiment sediment cores taken in the field edge and placed them underwater 858.5ss 37 m were placed in plastic sample bags and either from the shoreline table 1 by pushing each sorted or frozen as soon as possible before set into the sediment until the bottom edge of sorting we sievedsievek samples 025 mm mesh the hardware cloth was at least 2 cm below the sieve leaving only invertebrates and organic sediment surface wooden stakes of the open matter all invertebrates were sorted from the control were pushed down to a depth equal to samples preserved in 95 ethanol and later the other 2 treatment types we carried all identified and counted with the use of a binoc- treatment sets to the placement area from a ular dissecting microscope three distinct size downshore point to minimize sediment disturdastur classes of chironomid larvae differentiated by 199711997 SHOREBIRD PREDATION ON BENTHIC invertebrates 247

TABLE 1 summary of experimental design parameters including water depth in which treatments were placed and initial distance of treatments from shoreline placement sampling placement distance from site name date date depth cm shore m bronco 18 july 20 july 23 37 back bay 26 july 28 july 40 85 silo 6 august 9 august 40 30 aberdeenAbeideen mouth 9 august 14 august 27 10 willow experiment 1 14 august 16 august 19 31 willow experiment 2 18 august 20 august 22 35

TABLE 2 sediment slope and sediment composition at each study site sediment silt sand gravel site name slope 62 mm 62 161.61iglg 6 mm I1 161.6lgig16mmmm bronco 035 288 712 0 back bay 269 627 372 01 silo 076 590 406 04 aberdeen mouth 155 966 34 0 willow experiment 1 036 734 266 0 willow experiment 2 036 782 218 0

ac 0c E EC ffifaffif 97n E EC feisfets ftrrfarr K 99111 0cac FTFN NI I1 liiIII111 X ai1i L lil 1 IT U

distance from shoreline shoreline

fig 2 depiction of the placement of 2 treatment sets parallel to the shoreline A treatment set consisted of I1 excio sure E 1 exclosure control EQEC and 1 open control OC ordering of the exclosure and controls in each treatment set was random head capsule size and body length small 5 was calculated beginning the day after sets mm medium 5 10 mm large 10 mm were were placed and continuing through the day counted separately we took I1 sample from each benthic sampling was conducted direct counts study site and analyzed it for sediment compo- were made of all probing species in a 100loom m sition this sample was wet sievedsievek through 2 area of shoreline in front of the treatment sets sieves to separate it into gravel iglg 161.6 mm this area was designated by 2 small flags placed sand 0620620.62 mm and silt 0620620.62 mm compo- at both ends of the transect for a period of 30 nents after removing invertebrates we dried min each morning we counted all shorebirdsshorebirds the sample at 60c60goc C for 72 h and weighed it at 5 min intervals wilson 1988 counts of prob- percentages of gravel sand and silt at the study ing species were used to calculate a mean and site were then calculated from the components standard deviation which when doubled rep- and total weight of the sample table 2 resented an hourly estimate multiplying the A daily estimate of shorebird abundance in hourly estimate by 24 yielded a daily estimate of the immediate vicinity of the treatment sets shorebird abundance during each experiment 248 GREAT BASIN naturalist volume 57

20000 20000 BRONCO SILO

15000 15000 JL 10000 10000 5000 i 5000 B6 20000 20000 BACK BAY ABERDEEN MOUTH

15000 15000

W 10000 10000

5000 t 5000 h i

20000 20000 WILLOW 1 WILLOW 2

15000 15000

10000 10000

5000 5000

I1 I1 exclosure XA exclosure control gi open control

fig 3 mean total invertebrate densities in exclosures and controls during each experiment error bars indicate 1 standaidstanstandarddaiddald deviation from the mean an asterisk indicates a significant difference FP 005oos0 05 between total invertebrate densities in exclosures and controls comparison of shorebird abundance across all analysis to address the possibility of shorebird study sites was possible because daily estimates feeding preference for either size class and a were based on counts taken at a similar time at linear regression to investigate the relationship each study site between sediment composition and inverte- an independent t test was used to compare brate densities at each site invertebrate density data from the 2 types of controls addressing the effect of exclosure RESULTS presence on shorebird predation an ANOVA was used to compare invertebrate density data benthic prey items available to shorebirdsshorebirds at flomfrom all 3 treatment types at each site total american falls reservoir included chironomid invertebrate density data were analyzed as well larvae and tubificid worms invertebrate den- as large and medium size class density data for sities differed across study sites fig 3 most chironomid larvae we used size class data chironomid larvae consisted of chironomus sp 199719971 SHOREBIRD PREDATION ON BENTHIC invertebrates 249

12000 1200012000i BRONCO SILO T 10000 10000 1 I1 8000 8000 6000 T 6000 li 4000 A 1 aalapl 4000 li 2000 II11 2000 0 0 large medium large medium

12000 12000 BACK BAY ABERDEEN MOUTH S i uQ 10000 si 10000 sQ S Q 8000 9 8000 S A PC 15 6000 0oPC 6000 6 0 4000 0 4000 1 B f 2000 2000 u 1 & 4 anaAHAL ajia I li S 0 0 40 ouloui0 ft large medium largelailarlarilarj jg medium

12000 WILLOW 1 12000 WILLOW 2 10000 10000

8000 8000

6000 6000 T

4000 4000 ABIH aljkij T 2000 2000 i A IHN ihmIBM11 iiilii111 0n m ihn 0n large medium large medium

I1 I1 exclosure izavya exclosure control open control

fig 4 mean densities of large size class chironomid larvae and medium size class chironomid larvae in exclosures and controls during each experiment error bars indicate I1 standard deviation from the mean an asterisk indicates a sig- nificantnificant difference P 005oos0 05 between chironomid densities in exclosures and controls

90 95 but some Proprocladiuscladius sp also were of chironomid larvae were found at the silo found chironomidChiYonochiyonomidmid larvae were the predomi- site the smallest number at back bay nant benthic prey item constituting 40 100 no differences P 0050.05oos in total invertebrate of total benthic invertebrates found in excio densities occurred between the 2 control types sures during each experiment the majority of in any experiments A difference P 0050.05oos chironomid larvae in samples were represented between total invertebrate density among the by the large and medium size classes individ- 3 treatment types occurred only during the uals of these 2 size classes were 1 2 minmm larger aberdeen mouth experiment fig 3 com- in diameter than tubintubincidtubificideidcideld worms densities of parison of chironomid size class data from the large and medium size class chironomid larvae treatment types revealed a difference P were similar at each site but differed some- 0050.05 only in the medium size class of chiranochirono what across sites fig 4 the largest number mids at aberdeen mouth fig 4 the impact 250 GREAT BASIN naturalist volume 57 of shorebird predation on the large size class was found only at aberdeen mouth shorebird of chironomids at this site was noticeable but predation at this site had a greater impact on not significant P 0100.10olo at the silo site the medium size class chironomids fig 4 sug- medium size class of chironomids suffered gesting a feeding preference for size class the greater predation impact than the large size possibility of feeding preference should not be class but this difference was not significant P overlooked in management decisions concern- 0100.10olo fig 4 ing shorebirdsshorebirds further study of shorebird sand and silt dominated all sites table 2 preference for prey size or species in fresh- the percentage of these 2 fractions varied water environments is needed from site to site but linear regression results comparing conditions at the aberdeen indicated no effect of sand r 0220.22 P 0050.05oos mouth site to those at other study sites was or silt concentration r 0220.22 P 0050.05oos on useful in identifying factors that influenced benthic invertebrate densities because of the shorebird predation impact during this study rapid water drawdown during the experiments at aberdeen mouth treatment sets were ex- more than 10 m of sediment was exposed or posed to shorebird predation at least 24 hb under shallow water 5 cm each day at 4 of longer than at other sites because the mouth the 5 sites at aberdeen bay where 2 experi- of the small bay has a steeper sediment slope ments were conducted back bay aberdeen than all other sites except back bay the com- mouth 0 5 m of sediment was exposed each bination of steeper slope and drainage of water day daily estimates of shorebird abundance from the bay resulted in slower water rece- varied widely ranging from 14 during the silo dence shorebird predation at the aberdeen experiment to over 6000 during the bronco mouth site was concentrated on 0 2 m of experiment table 3 freshly exposed sediment each day rather than 5 10 m of freshly exposed sediment typical of discussion the other study sites the longer exposure and management of american falls reservoir concentrationconeconceoneentrationentration of shorebird predation on a of sediment probably for for irrigation purposes has created a unique smaller area accounts and dynamic environment with gradually slop- the observed impact on invertebrate densities ing sediment constant rate of summer draw- at the aberdeen mouth site although sediment down and little submerged vegetation taylor slope was steepest at the back bay site table et al 1993 these characteristics represent ideal 2 the minimal impact of shorebird predation habitat for foraging shorebirdsshorebirds during migra- was probably due to much lower invertebrate tion rundle and fredrickson 1981 kushlan densities fig 3 and shorebird abundance 1989 helmers 1991 shorebird abundance table 3 compared to the other study sites data were not useful predictors of predation our results suggest that sediment slope inver- impact on invertebrate densities at the study tebrate densities and shorebird abundance sites this may be because birds were foraging should all be monitored and considered in over large areas of freshly available sediment combination when making management deci- each day and counts were taken during only I1 sions regarding shorebird predation window of time daily significant shorebird conditions at american falls reservoir are predation effect on benthic prey populations a contrast to conditions in coastal areas the

TABLE 3 daily abundance estimates mean standard deviation of shoreshorebirdsbirds during each experiment numbers of counts reflect different lengths of each experiment site name daydayldayiI1 day 2 day 3 day 4 bronco 6034 1482 630 199 back bay 15 1 60 44 silo 14 36 240 40 41 109 aberdeen mouth 1008 469 254 151 117 67 103 81 willow experiment 1 3408 1761 2722 siisli51151111 willow experiment 2 199 216

listlasthist day linsinvin veyedkeyedyed 199719971 SHOREBIRD PREDATION ON BENTHIC invertebrates 251 latter are characterized by many shoreshorebirdsbirds acknowledgments feeding on a limited shoreline exposed during low tide myers 1983 burger 1984 coastal this research conducted as part of a mas- studies have documented a significant impact ter s thesis was funded by grants from the of shorebird predation on invertebrate densi- bureau of reclamation sigma xi and idaho ties schneider 1978 schneider and harring- state university graduate student research ton 1981 quammen 1984 at american falls and scholarship committee dr L ferrington reservoir the constant summer drawdown rate provided firmationformationconfirmationconeonfirhirbirmationmatlon of identification of chi- provides large areas of newly exposed sediment ronomid larvae dan taylor and 3 anonymous daily and shorebird densities are lower than at reviewers provided valuable critiques of the coastal areas our results suggest that current manuscript prey densities in both sandy and silty mudmudflatsflats are adequate to support shorebirdsshorebirds using amer- literature CITED ican falls reservoir this potential is encour- BLOOM SAS A 1980 an intertidal soft sediment avian exclo- aging because if shorebird densities were to sure which minimizes sediment alterations marine increase use of the traditional feeding areas ecology progress series 3793 79 81 might gradually increase over time myers et al BURGER J 1984 abiotic factors affecting migrating shore birds pages 1 71 in J burger and B olla editors 1987 further study is although of inland sites behavior of marine animals volume 6 shorebirdsShorebirds needed conservation and management of these migramigiamigrationtion and foraging behavior plenum press new sites should continue york our results and those of other studies have HANDS HMH M MRM R RYAN AND JW SMITH 1991 migrant shorebird use of marsh moist soil and flooded agri- identified criteria for evaluating inland areas cultural habitats wildlife society bulletin 1945719 457 464 as potential shorebird staging areas impound- HELMERS DLD L 1991 habitat use by migrant shorebirdsshorebirds ments currently used for irrigation purposes and invertebrate availability in a managed wetland may need only adjustments to complex unpublished mastermastey s thesis university of minor accom- missouri modate migrating shorebirdsshorebirds A gradually slop- KUSHLAN JAJ A 1989 avian use of fluctuating wetlands ing silty or sandy sediment with little or no pages 593 606 in RRR R shantzsharitz and JWJ W gibbons vegetation is most favorable helmers 1991 editors freshwater wetlands and wildlife DOE sym- no 61 S an annual drawdown would favor coloni- posium series USDOE office of scientific prey and technical information oak ridge TN zation by tubificid worms and certain chirono- MYERSMYEBS JPJ P 1983 conservation of migrating shorebirdsshorebirds mid species and discourage colonization by staging areas geographic bottlenecks and regional aquatic vegetation helmers 1991 the drop in movements american birds 372337 23 29 MYERS P AL water level must coincide with known JPJ ET 1987 conservation strategy for migra- migra- tory species american scientist 751975 19 26 tion periods of shorebird species that would PAULSON D 1993 shorebirdsShorebirds of the pacific northwest potentially use the area hands et al 1991 tay- university of washington press seattle lor et al 1993 A gradual but continual drop in QUAMMEN MLM L 1981 use of exclosures in studies of predation by shorebirdsshorebirds on intertidal mudmudflatsflats auk 98 water level would insure a renewing source of 812 817 available prey rundle and fredrickson 1981 1984 predation of shorebirdsshorebirds fish and crabs on kushlan 1989 prey densities will likely vary invertebrates in intertidal mudmudflatsflats an experimental across space and time and should be monitored test ecology 6552965 529 537 RAFFAELLI D AND H MILNE 1987 an experimental throughout the migration period ability to investigation of the effects of shorebird and flatfish manipulate the drawdown rate on a seasonal predation on estuarineestuanne invertebrates estuarineEstu anne or yearly basis is important faster drawdown coastal and shelf science 24124 1 13 may be necessary to compensate for sediment RUNDLEBUNDLE WD AND LH fredrickson 1981 managing seasonally flooded impoundments for migrant rails slope in some impoundments the drawdown and shorebirdsshorebirds wildlife society bulletin 9809 80 87 rate may also be manipulated to accommodate SCHNEIDER D 1978 equalisationequalization of prey numbers by numbers of shorebirdsshorebirds using the area in rela- migratory shorebirdsshorebirds nature 271353271 353 354 SCHNEIDER D C AND B A harrington 1981 tion to densitiestodensities of available prey monitoring DC BA timing of shorebird migration in relation to pleyprey depletion the impact of shorebird predation on inverte- auk 9880198 801 811 brate populations could be accomplished using SENNER SES E AND MAM A HOWE 1984 conservation of exclosures thus providing data for decisions neanearcticreticretieretle shorebirdsshorebirds pages 379 421 in J burger and B olla editors behavior of marine animals volume concerning drawdown rate skagen and knopf 5 shorebirdsShorebirds breeding behavior and populations 1993 plenum press new york 252 GREAT BASIN naturalist volume 57

SKAFNskagenSKAGLN SKS K ANDAN FLEL KNOPIKNOPE 1993 toward conservation WILSON WH JR 1988 methods of measuring wader of continentalmidcontinentalmid shorebird migrations conserva- abundance wader study group bulletin 5449 50 tion biology 75337 533 541 IAYLORTAYLOR DMD M CHC H trosiTKOSITROST AND B JAMISON 1992 abun- received 24 may 1996 dance and chionchlonchionologychronologyology ofofmigiantmigrant shorebirdsshorebirds in idaho accepted 21 april 1997 westeinwestern budsbirds 234923 49 78 1993 migrantmiglmigi ant shorebird habitat use and the influence ofofwateiwater level at americanamerlean falls reservoir idaho northwest naturalist 743374 33 40 great basin naturalist 573 0 1997 appp 253 258

lagomorphslagomorpha AND THE DISPERSAL OF SEEDS INTO communities DOMINATED BY EXOTIC ANNUAL WEEDS

eugene W Schuppschupplischuppl2schupp1212 hoyt J heatonl3Heatonheaton1313 and josejosa M g6mezl4gomez14

ABSTRACT large areas of western rangeland are presently dominated by alien annual weeds such as bromus fectotecto rum cheatcheatgrassgrass these communities resist succession to perennial communities primarily because the annuals are com- petitpetitivelyively superior to establishing perennial seedlings and they promote fires that favor weeds over perennials succes- sionslon may be further slowed however by low rates of seed dispersal into annual grasslands we investigated the role of lagomorphslagomorpha sylvilagusSylvilagus nuttalnuttalliinuttallialii nuttall s cottontail S audubonauduboniiandubomiandubomiii desert cottontail and lepus califorcalifomicuscalifornicusnicus black tailed jackjackrabbitrabbit in seed dispersal across an ecotone between an open juniper woodland and an annual grassland we collected pellets along five 100 x 2 m transects parallel to the ecotone 50 m into woodland border and 20 m 50 m and 100 m into grassland we searched pellets for juniper seeds visually and for any other species through germination from crushed pellets after cold moist stratification pellets were not evenly distributed across transects but there was no trend with respect to position of transect juniperusJumperus osteospennaosteosperma utah juniper was the most abundant seed both the number of juniper seeds and the proportion of pellets with juniper seeds decreased steadily from a high in woodland to absence at 100 m into grassland only 2 dicot seedlings emerged from pellets I1 salsola pespettiferpestifertifer and 1 unknown that died prior to identification consequently there was little seed movement into the grassland 72 of all seeds were collected from either woodland or border transects lagomorphslagomorpha apparently do not effectively replenish the native perennial seed pool of cheatcheatgrassofcheatgrassgrass dominated disturbances at dugway

key words juniperus osteosperma seed dispersal lagomorphslagomorpha bromus tectoriumtectorumtectorum range restoration degraded rangeland

overgrazing fires and other disturbances although competition and increased fire from human activities have degraded exten- frequencies make it difficult for native species sive areas of native vegetation in the inter- to reestablish given enough time between fires mountain region of the western USA leading at least some native species such as sitanionSitanion to the domination of many rangelands by alien hystrix bottlebrushbottlebrush squirreltail hironaka and annual weeds such as bromus tectoriumtectorumtectorum cheat sindelar 1973 and agropyron smithiismithia western grass taeniatherumtheniatherumTaeniaTheniatherum esperumasperum medusaheadmedus ahead wheatwheatgrassgrass monsen 1994 appear able to and salsola pespettiferpestifertifer russian thistle billings invade these weed communities in addition 1990 young 1994 over 131.3 million ha are com- to competition and repeated fires then succes- pletely dominated by B tectoriumtectorumtectorum or T esperumasperum sion to native perennial rangelands may be and 30830.8 million ha more are infested or sus- partly limited by low seed availability due to cepceptibletible to invasion pellant and hall 1994 depletion of the native seed bank over time once established these annual grasslands large- pyke 1994 LD humphrey and EW schupp ly resist succession to native rangelands be- unpublished data and potentially low levels of cause the weeds are highly competitive with dispersal into the annual grasslands establishing perennials monsen 1994 pyke and lagomorphslagomorpha however may effectively dis- novak 1994 young 1994 and because of the perse seeds from native rangelands into annual initiation of a cheatcheatgrascheatgrassgras s wildfire cycle where- grasslands they range widely feeding in shrub- in annual weeds promote fire that favors the land woodland and grassland smith 1948 further spread of weeds and thus of more fires Kundakundaelieli and reynolds 1972 westoby and wag- billings 1990 whisenant 1990 peters and ner 1973 mcadoo et al 1987 zeveloff 1988 bunting 1994 smith 1990 and in the process disperse viable

apaiidepartmentepai tmentament of rangeland resources and the ecology center utah state university logan UT 84322 USA 2addressaddie&iiwaddress correspondence to this author at departmentdepar tmentament of rangeland resources utah state university logan UT 84322523084322 52306230 spiesent3presentSpie sent addressaddless department of botany and plant sciences university of california riversideRive iside CA 92507 4presentaddress4plcsent addless depaitamentodepartamentoDepartamento de BiologiA animalammal y Ececologiaecologicologia racultfacultadad de cienciakcienciasCiencias umwsidadumversidad de Gigranadaanada E 18071 Gigranadamadamadd spain

253 254 GREAT BASIN naturalist volume 57 seeds 1 of grasses and forbs consumed inci-inciinel the native perennials listed from the burn all dentally while feeding on foliage welch 1985 annual weeds from the burn are also found in zedler and black 1992 malo and suarez 1995 low densities scattered through the adjacent malo et al 1995 and 2 of fleshy fruited plants woodland the lagomorphslagomorpha sylvilagusSylvilagus nuttalnuttalliinuttallialii whose fruits are intentionally consumed smith nuttall s cottontail S audubonii desert cot- 1948 dantonioDAntomo 1990 schupp et al 1996199619971997 tontontailtail and lepus califomicuscalifornicus black tailed the objective of this study was to examine jackrabbitjackrabbit are present at the site the role lagomorphslagomorphaoflagomorphsof in the dispersal of seeds across an ecotone between a relict open juni- METHODS per woodland and a burned area dominated by exotic annual weeds we were especially in february 1995 we placed a transect 100 concerned with 1 the diversity and quantity in long X 2 in wide in the annual grassland of native and exotic seeds found in pellets and directly adjacent to the border of the juniper 2 the spatial pattern of native seed deposi- woodland transect 0 successive transects tion in the grassland relative to distance from at 20 in 20 50 in 50 and 100 in 100 the woodland into the grassland and a final transect 50 in into the woodland 50 all transects were STUDY SITE parallel to the border after clearing S pespettiferpestifertifer skeletons with a pitchfork we collected all the study site is an open juniperjumper woodland intact lagomorph feces pellets encountered and adjacent annual grassland the U S armarmy on US y in each transect after completing collections grounds county dugway proving thoeletooele utah we realized that we had used a more thorough USA 40 N at an elevation of 4015n 11250w technique on the 100 in transect pulling up roughlylawlwwwhere old bonneville 14604460 in lake cheatcheatgrassgrass to expose pellets trapped within sand dunes meet the lower slopes of the cedar clumps of vegetation or within the upper few mountains climate is andaridarld with mean the cm of soil consequently we collectedrecollectedre re- annual precipitation of 19219.219 2 cm bagley 1991 maining transects with the same thoroughness aerial photographs show that wildfire con- decomposition of lagomorph pellets in andaridarld verted a shrubland adjacent to the woodland environments is slow in west texas flinders into an annual grassland sometime between and crawford 1977 estimated time for com- 1978 and 1985 R johnson personal commu- plete decomposition to be 444.4 yr for L califor ninication the burn was seeded with agropy- pellets and 95gs for S audubonii pellets ron cristatumlacnstatuma desertorumdesertorum crested wheat nicus 959.5 yr audubonii these collections long- grass and kochia proprostrateprostratastrata prostrate kochia consequently represent after the fire but the seeding was not very term patterns of deposition suggesting the for single rela- successful and densities of these exotic peren- results are at least this site nials are low native perennials such as the tively robust accumulations are not so long from grasses oryzopsis hymenoideshymenoides indian ricriericegrassncncegrassegrass term however that collections the annual and sporobolus cryptcryptandrusandrus sand dropseed grassland 10 yr old would be biased by in- and the forbs sphaeralcea munroanamunroana munroe s clusion of significant numbers of relict pellets globemallowglobemallow and oenothera hallidapallida pale eve- deposited in the former shrubland pellets ning primrose are also present at low densities were kept in plastic bags in a refrig- the overwhelmingly dominant species how- erator until processing because larger pellets ever are exotic annual weeds mostly B fectotecto are more likely to contain seeds than small rum with variable quantities of S pettiferpespestifertifer and pellets EW schupp M fuentes and JM sisymbrium altissimum tumbling mustard gomez unpublished data we randomly se- vegetation within the adjacent woodland lected 25 pelletstransectpellets transect and measured length consists of scattered large juniperus osteoosoteo to the nearest 0010.1oiol 1 mm with dial calipers to sperma utah juniper with a well developed compare pellet size across transects we then shrub understory of predominantly atriplex processed the entire sample from each tran- canescentcanescenscanescens four wing saltbush and sarcobatus sect by cleaning the surface of each pellet with vermiculatus greasewood and a diverse her- a stiff camel hair brush to dislodge any seeds baceous layer including erysimum esperumasperum clinging to the surface and then crushing all wallflower eriogonum sppapp buckwheats and pellets to search for the relatively large and 199711997 LAGOMORPH SEED DISPERSAL AND ANNUAL grasslands 255 obvious seeds of juniperjumper and the shrubs we in number of pellets number of seeds and did not open juniper seeds to determine percent proportions of pellets with seeds as functions filled because the only cone crop available for of distance from the woodland transect we dispersal in the 2 yr preceding this study had analyzed differences among transects in pellet 1 filled seeds when mature EW schupp size length with one way ANOVA followed unpublished data because lagomorphslagomorpha do not by a tukey HSD post hoc test to determine appear to discriminate among cones based on which transects differed because a normal seed filling EW schupp M fuentes and JM probability plot suggested the data were nor- gomez unpublished data results should also mally distributed and a bartlett s test x2oc2xa 4 be representative of years with high levels of 61226.122 P 0190olgo0.190 demonstrated homogeneity filled seeds of group variances we used untransformed due to the difficulty of locating small seeds data analyses were performed with SYSTAT in fibrous remains of pellets we searched for 505.0so wilkinson 1990 the presence of species other than juniper with germination tests after breaking pellets apart RESULTS further we spread them on wet washed sand in 0250.25 x 050osom m plastic nursery trays wet the we collected a total of 8425 pellets from pellet materials covered them with plastic lids the 1000 m2ma sampled pellets were not evenly to prevent drying and placed them in a refrig- distributed across transects xax2 L4 79797797.97 erator at 3 4c4ac C from 16 june to 28 august P 0 000010.0001 there was however no relation- 1995 73 d on 28 august we placed the trays ship between number of pellets and distance in a growth chamber with a 24 h cycle of 12 h from the woodland transect r 02000.200 n 5 light at 15c15 C and 12 h dark at 10c10log C after 5 wk P 050.5os the greatest number of pellets was we increased temperatures to 20c20 C and WC15 C found at 20 m and the least at 100 in respectively but terminated the experiment table 1 after 3 d because temperature began fluctuat- pellet size as measured by length differed significantly ing wildly while the experiment was in pro- among transects faf4fr4 1201Q 11811.8 FP gress we misted pellet material daily and trans- 00010.001 the difference was dedue entirely to planted emerging seedlings to small pots to the 50 transect having larger pellets than all grow until identified other transects which did not differ from each we used chi square goodness of fit tests to other table 1 analyze distributions of pellets and seeds across fifty seven pellets 07070.7 contained a total transects with an expectation that they would of 61 whole and apparently undamaged juni- be evenly distributed and a likelihood ratio per seeds I1 contained 3 seeds 2 contained 2 chi square test of a 2 x 5 contingency table to seeds and the remainder contained a single analyze for differences among transects in pro- seed an additional 17 pellets contained broken portions of pellets with and without seeds seeds or seeds separated along the seam joining spearman rank correlations tested for trends the 2 halves of the seed coat no other large

TABLE 1 number of pellets mean pellet length arnmmmrn number of pellets with whole seeds proportion of pellets with whole seeds and number of whole seeds by transect for pellet length values followed by the same letter are not significantly different at P 00500.05oos05 based on a tukey HSD test pelletspelpeibetshets length number proportion number transect number mean s with seeds with seeds ofseedsof seeds

WOODLAND 50 1588 iiilii11111 1 13131.31 3 a 25 00157 28 ECOTONE 0 1436 989.89 8 14141.41 4 b 16 00111 16 GRASSLAND 20 2597 959 5 1 1 b 11 00042 12 50 1777 909 0 090 9 b 5 00028 5 100 1027 979.79 7 101 0 b 0 00000 0 256 GREAT BASIN naturalist volume 57 seeded species were found in the pellets con- seeds were apparently old unfilled ones begin- sisidering only whole seeds a 2 X 5 contingency ning to split along the seam after several years table analysis showed the proportion of pellets in a pellet containing seeds differed among transects the percentage of pellets with juniper seeds 2 likelihood ratio x2xa 4 384438.4438 44 FP 00010.0010 ooi001 table in this study was considerably lower than the 1 similarly a chi square analysis showed that 4 7 reported for the 2 nearby woodland sites numbers of whole seeds were not evenly dis- schupp et al 1996 this is probably a func- tritributed across transects x2oc2xa 4 381038.1038 10 FP tion of irregular fruiting of JJ osteosperma 00010.0010 ooi001 table 1 in contrast to the number of while pellet collections for the previous study pellets both the proportion of pellets with seeds corresponded to a period of heavy fruitfruitfallfall the and the number of seeds decreased steadily present study includes pellets deposited over with increasing distance from the woodland rfg many years encompassing years with both loo1001.001 00 n 5 P 005oos0.050 05 inm both cases table heavy and light fruit production even if more 1 analyses using all seeds whole broken seeds were dispersed however few would be yielded virtually identical results dispersed much beyond the boundary of the only 2 seedlings emerged from the rabbit woodland 72 of all seeds were found in either pellet fragments during germination tests after the woodland transect or directly along the 7 d 1 S pespettiferpestiferpesttfertifertiner emerged from the 20 transect border and the number of seeds dropped off material and after 35 d I1 unidentified dicot rapidly to zero at 100 in emerged from the 100 transect material but reasons for this pattern of dispersal are died before growing to an identifiable size uncertain based on pellet counts L californi cus use of seeded grasslands has been docu- discussion mented to decrease with distance from native shrubland although in 2 of 3 sites the decrease although the lack of replication across sites was not evident within the 100loom m range we makes it difficult to generalize results beyond consider in this study westoby and wagner our site the fact that our samples represent 1973 mcadoo et al 1987 similarly numbers relatively longtermlong term accumulations of pellets of pellets and thus presumably activity did strengthens interpretations for this site our not decrease with distance from woodland in results strongly suggest that lagomorphslagomorpha do this study overall decreased lagomorph activ- not effectively disperse seeds of native species ity with distance cannot alone explain the into weed infested communities at dugway detected pattern of juniper seed deposition consequently they appear to contribute little variation in pellet size also cannot fully explain to replenishing the native seed bank in these the pattern larger pellets are more likely to degraded areas contain juniper seeds EW schupp M though not abundant the most frequently fuentes and JM comezgomez unpublished data encountered seed was that of juniper this and pellets were larger in the woodland but supports the growing realization that lago pellet size did not change from 0 to 100 morphsmorphe are involved in seed dispersal of while the number of juniper seeds dropped fleshy fruited species in general dantonioDAntomo rapidly it is possible that the lagomorph species 1990 nogales et al 1995 and of junipersjunijuniperuspers in differ in both the importance of juniper in the particular they have been recorded dispers- diet and in habitat use such that the species ing f pinpinchotipvnpvnchotipinchottichottchoti and J ashet in texas and okla- dispersing the most juniper seeds is are also homa smith 1948 J osteosperma in utah least likely to forage out into the grassland we this study schupp et al 1996 fJ occidentoccidentalisoccidentahsoccidentalistalisallsails have no data on abundances of the 3 species in oregon schupp et al 1997 andaandjand I1 phoeni nor species specific data on either juniper feed- cea in southwestern spain munoz reinoso ing or foraging location but known differences 1993 although the european rabbit orycto- between sylvilagusSylvilagus sppapp and L califcalicallcailfornicusraicusornicus in lagus cuniculus breaks many juniperjumper seeds habitat preference and predator escape strate- munoz reinoso 1993 the large number of gies zeveloff 1988 suggest sylvilagusSylvilagus sppapp damaged seeds in this study was not found in might be less likely to venture into the grass- oregon schupp et al 1997 or at 2 other dug- land if they also disperse more juniper seeds way sites only a few kinkm from the present site than L californicuscalifornicus the pattern of dispersal is schupp et al 1996 many of these broken easily explained since L califomicuscalifornicuscalifornicus is a larger 199719971 LAGOMORPH SEED DISPERSAL AND ANNUAL grasslands 257 species however it should produce the larger vide cover from predators these shrubs may pellets if so the distribution of pellet size then become foci for lagomorph mediated across transects and the generally greater occur- invasion of native species rence of seeds in larger pellets argue that it is in fact L californicuscalifornicus that is least likely to for- acknowledgments age away from the woodland and most likely to disperse juniper seeds although this is con- we acknowledge the ecology center and trary to expectations based on habitat affini- utah agricultural experiment station utah ties it could explain the pattern of dispersal A state university logan utah EWS and HJH final possible explanation is that gut passage is and the spanish ministry of science JMG for rapid relative to the rate at which lagomorphslagomorpha support of our research we thank M brooks move out into the grassland and that although and 2 anonymous reviewers for improving the they continue to move and feed they void most manuscript and J martin and R johnson of juniper seeds near the woodland no data exist the environmental protection office USU S army for evaluating this possibility dugway proving ground for access to the site we were especially surprised by the low air photo verification of the age of the annual number and diversity of other species in pel- grassland and cooperation with our research lets in southern california zedler and black on range restoration approved as utah agri- 1992 germinated 10 herbaceous species from cultural experiment station journal paper no sylvilagusSylvilagus sppapp pellets at a rate of 87 seed- 5023 lings per 1000 pellets in central spain malo and suarez 1995 germinated 2034 seeds of literature CITED 52 herbaceous species from 312 g of cun- 0 BAGLEY CFC F 1991 LCTA interim report for dugway prov- iculus pellets equivalent to 2200 of our pel- ing ground 1989 1990 USU S army dugway proving lets more similar to our results welch 1985 ground report found fairly low levels of emergence of 8 BILLINGS WD 1990 bromus tectoriumtectorumtectorurntectortecthoteotorumurn a biotic cause of great pages species from hare lepus europaeuseuropa eus and L ecosystem impoverishment in the basin 300 322 in GMG M woodwell editor the earth in timidus and rabbit 0 cuniculus pellets in transition patterns and processes of biotic impover- scotland part of the difference is likely due to ishment cambridge universityumveisity press cambridge methods other studies used repeated collec- UK tions of fresh pellets up to several months DANTONIO CMC M 1990 seed production and dispersal in the non invasive succulent carpobrotus eduhsedulis old nonnativenative invasive eduis while ours was a single collection of pel- aizoaceae in coastal strand communities of central lets spanning at least several years of age california journal of applied ecology 2769327 693 702 repeated collections of newly deposited pel- FLINDERS JT AND JAJ A CRAWFORD 1977 composition and lets may have yielded greater numbers of degradation ofjackrabbit and cottontail fecal pellets texas high plains journal of range management 30 emerging seedlings at our site additionally 217 220 our germination experiments ended after 38 d HIRONAKA M AND BWB W SINDELAR 1973 reproductive nonetheless the nearly total lack of seedling success of squirreltail in medusaheadmedusahead infested range emergence from 8425 pellets is striking lago- journal of range management 2621926 219 222211 KUNDAELI N REYNOLDS morph dispersal of herbaceous species at JNJ AND HGH G 1972 desert cotton- dug- tail use of natural and modified pinyon juniper wood- way is apparently an unusual event land journal of range management 2511625 ilg116 118 one last observation is worth noting the LONGLAND WS 1991 risk of predation and food con- peak in pellet density at 20 was associated sumption by black tailed jackrabbitsjackrabbits journal of range management 4444744 447 450 with 2 relict shrubs I1 on and the other adja- MALO JEJ E AND F SUAREZ 1995 herbivorous mammals as cent to the transect pellets were extremely seed dispersersdisperdispenserssers in a mediterranean dehesa oecologia abundant in the vicinity of these shrubs which 104246104 246 255 is compatible with the notion that jackrabbitjackrabbit MALO JEJ E B JIMENEZ AND F SUAREZ 1995 seed bank activity in open habitats is concentrated in the buildupbuild up in small disturbances in a mediterranean pasture the contribution of endozoochorous disperdispel vicinity of protective shrubs longland 1991 sal by rabbits ecographyEcography 187318 73 82 in other systems where lagomorphslagomorpha may be mcadoo JK WS LONCLANDLONGLAND GJ CLUFF AND DA more important in dispersing seeds of native KLEBNOW 1987 use of new rangeland seedings by species their use in restoration of open grass- black tailed jackrabbitsjackrabbits journal of range management 4052040 520 524 land may enhanced by planting be first scat- MONSEN SBS B 1994 the competitive influences of cheat tered shrubs throughout the grassland to pro grass bromus tectoriumtectorumtectorum on site restoration pages 258 GREAT BASIN naturalist volume 57

43 50 in SBS B monsen and SGS G kitchen editors schuppewjmschupeSCHUPP EW JM GOMEZJEGOMEZ JE JIMINEZjlmenezandmAND M FUENTES proceedings ecology and management of annual 1997 dispersal of juniperusJumperus occidentoccidentalisoccidentahsoccidentalistalisallsails western rangelands USFS general technical report INT juniper seeds by frugivorous mammals on juniper GTR 313 mountain southeastern oregon great basin natu- MUNOZ REINOSO JCJ C 1993 consumeconsumo de galbulosgdlbulosgalgaigambulosbulos de ralist 577457 74 78 sabina juniperusJumperus phoeniciaphoeniceaphoemceaphoenicea sspasp turbinateturbinataturbinata guss SMITH GWG W 1990 home range and activity patterns of 1891 y dispersion de semillas polpoipotpor el conejo orycto- black tailed jackjackrabbitsrabbits great basin naturalist 50 lagus cuniculus L en el parque nacional de Dodonanadofianadolianafianaflana 249 256 donanadolianadofianaDofianagiana acta vertebrata 204920 49 58 SMITH HNH N 1948 rabbits spread cedar menace cattle- NOGALES M A VALIDO AND FM MEDINA 1995 fru man 35397353 97 98 givorydivory of plocama bendulapendula rubiaceae by the rabbit WELCH D 1985 studies in the grazing of heather moor- oryctolagus cuniculus in xerophytic zones of tener land in northeastnorth east scotland IV seed dispersal and ife canary islands acta oecologica 1658516 585 591 plant establishment in dung journal of applied PELLANT M AND C HALL 1994 distribution of exotic ecology 2246122 461 472 glassesgrasses on intermountain rangelands status in 1992 WESTOBYWESTORY M AND FHEH WAGNER 1973 use of a crested pages 109 112 min SBS B monsen and SGS G kitchen wheatwheatgrassgrass seeding by black tailed jackrabbitsjackrabbits jour- editors proceedings ecology and management of nal of range management 2634926 349 352 annual rangelands USFS general technical report WHISENANT SGS G 1990 changing fire frequencies on INT GTR 313 idaho s snake river plains ecological and manage- PETERS EFE F AND SCS C BUNTING 1994 fire conditions pre ment implications pages 4 10 inm EDE D mcarthur and postoccurrencepostoccurrence of annual grasses on the snake EAE A romney SDS D smith and PT tueller editors river plain pages 31 36 in SBS B monsen and SCS G proceedings symposium on cheatcbeatgrasscheatgrassgrass invasion kitchen editorsediedltoistols proceedings ecology and man- shrub die off and other aspects of shrub biology and agement of annual rangelands USFS general tech- management USFS general technical report INT nical report INT GTR 313 GTR 276 PYKEDAPYKE DA 1994 ecological significance of seed banks with WILKINSON L 1990 SYSTAT the system foiforhorbor statistics special reference to alien annuals pages 197 201 in SYSTAT inc evanston IL SSBB monsen and SGS G kitchen editors proceed- YOUNG JAJ A 1994 history and use of semiarid plant com- ings ecology and management of annual range- munimunitiesties changes in vegetation pages 5 8 min SBS B lands USFS general technical report INTGTRINT GTR monsen and SGS G kitchen editors proceedings 313 ecology and management ofannualof annual rangelands USFS PYKEPYKL DAD A AND SJS J NOVAK 1994 Cheatcheatgrassgrass demogra- general technical report INT GTR 313 phy establishment attributes recruitment ecoecotypestypes ZEDLER PH AND C BLACK 1992 seed dispersal by a and genetic variability pages 12 21 in SBS B monsen generalized herbivore rabbits as dispersal vectors in and SGS G kitchen editors proceedings ecology and a semiarid california vernal pool landscape ameri- management of annual rangelands USFS general can midland naturalist 1281128 1 10 technical report INT GTR 313 ZEVELOFF SIS I1 1988 mammals of the intermountain west schupeschuppSCHUPPEWMEW M FUENTES AND JMJ M GOMEZ 1996 dis- university of utah press salt lake city 365 appp persal of utah juniper juniperusJumperus osteosperma seeds by lagomoiphslagomorphslagomorpha pages 496 497 in NEN E west editor received 3 december 1996 proceedings of the fifth international rangeland con- accepted 5 may 1997 gress volume 1 society for range management denver CO great basin naturalist 573 V 1997 appp 259 262

pseudocrossidium OBTUSULUM pottiaceae BRYOPSIDA NEW TO MONTANA WITH A KEY TO NORTH AMERICAN SPECIES IN THE GENUS

PM eckelleckelljaJA hoyahoy2 andicandjcand JC elhott3ejjiott3

ABSTRACT the moss species pseudocrossidium obtusulum lindblinda crum & anderson is reported foirolforhoi the state of montana recent systematics of the genus pseudocrossidium in north america is discussed

key words mosses bryophytes montana pseudocrossidium P cnnitumcrinitumcrinierinitum P hornschuchianumhomschuchianum P obtusulum P replica turn P revolutumrevolutum

montana has one of the richest recorded checklist anderson et al 1990 but without moss floras of the western united states with discussion zander manuscript in preparation 410 known species and varieties elliott in has determined that P revolutumrevolutwnrevolutum does not occur preparation although most collecting has in the arctic or greenland or elsewhere in occurred in the western portion of the state a north america because one of the obstacles to concentrated examination of particularly dry the convenient study of andaridarld bryophyte crypto habitats in ravalli county has yielded species floras is the scattered taxonomic treatments new to the state eckel hoy and elliott in an attempt has been made here to summarize preparation these species are members of a the genus to date in north america with a key cryptocryptofloraflora of minute bryophytes occurring on and illustrations to the 5 species various soil substrates of which one is the fol- historically species in the genus pseudo lowing species crossidium have been placed in the genera barbula hedw Desmadesmatodontodon brid and tor- pseudocrossidium obtusulum tula hedw however most are distinguishable Lindb crum & lindalindb anderson from these genera by the strongly and strik- USA montana ravalli co eastcast side ofofbitorbitbit ingly revolute leaf margins most evident on terrottterroot valley north birch creek watershed 7 transverse sections of the leaf resembling 2 miles SSE of stevensvilleStevens ville NW NE sec 12 cylinders separated by a groove zander per- t7ntan r19w 3600 elev from vertical S facing sonal communication as opposed to the usual cliff face N side of irrigation ditch just above plain or narrowly recurved leaves of other old water level hoy 306 april 6 1996 BUBUFF species of other genera the character of differ- there are 5 species of pseudocrossidium entiaentiatedted perichaetial leaves may differentiate williams presently listed in the moss flora of the genus in the broadest sense especially north america north of mexico anderson et al species in south america where the bulk of 1990 spence 1987 has given a good review the species occur but for the 4 taxa occurring of the literature and american distribution of in north america here excluding P revolarevolu 4 of these the ath5th being P obtusulum lindalindbLindb tum only P homshuchianumhornshuchianum has such a crum & anderson a species considered by species found to date in north america in only some to be a variety of P revolutumrevolutum brid in 2 botanical gardens I1 in british columbia tan schrad zand tan et al 1981 and by others et al 1981 the other in massachusetts mishler to be indistinguishable from the latter in local and miller 1983 populations in the field mcintoshmclntoshmeintosh 1986 the the genus is separated from barbula and variety was elevated to species status by crum other genera of the barbuloideae such as didy and anderson 1989 for the north american modon by the absence of a ventral stereidsteroid

clinton herbariumheihel bariumbarlum buffalo museum of science buffalo NY 14211 85822858SSS pheasant lane stevensvilleStevens ville MT 59870 3conservationsconservationSCon servation biology researchReseaichsealch 835 eighth avenue helena MT 59601

959259 260 GREAT BASIN naturalist volume 57 band in the costa neither desmatodonDesmatodon nor pseudocrossidium revolutumrevolutum is a european tortula has this feature either or it is occasion- species with oblong lanceolate to ligulate upper ally weakly present meintoshmclntoshmcintosh 1986 has also leaves more strongly and more narrowly revo- pointed out the useful character of the sharp lute margins extending nearly to the leaf base and deep costal groove or keeled leaf in pseudo margins as seen in section 112 times revolute crossidium whereas in tortula and desma with leaves strongly twisted and inrolled when todon the leaf is broadly channeled and the dry the leaf mucro is not stout and is shorter costa in section is nearly circular with an ele- than in P obtusulum whose leaves are imbri- vation sometimes distinctive on the ventral cated or only somewhat twisted when dry the surface composed of differentiated cells zander perichaetial leaves are strongly differentiated 1993 has indicated that the crescent shaped in P revolutumrevolutum and there is no record of pro dorsal stereidsteroid band as opposed to the semicir- paguia in this species whereas the perichaetial cular band of desmatodonDesmatodon and tortula is dis- leaves of the occasionally gemmiferous P ob tinctivetinctive in pseudocrossidium tusulumtusculum are weakly differentiated or undiffer- mcintoshmclntoshmeintosh 1986 has also distinguished P entiaentiatedted obtusulum as P revolutumrevolutum from didymodonDidymodon pseudocrossidium obtusulum is rare in the brachyphyllusbrachyphyllum sull in whippl zand a species arctic zander in preparation but distributed sometimes associated with P obtusulum by from greenland the northwest territories and the dense papillae as well as the lack of ven- alaska south to the yukon british columbia tral stereidsteroid band tortula muralis hedw and oregon and southern california just north of its closely related if not synonymous species the mexican border map in tan et al 1981 tortula brevipesbrevibrevipenpes fesqlesq broth and Desmadesmatodontodon in an extensive dry steppe vegetational study plithobius & also have broadly sull lesqfesq SPspi- by mcintoshmclntoshmeintosh 1986 the species was reported rally once revolute margins and densely papil- as locally common in the interior of british lose leaves but usually have a rather long columbia the montana station is an inland smooth awn tiny specimens may however be extension of its previously recorded range only mucronate and distinguished from P are mclntoshmcintosh s description of the area as the cor- obtusulum by the latter s flattened crescent dillerandilleran steppe or shrub steppe is extended shaped dorsal stereidsteroid band eastward to southern wyoming and colorado key to pseudocrossidium species in based on daubenmire 1978 the area of the north america and greenland collections presently reported also represents the steppe conditions as characterized in figs 1 5 in mclntoshmcintoshmeintosh s south central british columbia 1 leaves sholshortt- to long awned study it is probably also controlled by the 2 leaves rounded obtuse at base of long awn rain shadow effect of the western mountains ventral costal surface cells 4 concave guide no rhizoidal tubers as described and illus- mexico arizonaanzona new mexico texas utah pseudocrossidium crinicnnitumcrinitumtum P aureusaureum trated by tan et al 1981 were observed in the collections 2 leaves acute at base of short awn ventral montana costal surface convex guide cells 2 botanic As to pseudocrossidium obtusulum belong- galgaigardensdens in british columbia and massachusetts ing to floristicflonfionsticstie elements with distinctive distri- pseudocrossidium honischuchianumhomschuchianum bution patterns the reports of P revolutumrevolutum 1 leaves merely apiculate to shorshortt mucronate by mclntoshmcintoshmeintosh as a western north american P 3 leaves relatively long 101 0 151 5 mm ligulate to eurasian species must be revised as revolarevolu oblong lanceolate apex obtuse ventral costal tum has been excluded from the northnoi th ameri- surface concave guide cells 4 6 margins can flora that species would now be character- stionstronglygly spirally revolute inrolled margins of ized as a eurasian pseudocrossidium blightbright green thin walled cells with hollow species s papillae without propagulapiopagula mexico SW USA obtusulum would then belong to mclntoshmcintoshmeintosh

I1 I1 pseudocrossidium replicatumreplicatum western north american wewesternstern eurasian 3 leaves relatively short 070.70 7 121.21 2 mm ovate to element since this species occurs from the ovate deltoid apex broadly acute ventral costal arctic to southern california and 2 stations in surface convex guide cells 2 3 margins revolute what one might expect to be temperate steppe I1 time of undifferentiated cells sometimes with conditions in central germany and southern rhizoidal and costal piopagulapropagula southern cali- in fornia americanamerlean NW arctic and greenland sweden tan et al 1981 schofield 1980 dis- pseudocrossidium obtusulum cusses the disjunct flora of western north 199711997 pseudocrossidium OBTUSULUM IN MONTANA 261

0 0 0 00 00

3 ad000ooo

C 0 U

figs 1 5 1 pseudocrossidium hornschuchianum 2 P crinitumcrinitum 3 P replicatumreplicatum 4 P obtusulumobtusulmn 5 P revolrevolutumutum from european material showing the relative size of the perichaetial leaf to vegetative leaves scale banbars leaves a 050.5os mm cross sections b 100 amum 262 GREAT BASIN naturalist volume 57 america and europe describing species that also contributed valuable information we thank are widespread in america such as P obtuseobtusu WA albert for his assistance and for gaminggaining lum discussed here but which are restricted access to the collection site the owner of the to few sites throughout europe it is land dan cassin and ken mcbride forest apparent that this species is an element of this service soils specialist for doing the soil disjunct distribution the absence of these analysis and other species from most of asia strongly implies that these disjuncts have literature CITED been isolated for many thousands of years at least schofield 1980 ANDERSON LE HA CRUM AND WR BUCK 1990 list of the on fine mosses of north america north of mexico the specimens were collected very bryologist 9344893 448 499 sandy loam composed of 53 sand 40 silt CRUMCKUM H AND LEL E ANDERSON 1989 new names for and 7 clay with a bulk density of 1351.351 35 gcm3 some north american mosses bryologist 9253392 533 although volcanic ash deposits occur in the daubenmire R 1978 plant geography academic press new york region nearby that substrate was absent from mclntoshMCINTOSH TT 1986 the bryophytes of the semisemiaridarid the soil where specimens were collected the steppe of south central british columbia unpub- site is in dryland sagebrush bunchbunchgrassgrass habi- lished doctoral dissertation university of british tat with precipitation varying from 8 to 20 columbia vancouver mchesyrincbesyrmche syr plants were growing on a vertical MISHLER BDB D AND NGN G MILLER 1983 distributional face studies of massachusetts bryophytes rhodora 85 just above the waterline of a large irriga- 421 432 tion canal and consequently the soil is contin- SCHOFIELD WB 1980 phytogeography of the mosses of uously moist for nearly 5 months from may to north america north of mexico in RJR J taylor and september and extremely dry during the AEA E leviton editors the mosses of north america pacific other 7 months didymodonDidymodon vinealisneails brid division of the american association for the advancement of science san francisco CA var vinealisneails was associated with the zand SPENCE JRJ R 1987 pseudocrossidiumpseudocrossidzum aureusaureum bararbartr zan- specimens der pottiaceae musci new to utah great basin although pseudocrossidium obtusulum has naturalist 4734747 347 348 gemmae in north america it fruits and richly TAN BCB C RHR H ZANDER AND T TAYLOR 1981 pseudo crossidium P so only in greenland hornschuchianumbornshornsburns chuchianchuchianumum and revulutum var in obtusulum in the new world Lindlindbergialmdbergiabergia 7397 39 42 ZANDER RHR H 1993 genera of the pottiaceae mosses of acknowledgments harsh environments bulletin of the buffalo society of natural sciences 32132 1 378 vi we are grateful to richard zander for allow- us to information in his unpublished received 16 september 1996 ing use in accepted 24 march 1997 manuscripts the key has been considerably improved by his suggestions john spence has great basin naturalist 573 1997 appp 263 267

STICK NESTS ON A BUILDING AND transmission TOWERS USED FOR NESTING BY LARGE FALCONS IN UTAH

1 stephen T BunBunnellnelll clayton M whitelwhiteawhite1 don paulapaul2 and S dwight BunneBunnell133

ABSTRACT large falcons genus falco do not build their own nests and in north america at least usually nest on high cliffs occasionally they nest in abandoned stick nests built by another large bird on the cliff in asia and particularly south africa they sometimes nest in stick nests on electrical power transmission towers this use of electric trans- towers was ly mission recentlylecent 1980 reported for the prairie falcon falco mexicmexicansmexicanmmexicanusmexicanmanus in north america but is unknown for 1 except I anecdotically documented use of an electric power pole at the turn of century in california for the peregrine falcon falco peregrinus in north america here we report such nesting of the peregrine in north america and additional tower testingsnestingsnestings for the prairie falcon

key words electric transmission towers peregrine falcon nesting prairie falcon nesting falco peregrinus falco mexicanusmexicarmsmexiemexicanusarms

several species of raptorsrapraptorestors primarily the saker falcon falco cherruycherrugcherrug in mongolia ellis buteasbuteos buteo sp ospreys pandion and eagles et al 1995 and the ukraine del hoyo et al aquila and also the common raven corvus 1994 S sorokin and V flint personal commu- boraxcorax use electric power transmission towers nications and laggar falcon ealcofalcojuggerfalco jugger in as nesting platforms and substrates steenhof india rishadrashad maoroji unpublished manuscript et al 1993 blue 1996 most of these species and commonly by the lanner falcon falco however build their own stick nests on towers biarbiarmicusmicus in south africa tarbotontarbottonTarboton and allan frequency of use of towers varies from region 1984 in the latter study of 157 nesting lanner to region and may in part have to do with falcon pairs in the former transvaal province learning within a local population that such 22322.3 used stick nests on transmission towers structures are appropriate for nesting fre- and 131.3 used stick nests on buildings in quentquentlyly newly erected transmission lines cross stark contrast however in north america the regions where historically a species was absent use of this combination of stick nest and trans- as bleedersbreedersbreeders for there were no structures for mission tower or building is very rare in fact nest sites and thus use of such towers may there are no published reports to our knowl- then allow that species to move into an area edge of peregrine falcon falco peregrinus locally and exploit a previously unused food using stick nests on transmission towers for source white and tanner white 1988 use of nesting there is however a record early in such situations thus confers a selective advan- this century from california of a peregrine nest tage to individual pairs although falcons do on a platform on an electric power pole but not build their own nests they frequently use it was not documented adequately enough in stick nests generally on cliffs that were aban- the literature to interpret what platformplatpiatforn meant doned by the original builders RM bond in letter to hickey and anderson nesting by large falcons in abandoned stick 196918 the peregrine nest on osprey pan- nests of other birds on electric power trans- dion haliahaliaetushaliaeetusetus nests on 7 m tall navigation mission structures is not uncommon in the guidance towers on pacific coastal baja cali- eastern hemisphere As an array of examples fornia JB platt personal communication stick nests on power transmission poles are approximates the hack tower discussed below used rarely by the black falcon falco sub there are however at least 2 recently published niger in australia marchant and higgins 1993 records of the prairie falcon falco mexicanusmexicanus del hoyo et al 1994 more frequently by the using transmission towers I1 in new mexico

department of zoology brigham young university prosoprovo UT 84602 tahedh2utahbutahtdh division of wildlife resources 515 east 5300 south ogden UT 84403 butah3utahtah division of wildlife resources 1596 west northnor th temple salt lake city UT 84116

263 264 GREAT BASIN naturalist volume 57

blue 1996 and I1 in nevada roppe et al 1989 the GSL and was surrounded by a vast shallow in midlatitudemid latitude north america both species lake for approximately 3 km to the nearest land particularly the peregrine typically nest in the house was the only structure left standing inaccessible locations on cliff faces that are after several years of flooding and shear ice usually in excess of about 16 m ca 50 ft on a thus it and surrounding trees had become a cosmopolitan basis however the peregrine veritable mecca for birds paul confirmed the shows considerable flexibility in the use of dif- reports on 4 april 1988 permission was ob- ferent nesting substratasubsubstratalstrata even nesting on the tained to place a nesting box about 23 the gigroundaundound size of a hack box on the roof of the 2 story throughout the early 1970s and early clubhouse in hopes that the falcons would use 1980s the utah division of wildlife resources it on 14 april the pair was still in the area UDWR erected several structures around but it could not be determined whether they the edge of the great salt lake GSL from were using the nest box on 5 may the site which to release hack peregrine falcons in a was again visited and the female peregrine reestablishment program these structures flew from the southwest corner of the building were about 7 m tall with a platform approxi- and perched nearby protesting loudly it was at mately 2 x 2 m in size on which a nesting box this time that band information was obtained was placed young leleasedreleased from the towers for both birds female LL black anodized band wereweiwel e bred in captivity by the peregrine fund RL green band male RL dark band indicat- they had black anodized USU S fish and wild- ing that both were raised at hack boxes and the life service USFWS bands placed on the left female was from the peregrine fund after leg LL and either plastic colored bands or the agitated falcons quieted down the female colored aluminum bands with characters let- disappeared on the west side of the building ters andor numbers on the lightright leg RL there 2 wings of the clubhouse come together peregrinesPereperegrinusgrines returned to the towers as adults to form a narrow passage on investigation the and began breeding on them 4 yr after the ini-ini female flushed from a ravenravenss nest built on top tial releases young produced on the towers of an electrical circuitry box about 252.5 m above had a silver aluminum USFWS band placed ground and shaded by the eaves of the build- on the RL and in some cases after 1990 a ing at the end of the passage the nest contained black anodized coded band on the LL there a clutch of 4 eggs three young 2 males I1 was no evidence through at least 1994 that female were hatched and eventually fledged falcons from the towers either initially released one male and the female are known to have there or raised there as young occupiedrereoccupiedpreoccupied survived to dispersal the other male disap- nesting cliffs that were historically used on the peared somewhat earlier but is assumed also nearby mountains some of which are as close to have dispersed all young were leg banded as 10 km the peregrinesperegrinusperegrines however seemed to prior to fledging the nearest artificial nesting prefer and perhaps even compete for towers tower was some 32 km S at the harold crane or other similar structures wildlife management area during the early 1980s the GSL underwent an unprecedented rise in water level from transmissionTRANS mlsMIS slonSIONTOWERSION TOWER NESTINGNE STING 4203 ASLAS L to a historical high of 4212 ASL peregrinesperegrinusPERE GRINES which approximately doubled its surface area most towers and buildings otherwise around another peregrine falcon nest was located the lakeshorelakeshore were surrounded by water and on a 340 kv electric power transmission tower therefore secure from human intrusion immediately adjacent to the farmington bay waterfowl management area FBWMA on BUILDINGBUILDINGNNNESTINGE STING the eastern shore of the GSL davis county PEREGRINE FALCONS falcons were initially observed by SDB and a group of 11 other biologists from the UDWR in 1988 paul received reports of a pair of on 19 may 1994 when the male flew near the peregrinesperegrinusperegrines defending the vicinity of the his- nest in which the female was incubating this toric bear river clubhouse used by a duck and all subsequent observations were made hunting club the clubhouse had been sur- from the ground using binoculars and spotting rounded by a dike to protect it from waters of scopes the falcons were using a ravenravens s nest 199711997 TRANStransmissionmlsMIS slonSIONTOWERSION TOWER NESTING FALCONS 265 built at the intersection of the main part of the tower and the middle cross section useuseableable 77 photos are not available but see figure in roope et al 1989 and fig I1 herein for approximate nest placement we believe the ravens that built the nest may have been forced from it by hetthe peregrinesperegrinuspere grines because a pair of ravens were attempting to build a nest on the next nearest transmission tower but were continually harassed by the peregrinesperegrinusperegrines since ravens typically have young nearly ready to fledge by mid may it is unlikely that the ravens would have newly moved into the area especially to attempt building a nest so close to peregrinusperegrinesperegrines due to the location of the peregrine nest the contents could not easily be seen except from an aircraft so the initial number of eggs or 14 young was unknown A lone large downy nestling ca 14 d old was first observed in the nest on 26 june the nest appeared to be very exposed most of the day to sun wind and rain the female was seen frequently shading the A nestling with her drooped wings during the hottest part of the day in the absence of the female the nestling showed signs of heat stress we last saw the young in the nest 26 july it had fledged by 30 july we judged it to be a female based on size both adult birds had leg bands the male Q ah had a black band LQLL and a red band RL miilmill the female had a black band LQLL and an aluminum band RL based on the band configbonfigconfigu- ration the female was probably hatched at I1 of 7 nesting hack towers located around the GSLS 4 the male may have been released by the pere-er grine fund as part of the reintroduction prpro- gram several attempts to trap the adults to fig 1 nest placement of the raven s nest used by the determine band numbers were unsuccessful prairie falcon nest was in the center cross arm the adult because of the location of the tower relative to falcon can be seen standing on the edge of the nest the marshlands and appropriate locations to place placement is similar to that of the nests used by peregrine a trap falcons near the GSL in 1994 the nearest established nesting hack tower used by peregrinusperegrinespere grines was at the ambassador gun club about 11 12 kinkm away it was occu- by the presence of the early nesting barn owls pied by a pair of barn owls tyto alba that however another pair of falcons was present same year 1994 As soon as the young owls in the ambassador club area and seen occa- were discovered they were removed from the siosionallynally on the tower R walters personal nesting box and placed in another nearby owl communications nest but no peregrinesperegrinusperegrines attempted to nest in to encourage the pair to return to the same the box we initially thought the pair of fal- transmission tower area we had hoped to place cons using the stick nest on the power trans- a nesting hack box on or near the same transmission tower was the same pair that had used mission tower but were unsuccessful the fal- the ambassador hack tower in the past and cons were not observed trying to reoccupy the had simply been prevented from nesting there nest in 1995 even though a single female was 266 GREAT BASIN naturalist volume 57 there in early spring and a pair of ravens nested in the same nest in 1995 in 1996 a pair of peregrinesperegrinusperegrines occupied the ambassador club nest hack tower and produced 3 young males the adult female was handedbandedunbandedun and the male had no band on RL but we could not be certain about LL during this same time justin dolling superintendent of the FBWMA found a pair of peregrinesperegrinusperegrines in alk may nesting in a ravenravers s nest on the ist horizontal bar of the ath4th transmission tower about 080.80os 8 km S of the FBWMA transmission tower used in 1994 paul saw 2 young ca 3 wk old in the nest on 21 june 2 were there on 7 july and on 16 july the adults were seen with 1 female fledgling the adult female had a black anodized aluminum band LL and a regular aluminum USFWS band RL the adult male was handedbandedununbanded the female could have been the same female that nested on the transmis- sionslon tower in 1994 based on the band configuration the males however were different birds because the 1994 male was banded in 1997 presumably the same pair based on bands or lack of them was back at the same transmission tower as in 1996 they occupied a different location on the tower but it also was a ravenraversraveis s nest the female was still incubating at the time this went to press

transmissionTRANS MISS lonIONTOWERION TOWER NESTINGNE STING PRAIRIE FALCONS

mid may 1994 mark allman a falconer in fig 2 view of the prairie falcon on the nest edge from provo and 2 colleagues found a pair of prairie falcons nesting in a ravensraveiisravenis nest on a transmission tower on lines coming from the intermountain power plant about 16 km NW prairie falcons that traditionally use deserted of delta millard county utah figs 1 2 the stick nests on cliffs this behavior by peregrinesperegrinusperegrines falcons defended the nest and tower allman is most likely a result of the use of hack towers was unable to return to the nest to determine in the reintroduction effort the frequency and its success when we checked the area in sep- availability of ravenravens s nests on power transmis- tember we found a few boltedmolted prairie falcon sion towers and the lack of acceptable cliff feathers and some regurgitated pellets like sites along the shores of the GSL there are those cast from prairie falcons below at least however abundant cliffs some used histori- I1 other tower with raven nests farther west cally by peregrinesperegrinusperegrines for nesting 5 10 km E of along the line suggesting that perhaps a and2nd GSL of interest is the fact that all but I1 of the pair of falcons may also have used that stretch peregrine falcons nesting on the transmission of transmission towers towers or buildings were from previous artificial nesting situations the handedbandedunbandedun male at the discussion transmission tower in 1996 and 1997 may have been produced from a normal cliff nesting situa- the use of stick nests on transmission tow- tion if so the fact that it would occupy a trans- ers by peregrine falcons represents a more mission tower and a ravenraversraveis s nest on the tower significant behavioral shift for them than for attests to the species ecological amplitude in 199719971 TRANStransmissionmlsMIS slonSIONTOWERSION TOWER NENESTINGSTING FALCONS 267 nesting alternatively that male may simply world vultures to guineaguineafowlfowl lynx editionsedicionsEdicions baicebalcebaleebarce have been raised at a hack tower that contained lona spain ELLIS ELLIS produc- a stick at some more distant unknown release DHD H MHM H AND P TSENGEG 1995 tivity of saker falcons raicofalco chercherrugcherruyrug in mongolia location outside of utah where the young had pages 117 130 in proceedings of the specialist work- not been banded shop middle east falcon research group abu dhabi united arab emiratesEmiemigratesrates 14 16 november 1995 addendum peregrinesperegrinusPeregrines were recently re- HICKEY jjJ J AND DWD W ANDERSON 1969 the peregrine life ported using similar pylon situations in hol- falcon history and population literature pages 3 42 in jj hickey editor peregrine falcon popula- land 1993 and germany 1995 in newsletter tions their biology and decline university of wis- 2324 1996 world working group of birds of consincoasin press madison prey and owls BUB U leyburgmeyburgMeyburg editor berlin MARCHANT S AND PJ HIGGINS EDITORS 1993 handbook germany of australian new zealand and antarctic birds volume 2 oxford university press melbourne australia ROPPE JAJ A SMS M SIEGEL AND SES E WILDER 1989 prairie acknowledgments falcon nesting on transmission towers condor 91 711 712 we especially thank carl johansson kevin STEENHOF K MNM N KOCHERT AND JAJ A ROPPE 1993 nest- bunnell donald haney frank howe robert ing of raptorsraptores and common ravens on electrical trans- walters joseph platt and christian gonzales mission line towers journal of wildlifeofwildlife management for help in banding or for the use of their 572715797157 271 281 TARBOTON W AND D ALLAN 1984 the status and conser- observations lloyd tiffandkiffandkiff and mark fuller pro- vation ofbirdsof birds offreyofpreyof prey in the transvaal monograph of vided most helpful reviews of the manuscript the transvaal museum number 4 pretoria south africa literature CITED WHITE CMM AND M TANNER WHITE 1988 use of inter- state highway overpassesoverpasses and billboards for nesting by the common raven great 48 BLUE R 1996 documentation of raptor nests on electric basin naturalist 64 67 utility facilities through a mail survey pages 87 95 in D bird D varland and J negro editors raptorsraptores in human landscapes academic press new york received 18 november 1996 DEL HOYO J A ELLIOTTELLIOTF AND J SARCATALSARGATAL EDITORS 1994 accepted 26 february 1997 handbook of the birds of the world volume 2 new cleatcledtcreatgreat basin naturalistnatuidlist 573 C 1997 appp 268 272

EFFECTS OF myofibrogranuloma ON SERUM CALCIUM LEVELS IN WALLEYE stizostedion VITREUM

craig A Shoemakershoemakerl2shoemaker1212 and harry L holloway jr 1

ansABSABSIRACTRAur1 the effect of myofibrogranulomamyofibi ogranuloma skeletal muscle degeneration on serum calcium levels in spawning walleye stizostedionstizo&tedion vitreum was examined mean serum calcium levels for healthy male walleye 11711711.711 7 151.5isls1 5 mg100 ml serum was significantly lower P 0050 05 than calcium levels for healthy female walleye 15415 4 181 8 mg100 ml serum significant mcimclincreaseseases P 010.10oi 1 in seiummeiumserum calcium weiewere seen between healthy male and myofibrogranuloma diseased male walleye 13613613.613 6 2122.1gi91 1 mg100 ml seiummeiumserum and between healthy and myofibrogranuloma diseased female walleye 20220 2 37 mg100 ml serum elevations seen in mean serum calcium levels suggest the muscle degeneration and subsequent gigranulornaanuloma formationfoiboibol mationmatlon in later stages of myonbrogranulomamyofibrogranulorna have a significant effect on serum calcium

kenkeykeixhi words calcium walleyemalmaiwalleieleteleih myofibrogranuloma colorimetric determination stizostedion vitreum

calcium ca levels of osteichthyan extra- relatively more ca positive fibers in normal cellular fluids are regulated to a finer degree muscle of MFG positive fish and MFG positive than those of moiemore primitive fishes dacke tissue than in muscle from healthy walleye 1979 hormonesholHoi mones from the pituitary ultimo kelly et al 1987 holloway and shoemaker branchibranchialsals stanciusstanniusstannius corpuscorpusclescles and monadsgonads 1993 used xrayX ray technology to demonstrate affect osteichthyan ca metabolism dacke 1979 increased opacity presumably due to concen- since ca concentration is essential for cell trated ca in MFG diseased tissue the cause membrane permeability to water neuromus- of MFG is unknown fisheries managers and cular irritability and blood clotting growth and anglers are concerned about the disease be- enzyme reactions it is less variable than other cause the involved muscle has a yellow color ions in serum urist and van de putte 1967 with sandy texture anglers discard these fish myofibrogranulomamyofibrogramiloma MFCMFG is a unique form adult osteichthyans bony fishes exhibit of skeletal muscle degeneration recognized seasonal variations in plasma ca levels associ- only in adult walleye stizostedion vitreum ated with breeding cycles dacke 1979 sex mayes 1976 economon 1978 holloway and differences in plasma ca levels of fish were smith 1982 this myopathy is characterized first reported by hess et al 1928 booke 1964 by profound alterations of the trunk musculamascula found differences in the ca levels of spawning tuitulturee produced by extensive hypertrophy of the male and female brook trout salvelinus fonti- muscle fibers economon 1978 holloway and nalis hunn 1972 presented values of vari- smith 1982 noted 2 degenerative processes ous fishes including spawning male walleye the ist and most pronounced lesion consisted from the upper mississippi river our objective of coagulation necrosis of muscle fibers accom- was to establish ca values for spawning male panied by an inflammatory response and the female and MFG diseased walleye using a formationfoiroibol mationmatlon granulomasofgranulomasof muscle tumors the simple colorimetric procedure and2nd was noninflammatory and characterized by focal areas of acute myolysis mineralization MATERIALS AND METHODS or calcification was evident in the central portion of some fibers in the more granular stage spawning walleye were collected by gill and of degeneration economon 1978 kelly et al frame net from merrit reservoir nebraska 1987 showed an increase in muscle ca associassoniassoci- 1991 and 1992 and lake sakakawea north ated with myofibrogranulomamyofibrogramiloma 55 times normal dakota 1991 only walleye greater than 500 muscle fibers histochemical staining indicated mm total length were sampled to eliminate

I1 dqartnunofocpaitiik lililllit ot biology university ofot northnoi th dakota box 9019 university station clandcrandglandgiandgrand forks ND 58202 2presotn sfiit address USDAUSOA ARSAHS rishfishfisli discasesanddiseaseDiscasediseasessandand parasites research laboratorylaboi atoilatoiy PO box 0952 auburn AL 36831

268 199719971 myofibrogranuloma AND SERUM CALCIUM 269 killing smaller fish that would not show macro- mixed the spectrophotometer bausch and scopic signs of disease blood was sampled via lomb spectronicSpectronic 20 was set at 50 transmit- cardiac puncture with heparinized lomi10 ml sy- tance at 620 nm with the blank and the test ringe and 21 gauge needle centrifugation was and standard cuvettes were then read after carried out within 24 h to minimize sample this 0250.25 ml of titranttitrent was added to each hemolysis serum was placed in dry ice for cuvette and the spectrophotometer reset at transport and stored at 80c80 C until analyzed 50 transmittance with the blank before heavily hemolyzed samples were discarded reading the test and standard this step titra- walleye were filleted and examined with un- tion was repeated and followed colorimetri- aided eye for MFG MFCMFG diseased fish were cally until the endpoint was reached all determined by observing yellow colored sandy serum ca bound to EDTA calculations were textured muscle then carried out using the following formula as the procedure for measuring serum ca con- described by oser 1965 to determine serum centcentrationration was modified from fales 1953 by ca level oser 1965 we used the latter protocol but and reagent volumes halved due volume of titranttitrent for serum serum were X 10 mg ca100 ml serum to small sample volume A 1 stock ca standard volume of titranttitrent for standard solution was prepared by dissolving 24972.497 g calcium carbonate in 6 N hydrochloric acid and A student s t test compared mean serum evaporating on a steam bath to dryness the ca levels of healthy male and female walleye residue cac12caclg was then dissolved in distilled significance was determined at P 0050.05oos due water to make 100 ml A ca standard oi01010.1 mg to small sample size and differences between ml was prepared by diluting I1 ml of the stock variances healthy male walleye and MFGMFC posiposl ca to 100 ml with 141.4 N sodium chloride A tive male walleye as well as healthy female light sensitive stock solution of murexide was walleye and MFGMFC positive female walleye were prepared by dissolving 0250.25 g ammonium pur- compared by unequal variance t test sokal purate in 5 ml distilled water and 25 ml 95 and rohlf 1981 the critical significance level ethanol the working murexide solution was was P 0100.10 for tests between MFCMFG positive prepared by adding stock murexide solution to and healthy male and female walleye because 100 ml 0050.05oos N sodium chloride until an OD of of the low numbers of MFCMFG positive fish 6 0390.39 0480.48 was reached at 620 nm the titranttitrent males and 4 females was prepared by mixing 18 volumes of the working murexide solution 1 volume of 0140.14 RESULTS AND discussion N sodium chloride and I1 volume of stock di- sodium dihydrogen ethylene diaminetetraacetic selected serum ca titration curves standard acid EDTA solution 000500050.005 M EDTA titranttitrent spawning female nb201 spawning male was mixed prior to each set of determinations 006600661 MFCMFG positive spawning female 0043 kept in brown bottles and used within 151.5isls h and MFCMFG positive spawning male 025202521 are the principle of the procedure as described shown in figure 1 and serum ca eoncentraconcentra- by oser 1965 is that the ca in serum mixed tions are described in table I1 for healthy and with murexide forms a red colored complex in MFCMFG positive walleye A significant differ- equilibrium with free ca ions titration of the ence was found between serum ca levels of ca murexide complex with EDTA chelateschchelakeselates male and female walleye FP 0050.05 serum ca frees ca ions causing the release of ca ions levels of male walleye differed significantly from the complex As the murexide ion calcium from mean serum ca levels of MFCMFG positive murexide ratio increases there is a shift in male walleye P 0100.10olo A significant differ- color from red to purple endpoint the pro- ence was found between serum ca levels of cedure followed is described briefly 0250.25 ml healthy female walleye and MFCMFG positive of 0140.14 N sodium chloride was placed in a female walleye FP 0100.10 blank cuvette and equal amounts 0250250.25 ml of serum ca values were higher in this study serum and working ca standard were placed than in others table 2 one explanation may in test and standard cuvettes respectively be species differences hunn 1972 examined then 2252.25 ml of working murexide and 050.5 ml spawning male walleye from the upper missis- of titranttitrent were added to each cuvette and sippi river and found serum ca lower 9529529.52 270 GREAT BASIN naturalist volume 57

al 1943 shell 1961 booke 1964 and atomic 0 standard absorption spectrophotometry grant et al 1970 81 nb201 hunn 1972 the colorimetric method described by faleshaleshaiespales 1953 determined total serum ca ionic 00066 777 plus protein bound and he found excellent 150043130043 the method of and 73 agreement with clark col x0252 lip 1925 which also measured total ca in 69 addition the method is inexpensive low cost of equipment spectronicSpectronic 20 and reagents 65 the difference between serum ca levels of healthy male and female walleye was due to 61 egg production sex differences in ca levels in a osteichthyan fish were first reported by hess 57 et al 1928 who found a range of 9 12512.5 mg 100 ml serum in male cod cadusgadus morheamorhuamorhua 53 while mature females had a range of 12712.7 29 49- mg100 ml serum dacke 1979 suggested that hypercalcemia is related to the influence of

45 1 estrogen and ovarian follicle maturation which 00 05 10 15 20 25 30 35 40 45 50 stimulate synthesis of yolk protein and hence ml ttrantfitrantstrant an increase in protein bound plasma ca urist van de putte 1967 found similar results fig 1 selected ca titration curves of serum collected and fromfi om spawning walleye stizostedion vitreum using a col for spawning sturgeon spawning brook trout oiimetnctrimetricorimetricoriorlmetric assay standard 01 mg caalcamlcami nb201 females also exhibited increased levels of ca healthy female 0066 healthy male 0043 MFG posi booke 1964 hunn et al 1992 found ca lev- tive female 0252 MFCMFG positive malemaie els were significantly higher in gravid female than in male golden trout oncorhynchus aguaguabonitaabonita mg100 ml than our mean value 116811.6811 68 mg dacke 1979 stated that ca levels through- 100 ml for spawning male walleye this may out the osteichthyan subphylum are regulated be a result of the blood sampling procedure within narrow limits this is accomplished in caudal peduncle punctulepuncturepunctuieule vs cardiac punc- part by exchange with ca in the aquatic envi- ture which is known to influence other blood ronment and by exchange with bone ca oser parameters ege g serum enzyme levels hille 1965 stated slight variations in normal levels 1982 or simply differences between river and of serum ca are indicative of pathology such as reservoir fish low serum ca levels in paddle- bone abnormalities and muscle tumors even fish polyodon spaspathulathula suggest physiological though our sample sizes of MFCMFG positive fish hypocalcemia and in this respect paddlefish were small elevations were seen in ca levels of show phylogenetic affinity to stursturgeonsgeons grant MFCMFG positive male and female walleye these et al 1970 the low levels seem to support elevations suggest ca is highly regulated in urist and van de putte s 1967 suggestion that walleye and that MFCMFG has a significant effect hypocalcemia is related to the absence of a bony on serum ca levels probably resulting from skeleton however gravid female white sturgeon acute myolysis holloway and smith 1982 acipenser transtransmontanusmontanus exhibit increased leading to increased extracellular ca serum ca low serum ca determined for channel catfish ictalurus punctpunctatusatus 929.29 2 mg100 acknowledgments ml may be a result of obtaining fish from cul- ture ponds rather than from natural waters we acknowledge federal aid sport fish warner and williams 1977 restoration funds from nebraska game and various techniques were used to determine parks commission project F 77 11 north serum ca levels which may have resulted in dakota game and fish department project differences noted methods of ca determina- F 2 11 and south dakota game fish and parks tion include automated methods warner and department project F 14 11 HLH principal williams 1977 colorimetriccolonmetriemetric methods field et investigator 199711997 myofibrogranuloma AND SERUM CALCIUM 271

TABLE 1 serum ca mg100 ml serum of healthy and MFG positive spawning male and female walleye stizostedion vitreum from merrit reservoir NE and lake sakakawea ND

fish sample size mean sl range healthy female 20 15418154 18at 119 195 healthy male 18 11711.711711515bt 91 14614.614 6 MFG positive female 4 20220 2 37f3 7 1711717.11 25425 4 MFG positive male 6 13613 6 2vav2 it 115 168 means nithvithwith different alphabetical lettelletterietter superscripts weiewelewere significantly different between sexes at P 005 the means marledmariedmarked with diffendifferentnt symbols were significantly diffrientdiffeientdifferent within the same sex iei e healthy female compared to MFG positive female and healthy male compared to MFG positive male at P 0100 10

TABLE 2 published serum ca values mg100 ml serum for freshwater fishes

sample size authors species mean s range warner and williams 1977 107 929 2 2659652ggs65 39 45 ictalurus punctpunctatusatus field etetalal 1943 5 10 115 9459.459 45 147714.7714 77 cyprinus carpio shell 1961 30 pooled sample 10210.210 2 19519.519 5 micropterus dolodolomiettidolormewdolormiettimemmew grant etalet al 1970 3 female 746 8 male 772 polyodon spaspathulathula grant etalet al 1969 14 males 126 carassiusCaras sius auratusauranus urist and van de putte 1967 6 gravidgi avidavld females 184 4 malenongravid female 72 acipenser transmontanustransmontanus hunn 1972 9 male 9529.529 52 030.30 3 stizostedionstzzostedton vitreum booke1964booke 1964 females 5545.545 54 27792727.7979 males 6346.346 34 106510.6510 65 salvelinusfontinalissalvelinus fontinalis

literature CITED GRANT beemBFPMBY PM mehrleandtrMEHRLE AND TR RUSSEL 1970 serum characteristics of spawning paddlefish polyodon BOOKE HEH E 1964 blood serum protein and calcium levels spaspathulathula comparative biochemistry and physiology in yearling brook trout progressive fish culturist 3732137 321 330 2610726 107 111 GRANT BYB F PKTPK T PANG AND RWR W GRIFFITH 1969 the CLARK EYE P AND JBJ B gollieCOLLIECOLLIP 1925 A study of the tisdall twenty four hour seminal hydration response in gold- method for the determination ofbloodofoxbloodblood serum calcium fish carassiusCaras sius auauratusauranusratus I1 sodium potassium cal- with suggested modification journal of biological cium magnesium chloride and osmolality of serum chemistry 6346163 461 and seminal fluid comparative biochemistry and DACKE CGC G 1979 calcium regulation in sub mammalian physiology 3027330 273 280 vertebrates academic press inc london 222 appp HESS AEA F CEC E BILLS M WEINSTOCK AND H RIVKIN ECONOMON PP 1978 A muscular dystrophy like anomaly 1928 differences in calcium level of blood between of walleye american fisheries society special pub- the male and female cod proceedings of the society licationlication 1122611 226 234 for experimental biology and medicine 253499534925 349 359 FALES FW 1953 A micromethod for the determination of HILLE S 1982 A literature review of the blood chemistry serum calcium journal of biological chemistry 204 of rainbow trout salmo gairdnergairdnengairdgairdnerinerinerlnen rich journal of 577 585 fish biology 205359053520 535 569 FIELD JBJ B CAC A ELVEHJEM AND C JUDAY 1943 A study HOLLOWAY HLH L JR AND CAC A SHOEMAKER 1993 roent- of the blood constituents of carp and trout journal genologygenology of myofibrogranulomamyofibrogranulorna in walleye stizoste of biological chemistry 148261148 261 269 dion vitreum canadian journal of zoology 7121971 219 222211 272 GREAT BASIN naturalist volume 57

HOLLOWAY HLH L JR AND CEC E SMITH 1982 A myopathy SHELL EWE W 1961 chemical composition of blood of small- in north dakota walleye stizostedionstzzostedion vitreum mouth bass united states fish and wildlife service mitchill journal of fish diseases 55275 527 530 research report 57 USU S government printing office HUNN JBJ B 1972 blood chemistry values for some fishes washington DC 36 appp of the upper mississippi river journal of the min- SOKAL RRR R AND ejFJ ROHLEROHLF 1981 biometry WH free- nesota academy of science 381938 19 212 1 man and company new york 859 appp HUNN JBJ B RJR J WIEDMEYER IEI1 E GREER AND AWA W GRADY URIST MRM R AND KAK A VAN DE PUTTE 1967 comparative 1992 blood chemistry of laboratory reared golden biochemistry of the blood of fishes identification of trout journal ofaquaticof aquatic animal health 42184 218 222 fishes by chemical composition of serum pages KELLYKLLLY RKR K JEJ F klavierkamp RXR V HUNT AND 0 NIELSEN 207 217 in RFR F mathewson and DED E railrallrali editors 1987 chemical analysis of muscle from walleye sti sharks skates and rays the johns hopkins press zostzostedionedion vitreum vitreum with myofibrogranuloma baltimore a chronicchrome myopathy canadian journal of fisheries WARNER MCM C AND RWR W WILLIAMS 1977 comparison be- and aquatic sciences 44142544 1425 1431 tween serum values of pond and intensive raceway MAYES MAM A 1976 muscular necrosis of the walleye stisto cultured channel catfish Ictalums punctpunctatusatus rafinesrabines zostzostedionedion vitreum in nebraska transactions of the que journal of fish biology 1138511 385 395 american microscopical society 9526995 269 270 OSER BLB L 1965 hawk s physiological chemistry the received I1 april 1996 blakiston division mcgraw hill book company accepted 17 march 1997 new york 1472 appp great basin naturalist 573 0 1997 appp 273 277

HELMINTHS FROM THE SONORAN SPOTTED WHIPTAIL cnemidophorus SONORAE AND THE WESTERN WHIPTAIL cnemidophorus TIGRIS SAURIA TEIIDAE FROM SOUTHERN ARIZONA WITH COMMENTS ON abbreviataabbreviate terrapenisTERRAPENIS NEMATODA physalopteridae

stephen R coldGoldGoldbergberglbergi1 charles R bursey2 and hay chearlcheamlcheam1

key words cnemidophorus sonoraexonoraesonorae cnemidophorus tigris teiidae helminthshelminthes nematoda cestoda acanthocephala arizona

cnemidophorus sonoraexonorae lowe and wright LACM and the university of arizona UAZ 1964 the sonoran spotted whiptail occurs and examined collection data are given in the from southeastern arizona to northeastern appendix the lizards were originally pre- sonora and east to western new mexico cne- served in 10 formalin or boulisbouins fixative and midophorus tigris baird and girard 1852 the stored in 70 ethanol the body cavity was western whiptail ranges from oregon and opened and the gastrointestinal tract was idaho south through california to baja califor- excised by cutting across the esophagus and nia and coahuilaCoahuila mexico and eastward to rectum the esophagus stomach small intes- western colorado new mexico and texas tines and large intestines were slit longitudi- stebbins 1985 helminthshelminthes have been previ- nally and examined separately under a dissect- ously reported from cnemidophorus sonoraexonorae ing microscope the body cavity and liver were by mcallister 1992 and cnemidophorus tigris also examined each helminth was removed by Grundgrundmannmaimmarm 1959 babero and matthias and initially placed in a drop of glycerol on a 1967 telford 1970 specian and ubelaker glass slide nematodes were identified from 1974a 1974b benes 1985 and lyon 1986 these temporary mounts cestodescustodesCestodes were stained abbreviateabbreviata terraterrapenispenis hill 1945 morgan with hematoxylinhaematoxylin mounted in balsam and iden- 1945 was originally described from specimens tified Acanthocephalaacanthocephalansns were cleared in xylene taken from 7 ornate box turtles terrapene mounted in balsam and assigned to genus omata collected from widely separated points terminology usage is in accordance with mar- in oklahoma hill 1945 the ist lizard host to golis et al 1982 be reported for this helminth was sceloporus cnemidophorus sonoraexonorae was found to har- jarrojarroviivii also collected from widely separated bor 2 species of cescestodescustodestodes oochoristica bivitel points in arizona new mexico and mexico lobata loewen 1940 and 0 macallistemacallisteriri bursey goldberg et al 1995 1996 the purpose of and goldberg 1996 and 3 species of nema- this paper is to report on a helminthological todes abbreviate terraterrapenispenis pharyngodon examination of cnemidophorus sonoraexonorae and carneriwarneriwomen harwood 1932 and thubunaea cne- cnemidophorus tigris from southern arizona midophorus babero and matthias 1967 cnemi- and the presence of A terrapenisterrapenis in these 2 dophorusdophorus tigris was found to harbor 1 species additional lizard hosts of cestode 0 bivitellobata 2 species ofofnemaounemanema twenty one female cnemidophorus sonoraexonorae todes A terraterrapenispenis and PF carneriwarneriwarneri and aystacysta mean snout vent length SVL 73273.2 mm canthscantes of a species of acanthocephala centro 565.6 s range 60 80 and 82 cnemidophorus rhonchusrhynchusrhynchus sp valencesPreprevalences and mean intensities tigris 28 females 54 males mean SVL 65665.6gsg for these helminthshelminthes are given in table 1 the in- mm 10loi10110.11 s range 34 82 mm were borrowed fection prevalence between males and females from the herpetology collections of the nat- of C tigris was not significantly different for ural history museum of los angeles county A terrapenisterrapenis x2xa 0170.17 1 df P 005oos0.05 for P

department of biology whittierwhittlerWhittieiiieriler college whittlerwhittierWhittiettiei CA 90608 213epartmentepaitment of biology pennsylvania state university shenangoShenango campus 147 shenangoshenadgoShenadeonango avenue shalonsharon PA 16146

273 274 GREAT BASIN naturalist volume 57

TABLETABLL 1 gastrointestinal helminthshelminthes of 21 cnemidophorus xonoraesonorae and 82 C tigris from pima county AZ

cnemidophorusdophorusdop horus sonoraexonorae cm midophoriischemidophoruscnemidophorus tigris heiihelihelminthHeIrninth prevalencepi evalence mean intensity prevalence mean intensity range sitelaitel range sitelaitelsite1 oochoristicaoochonstica bivitellobata 5 60 c 1 30 c oochonsticaoochoristica macallistemacallistermacallistenmacaimacalmacallisterilistenri 52 20 c abbreviateahbreviataabbreviata terrapenisterrapemsferraterrapenis 762 8018 01 24 ab 862 143114.3114 3 1 61 abcdab1c1dabad pharyngodon wamenwomen 142 227422 7 4 55 cd 42 50350.350 3 1 220 d thubunaea cnemidophorus 52 20 b centrorhynchus sp 4 10 b lad esophagus 1 stomach e small intestine d laigelarge intestine 2nanliewllewIILW listhost icloldd

carneriwarneriwatwarnenwarnetinerlneri x2xa5c2 0210.210 21 1 df P 005oos0.050 05 cnemianemi lizard sceloporus undulatesundulatus morgan 1941 dophorusdophorus sonoraexonorae is a new host record for 0 mcallister and trauth 1985 adults albrearbreofabbreof macallistemacallistermacallistenmacallisterimacaimacallistenri A terrapenisterrapemsterrapenis P warwarnenmarwaynenmaynennerlneri and T viataaviata terraterrapenispenis have previously been reported cnemidophorus C tigris is a new host record from sceloporusjarroviisceloporus jattojarrojarroviiuliullvii in arizona new mex- for A terrapenisterrapemsterfatetraterrapenis helminthshelminthes were placed inm vials ico and mexico goldberg et al 1995 1996 of alcohol and deposited in the USU S national this is also the ist report of 0 macallistemacallisteriri parasite collection USNPC beltsville mary- from a teiidtegid lizard although unidentified species land accession numbers in appendix of oochoristica have been reported from cne- oochonsticaoochoristica bivitellobata pharyngodon midophorus dixonidiboni C gularis and C tesselatustesselatus dameriwameriwarnen and thubunaea cnemidophorus have mcallister 1990a 1990d mcallister cordes been previously reported in north american and walker 1991 in arizona benes 1985 re- lizards table 2 and may be limited to tend ported oochoristica from cnemidophorus tigris lizards the occurrence of T cnemidophorus leonyxcoleonyxCo variegatusvariegatus phrynosoma solare scelo- in the crotcrotalidcrotaloidalid snakes Grocrocrotalustalus cerastes C porus magister and uta stansburianastansburiana but did mitclemitchemitchelhmitchelemitchellilh and C scutulascutulatustus by babero and not identify the species emmerson 1974 needs further study to abbreviateabbreviata terraterrapenispenis is a heteroxenous determine if the snakes are indeed hosts or if physalopterid helminth with an indirect life the parasites were present in lizards that the cycle involving an insect intermediate host snakes had ingested mcallister 1992 ques- anderson 1992 echternacht 1967 reported tioned the determination of 0 bivitellobata in that termites are the major dietary component sceloporus undulatusundulates and suggested that it for C sonoraexonorae and C tigris from the santa rita probably is oochonstica0ochoristica sceloponscelopori thus S un mountains pima county arizona represent- dulatus is not included in table 2 species of ing over 90 of all prey organisms consumed centrorhynchus typically use arthropods prob- mitchell 1979 reported a predominance of ably insects as intermediate hosts and primar- termites in the diets of C sonoraexonorae and C tigris ily birds of prey as definitive hosts petro- in cochise county southeastern arizona vitt chenko 1958 the occasional presence of a and ohmart 1977 similarly found that ter- cystcystacanthacanth in the stomach of an insectivore mites compose 76 of the diet of C tigris could be expected one nematode not found living along the colorado river in western in these lizards but frequently associated with arizona pianka 1970 reported that while tend lizards is parathelandros tetanustexanus specian southern C tigris populations eat large quanti- and ubelaker 1974 this helminth may be ties of termites northern populations idaho limited to west texas see baker 1987 nevada utah utilize other food types and this is the first report of adult abbreviateabbreviata consume few termites if termites serve as inter- terrapenisterrapemstettatetraterrapenis from tend lizards although larvae mediate hosts for abbreviateabbreviata terraterrapenispenis low of abbreviataabbreviateofabbreviata sp have been reported from C frequencies of these insects in the diets of C sexhneatussex1ineatus by mcallister trauth and conn tigris from northern populations might account 1991 larvae of abbreviateabbreviata sp have also for the absence ofaof A terrapenisterrapenis in the studies been reported from the crotaphytid lizard of these populations by grundmann 1959 crotaphytus colcoicollanscollariscollaraslaris and the phrynosomaphrynosomatidtid babero and matthias 1967 and lyon 1986 199719971 NOTES 275

TABLE 2 reports ofoochofisticaofoochonstwa bivitellobata pharyngodon wamenwomen and thubunaea cnemidophorus from tend lizards helminth host locality prevalence reference oochonsticaoochoristica bivitellobata cnemidophorus burti arizona 1572157 2 goldberg and bursey 1989 C dixonidixomdiboni texas 95816958 16 mcallister cordes and walker 1991 C exsanguisexsangws new mexico texas 7878787 8 mcallister 1990c C flagellicaudusflagelhcaudus new mexico 52322523 22 mcallister 1992 C gulansgularis new mexico texas 328913289 1 mcallister 1990d texas 1831183 1 mcallister et al 1995 C hyperythrus california 510455104 5 bostic 1965 canornatusCAnC inornatus arizona 1078131078 13 goldberg and bursey 1990 C neomexicanus new mexico texas 76111761 11 mcallister 1990b C sexhneatussex1ineatus kansas 91144 63 loewen 1940 nebraska 3310033 loo100 brooks and mayes 1976 2643264 3 shoop and janovy 1978 south dakota 1323571323 57 dyer 1971 C sonoraexonorae arizona 1166116ilg 6 mcallister 1992 C tesstesselatuselatus texas 32711 mcallister 1990a C tigris california 1349271349 27 telford 1970 idaho 1332411332 41 lyon 1986 nevada 5975597 5 babero and matthias 1967 utah 577157 71 grundmann 1959 C uniparensumuniumparensparens arizona 83126831 26 goldberg and bursey 1990 C velox colorado not stated douglas 1966 new mexico 2375237 5 mcallister 1992 pharyngodon mamenwomenmomen C exsanguisexsangws new mexico texas 108711 mcallister 1990c C gularisgulans oklahoma texas mexico 69289 24 mcallister 1990d texas 2832283 2 mcallister et al 1995 C inornatus arizona 1878231878 23 goldberg and bursey 1990 texas not stated specian and ubelaker 1974a C laredoensis texas 52223522 23 mcallister et al 1986 C neomexicanus new mexico texas 2613261 3 mcallister 1990b C sexlineatus texas 245024 50 harwood 1932 south dakota 1923831923 83 dyer 1971 arkansas 1551291551 29 mcallister trauth and conn 1991 C tesselatustesselatus texas 42715427 15 mcallister 1990a C tigris utah 577157 71 grundmann 1959 arizona nevada 63100 63 babero and matthias 1967 thubunaea cnemidophorus C tigris nevada 98710987 10 babero and matthias 1967 C burti arizona 2574257 4 goldberg and bursey 1989 C sexlineatussex1ineatus arkansas 3516351331 6 mcallister trauth and conn 1991 it has been shown that skrjabinoptera phryno- inelneme cnemidophorus sonorae and jeffrey feng soma also a member of the physalopteridae whittier college for technical assistance and a common parasite of phrynosoma sppapp is dependent upon ants pogonomyrmex sp as literature CITED intermediate hosts lee 1957 the possibility that termites may serve as intermediate hosts ANDERSON RCR C 1992 nematode parasites of vertebratesofvertebrates ofaof A terrapenisterrapenis needs to be investigated such their development and transmission CAB international wallingford oxonaxon UKU K 578 appp information would helpful in be determinindeterdeterminingminin9 BABERO BBB B AND FHEH EMMERSON 1974 thubtmaeathubunaea cne- distribution patterns ofaofA terrapenisterrapenis midophorus in nevada rattlesnakes journal of parasitology 6059560 595 acknowledgments BABEROBABEBO BBB B AND D MATTHIASMATFHIAS 1967 thubunaea cnemi-cnemianemi dophorusdop horus n sp and other helminthshelminthes from lizards authors thank charles H cnemidophorus tigris in nevada and arizonaai izonatzona trans- the lowe depart- actions of the american microscopical society 86 ment of ecology and evolutionary biology 173 177 university of arizona for permission to exam BAKER M R 1987 synopsis of the nematoda parasitic in 276 GREAT BASIN naturalist volume 57

amphibians and reptiles memorial university of new- ual whiptail lizards tehoehteiidaeTen dae in north america II11 foundfoundlandland occasional papers in biology 111iiiili11 1 325 the new mexico whiptail cnemidophorus neomexi BENEsBLNLS ESE S 1985 helminth parasitism in some centialcentral banuscanus journal of wildlifeofwildlife diseases 2640326 403 406 arizona lizards southwestern naturalist 3046730 467 473 1990c helminth parasites of unisexual and bisex- bostic DL 1965 parasites of the tend lizard cnemi- ual whiptail lizards teiidae in north america III111 dophorusdophorus hyperythrus beldingibeldingi southwestern natu the chihuahuan spotted whiptail cnemidophorus ralistcalistiahstl031310313 exsanguisexsangws journal of wildlife diseases 265449654426 544 546 BROOKS DRD R AND MAM A MAYESMAYLS 1976 morphological vari- 1990d helminth parasites of unisexual and bisex- ation in natural infections of oochonsticaoochoristica bivitellobata ual whiptail lizards oenteiidaeTen dae in north america IV loewen 1940 cestoidea anoplocephalidae trans- the texas spotted whiptail cnemidophorus gularisgulans actions of the nebraska academy of science 3203 20 21 texas journal of science 4238142 381 388 DOUGLAS CLC L 1966 amphibians and reptilesleptiles of mesa 1992 helminth paiapalaparasitessitessltes of unisexual and bisex- veideveldeverde national paliparipaikparkpalk colorado university of kansas ual whiptail lizards oehoenteiidaeTen dae in north america villviliVIII publications museum of natural history 1571115 711 744 the gila spotted whiptail cnemidophorusflagellicnemtdophoruscnemidophorus flagelh DYERDYLR WG 1971 some helminthshelminthes of the six lined lizard cacaiduscaudusudus sonoran spotted whiptail cnemidophorus cnemidophorus sexlineatussex1ineatus in south dakota pro- sonorae and plateau striped whiptail cnemidopho- ceedingsce of the unthologicaluntheologicalhelminthologicalhelnhein society of wash- rus velox texas journal of science 4423344 233 239 ington 382563885638 256 mcallister CT JE CORDES AND JM WALKER 1991 ecnrlrnaclneciiternachr AC 1967 ecological relationships of two helminth parasites of unisexual and bisexual whip- species of the lizard genus cnemidophorus in the tail lizards teiidae in north america VI the gray santa rita mountains ofofanzonaarizonaAnzona americanamerlean midland checkered whiptail cnemidophorus dibonidixoni texas naturalist 7844878 448 459 journal of science 4330943 309 314 GOLDBERGGOLDBLRC SRS R AND CRC R BURSEY 1989 helminthshelminthes of the 1995 helminth parasites of unisexual and bisex- giant spotted whiptail cnemidophorus burti stictosticco ual whiptail lizards teiidae in north america IX grammasgrammus sauriasauna teiidae pi oceoceedingsproceedingsedings of the hel the plateau spotted whiptail cnemidophorus gulans minthologicalminthological society of washington 568656 86 87 septemvittatus texas journal of science 478347 83 88 1990 helminthshchninthshelminthes oftheodtheof the arizonaanzona little striped whip- mcallister CTC T AND SES E TRAUTH 1985 endoparasitesEndo parasites tail cnemidophorus inomatusinomatus arizoarlzoarizonaearizonananzoanzonaenae and the desert of crotaphytus collariscolcollaraslaris collanscollariscollarascolcoilarislafis sauriasauna iguanidae giasslandgrassland whiptail cnemidophorus miunimiparensuniparensharengvarensparens sauriasauna from arkansas southwestern naturalist 3036330 363 370 teiidaeTenoendae flomfrom southeastsoutheasterneineln arizona journal of the mcallistermcallisier CT SES E TRAUTH AND DB CONN 1991 helminthological society of washington 578357 83 86 helminth parasites of unisexual and bisexual whip- goigolGOLDBERGDBLRG SRS R CRC R burseybubseyBURSLY AND RLR L BEZY 1995 tail lizards teiidae in north america VII the six helminthsheirninthshelminthes of isolatedofisolated montane populations ofyarrowof narrowyarrow s lined racerunnerracerunner cnemidophorus sexlmeatussex1ineatus texas spiny lizardd sceloporus farrojarrojarrovnjarroviiullvii phrynosomatidae journal of science 4339143 391 397 southwesternSouthwestein naturalist 4033040 330 333 mcallister CT SE trauthtbauihandjeAND JE UBELAKERUBELAKLR 1986 1996 gastrointestinal helminthshelminthes ofofyarrowmarrowyarrow s spiny nematode parasites of the parthenogenetic whiptail lizard sceloporusjarroviisceloporus jarroviijarrouitditvii phrynosomatidae in mex- lizard cnemidophorus laredoensis sauriasauna teiidae ico american midland naturalist 135299135 299999 309 from south texas proceedings of the helminthologi- GRUNDMANN AW 1959 parasitesPaiapalasitessltes recovered from six cal society of washingtonofwashington 5313853 138 139 species of utah lizards journal of parasitology 4539445 394 MITCHELL JCJ C 1979 ecology of southeastern arizona HARWOODHAKWOOO PD 1932 the helminthshelminthes parasitic in the am- whiptail lizards cnemidophorus teiidae popula- phibia and reptilia of houston texas and vicinity tion densities resource partitioning and niche over- pi oceoceedingsproceedingsedings of the USU S national museum 81181 1 71 lap canadian journal of zoology 57148757 1487 1499 hlllHILLiliIII111 L WC 1945 physaloptera terrapenisterrapenis a new nematode MORGAN BBB B 1941 additional notes on north american fromflom a tortoisetoitol toisetolse transactions of the americanamerlean micro- physaloptennaephysalopterinae nematoda proceedings of the scopicalscopical society 605960 59 64 helminthological society ofwashingtonofwasbidgton 8638 63 64 lelLEELLL SHS H 1957 the life cycle of skrjabinoptera phryno- petrochenko VI 1958 acanthocephala of domestic and soma oitlepportlepp schulz 1927 nematoda spiruroideaspiniroidea wild animals volume II11 israel program for scientific a gastric nematode of texas horned toads phryno- translations jerusalem 1971 477 appp soma corncornutumutum journal of parasitology 436643 66 75 PIANKA ERE R 1970 comparative autecology of the lizard loewiloewenLOLWI N SLS L 1940 on some reptilian cescestodescustodestodes of the cnemidophorus tigris in different parts of its geo- genus oochoristicaoochonstica anoplocephalidae transactions graphic range ecology 5170351 703 720 oftheodtheof the americanamerlean microscopical society 5951159 siisli511 518 SHOOP WL AND J JANOVY JR 1978 adult cestodescustodescestodes from LYON RER E 1986 helminth parasitespaiaparasitessltes of six lizard species the coelomic cavity of the teidtcidacid stcsic lizard cnemi- flomfrom southernsouthernthein idaho proceedings of the helminthol- dophorusdop horus sexlmeatussex1ineatus journal of parasitology 64 ogical society of washington 5329153 291 293 561 562 maltmaitMAtiMAKCOIISbuswusmus L GW escriescilESCII JC HOLMES AMA M KURIS AND SPECIAN RDR D AND JEJ E UBELAKER 1974a two new species GA schanSCHADsc hanHAD 1982 the use of ecological terms in of pharyngodon diesing 1861 nematoda oxyuri- paiapalaparasitologysitology lepoltreport of an ad hoehocbocboe committee of the dae from lizards in west texas proceedings of the americanamerlean society ofparasitologistsofpaiasitologists journal ofofparaokparapara helminthological society of washington 41 46 51 sitology 6813168 131 133 1974b parathelandros tetanustexanus n sp nematoda mcallister CT 1990a helminth paiapalapaiasitesparasitessitessltes of unisexual oxyuridae from lizards in west texas transactions and bisexual whiptail lizards oenteiidaeTendae in north amer- of the american microscopical society 9341393 413 415 ica I1 the Coloiacoloradodo checkered whiptail cnemido- SIEBBINSSTEBBINS RCR C 1985 A field guide to western reptiles and phorus tesstesselatuselatus journal of wildlife diseases 26 amphibians houghton mifflin company boston 139 142 336 appp 1990b helminth parasitespaiapalasitessltes of unisexual and bilexbisex TELFORD SRS R JR 1970 A comparative study of endopar 199711997 NOTES 277

asitism among some southern california lizard pop- 20666 67 20677 20681 20687 29637 30087 30090 ulationsulations american midland naturalist 8351683 516 554 30682 VITT LJ AND RD OHMART 1977 ecology and repro- duction of lower colorado river lizards II11 cnemi- natural history museum of los angeles county dophorusdophorus tigris oehoenteiidaeTendae with comparisons herpe cnemidophorus tigris N 77 27 females 50 males tologicatologica 3322333 223 234 foothills santa catalina mountains 822 m elevation 3220n 11007w pima county arizona collected 1962 received I1 october 1996 LACM 143588 1963 LACM 143587 143589 1964 accepted 5 december 1996 LACM 143590 93 1966 LACM 143586 143594 634 1969 LACM 143558 85 N 5 1 female 4 malesmaies avra valley 457 m elevation 3220n 11120w pima county collected 1964 LACM 14365 69

APPENDIX US national parasite collection MUSEUM ACCESSION NUMBERS cnemidophorus sonoraexonorae oochonsticaoochoristica bivitellobata 86861 oochonsticaoochoristica macallistemacallistermacallisterimacallistenmacaimacallistenri 86862 abbreviateabbreviata ter university of arizona rapenis 86863 pharyngodonpharifngodon wamenwomenwarnerinerl 86864 thubunaeathubiinaea cnemidophorus sonoraexonorae N 21 all females sabino cnemidophorus 86865 canyon 883 m elevation 3220n32 20n I11049w11049w santa cata- cnemidophorus tigris oochoristicaoochonstica bivitellobata 86866 lina mountains pima county arizona collected 1953 abbreviateabbreviata terrapenisterrapenis 86867 pharyngodon wafwarwamenwomennerinerl 86868 UAZ 4810 12 4861 1960 10903 10971 1961 11034 centrorhynchus sp cystcystacanthacanth 86869 1964 15252 15258 15471 15541 15708 1967 1969 great basin naturalist 573 C 1997 appp 278 280

RANGE EXPANSION OF WHITE TAILED DEER odocoileus virginianusVIRGIN IANUS INTO URBAN AND agricultural AREAS OF UTAH

mark FE McCluremcclurellmcclurel2mcclure1212 john A Bissonettebissonettel2bissonette1212 michael R ConconoverlConoverconover1overl1 and dennis D austinaaustin3

keyketkeikhi words range expansion urban areas utah white tailed deer

numbers of white tailed deer odocoileus their presence has been reported in the litera- virginianusvirgimanusvirgivirginlanusianusmanusmanns in the united states have increased ture and that some persons think they occa- to an unprecedented level over the past 50 60 siosionallynally see them durrant however did not yr mccabe and mccabe 1984 curtis and provide any references hall and kelson 1959 richmond 1992 harlow and guynn 1994 1011 list an account of white tailed deer north during this time range expansion by white of ogden utah they reference miller and kel- tailed deer has occurred in portions of the logg 1955802 who in turn reference bailey western united states regions dominated his- 1932 bailey however does not mention utah toritoricallycally by mule deer 0 hemionushermonus martiniamartinka in his account consequently confirming the 1968 baker 1984 wiggers and beasom 1986 existence ofwhite tailed deer in utah has been mackie 1995 because white tailed deer adapt problematic and until now appears not to have readily to man altered environments range ex- been documented definitively pansion has been linked with land use changes this note documents our observations of brought about by humans baker 1984 dusek 2 white tailed deer in cache valley of north- et al 1989 wood et al 1994 accordingly ern utah on 8 february 1996 we sighted range expansion into the west by white tailed and photographed a male white tailed deer deer has been most conspicuous in agricul- in an urban environment near logan UTM tural kufeld and bowden 1995 and urban 4623500n 433100e this deer accompanied vogel 1989 environments where white tailed a radio collared doe mule deer and we were deer may now outnumber mule deer therefore able to relocate him until 10 april at white tailed deer currently occupy the lar- which time he was killed by an automobile gest geographic range of any native terrestrial during those 2 mon this deer confined his mammal in north america pagel et al 1991 activities to a 1 km2 area and we frequently thentheir present distribution in the united states observed him from 20 in away we identified includes every contiguous state halls 1978 him as a white tailed deer based on his white hall 1981 baker 1984 smith and rhodes 1994 underunderpartsunderpantsparts tail chin and neck the distinctive although halls 1978431978 43 and hesselton and white band around his muzzle the white rings hesselton 19828781982 878 question whether their around his eyes the white fringe around his distribution into utah is verifiable undocu- large tail as well as his tail flagging behavior mented sightingssigh tings of white tailed deer have been when startled or in flight when in flight this reported in northern utah in the last 50 60 yr deer galloped a gait that differs from the stot however we have been unable to find sub- of a mule deer and the bound of a whitetail stantial evidence ege g harvest records photo- mule deer hybrid lingle 1993 his antlersanglers graphs that confirmed their existence durrant 3 pt X 2 pt also characteristic of white tailed 19524561952 456 did not list white tailed deer as a deer were formed from I1 main beam curving species occurring in utah but did suggest that outward and forward the unbranched tines

I1 d1dadcpiitmcntotrishciicstnentof fihericFi herlehericherie andinclinci wildlife utah stitestatestute university loginlogloganin UT 84322 united stitesstates geologicgeological il suivcysurvey Biologicbiologicalil resource division utah cooperative fish mdand wildlife research unit utah state university logan UT 84322529084322 5290 utihrutih3utaljgutalj division ofolwildlitcrcsouiccsWildlife resources 515 eistelsteast 5300 south ogden UT 84405

278 199719971 NOTES 279

extended from this main beam after he died municationmunication in idaho numbers of white tailed we estimated his age to be 151.5lsis 2 yr based on deer have increased around idaho falls within tooth eruption and wear the past 6 yr R smith personal communica- on 2 march 1996 we sighted a and2nd white tion and they are known to exist as far south tailed deer in an agricultural environment as american falls C anderson personal com- near smithfield UTM 4635200n 431100e municationmunication 11911.9ilg km from the ist deer using binoculars white tailed deer may be better adapted from an elevated position we observed this deer than mule deer to survive in intensively culti- for 2 h at a distance of 50 200 m because this vated agricultural and urban areas vogel 1989 deer did not have antantlersanglerslers nor was there any we therefore speculate that changes in land indication of antler loss we concluded it was a use practices related to agriculture and urban- female we did not observe this deer urinate ization brought about by man during the last which would have provided further evidence of century have created patches of habitat in utah sex we identified her as a white tailed deer that appear to be suitable for white tailed deer based on the same defining characteristics listed the environments in which we observed these above except antler shape throughout the ob- 2 white tailed deer provide preliminary support servation period this deer associated with a for this speculation As the connectivity of urban group of 16 mule deer an occurrence that and agricultural developments increases and accentuated her defining behavioral and mor- the open landscape of the west is fragmented phphological characteristics based on her rela- continued expansion by white tailed deer into tively large size and coloration pattern we formerly unoccupied regions is conceivable believe she was an adult 252.5 yr although both deer appeared to be pure literature CITED bledsbreds based on their behavioral and morphological characteristics it is possible that they BAILEY V 1932 the northwestern white tailed deetdeer pro- could have been whitetail mule deer hybrids ceceedings of the biological society of washington 45 43 hybrids will frequently exhibit intermediate 44 BAKERD AKER RHR H 1984 origin classification and distribution morphological and behavioral traits but these pages 1 18 in LKL K halls editor white tailed deer traits are not always obvious and accurate ecology and management stackpole books harris- identification may be difficult in the field day burg PA 1980 genetic studies eg protein electro- CRONIN MAM A ERE R VYSE AND DGD G CAMERON 1988 genetic relationships between mule and white phoresis or mitochondrial DNA analysis pro- deer tailed deer in montana journal of wildlife manage- vide more definitive evidence of hybridization ment 5232052 320 328 cronin et al 1988 but we did not perform curnsCURTIS PD AND MEM E RICHMOND 1992 future challenges these studies consequently our conclusion that of suburban white tailed deer management transac- these 2 deer were purebred should be viewed tions of the 57th north american wildlife and nat- resources with caution ural conference 5710457 104 114 DAY GIG I1 1980 characteristics and measurements of cap- if the unconfirmed sightingssigh tings of white tailed tive hybrid deer in arizona journal of wildlife man- deer within the past 60 yr were accurate the agement 2543425 434 438 animals we observed would not represent the DURRANT SDS D 1952 mammals of utah university of ist record of this species to exist in the state kansas publications lawrence 549 appp DUSEK GLG L R J MACKIE J D HERRIGES JR AND BBB B to our knowledge however these are the first 2 RJ JD COMPTON 1989 population ecology of white tailed documented accounts of white tailed deer in deer along the lower yellowstone river wildlife utah although we cannot ascertain how these monographs 1041104 1 68 2 deer arrived in cache valley it is possible HALL RER E 1981 the mammals of north america john that they or their progenitors emigrated from wiley and sons new york 1175 appp either or idaho HALL RER E AND KRK R KELSON 1959 the mammals of wyoming white tailed deer north america ronald press company new york in wyoming appear to be increasing and ex- 1083 appp panding their ranges westward through agri- HALLS LKL K 1978 white tailed deer pages 43 65 in JLJ L cultural and riparian areas pauley and lindzey schmidt and DLD L gilbert editors big game of northnor th 1993 within the past 10 yr sightings of white america stackpole books harrisburgHarnsburg PA HARLOW R F AND C GUYNN tailed deer and hybrids have occurred in the RE DCD JR 1994 population change and loss of habitat pages 218 223223min D gerl- southwest green comer of wyoming near river ach S atwater and J schnell editors deer stack- and kemmerer T christiansen personal com pole books mechanicsburgMechanicsburg PA 280 GREAT BASIN naturalist volume 57 illliiHESSEXFONssi llonLION WT AND RM HLSSLLIONHLSSELTON 1982 white tailed PAGELGEL MDM D RMR M MAY AND ARA R COLLIE 1991 ecological deer pages 878 901 in JAA chapman and GAG A feld aspects of the geographical distribution and diversity hamelharnerhanner editorsediedltoistols wild mammals of north america of mammalian species american naturalist 137 biology management and economics johns hopkins 791 815 university fresspipressess baltimoreBaltimoiemole MD PAULEYUJLEY G AND F LINDZEY 1993 historic and current kufielddufieldKukuikul ILLDFIELD RC AND DC BOWDENBOWDCN 1995 mule and white distribution of white tailed deer in wyoming wyom- tailed deeldeer inhabiting eastern Coloiacoloradodo plains aveinveiriver ing cooperative fish and wildlife research unit botbottomstorns technicalreehrechnical publication 41 colorado division university of wyoming laramie 69 appp of wildlife fort collins 58 appp SMITHMITH MHM H AND 0OEE RHODES JR 1994 the subspecies LINGILINGLE L S 1993 escape gaits of white tailed deer mule page 90 in D gerlach S atwater and J schnell deeldeer andtindkind thentheir hybrids body configuration biome- editorsediedltoistols deer stackpole books mechanicsburgmechanicsbuigMechanicsburg PA chanicschanieschanlescs andkind function canadian journaljouinal of zoology VOGELOGEL WO 1989 response of deer to density and distri- 7170871 708 724 bution of housing in montana wildlife society bul- mackleMACKIEMAC kitkiekle RJR J 1995 mule deer elk and whitewhitetailstails recent letin 1740617 405406 413 ti endstrends and futuree management in an ecosystem con- WIGGERsIGGERS EYE P AND SLS L BEASOM 1986 characterizationCharaccharactercharacteiteitel ization text page 97 in pi oceoceedingsproceedingsedings of the western states ofofsympatncsympatric or adjacent habitats of 2 deer species in and provinces 1995 joint deeldeer and elk woikwolkworkshopshop west texas journal of wildlife management 50 idaho department of fish and game boise 129 134 MARTINKAmaiuinkacjCJ 1968 habitat relationships of white tailed WOODOOD AKA K RJR J MACKIE AND GLG L DUSEK 1994 where deeldeer and mule deeldeer in northern montana journal of the species come together pages 344 350 in D gerl- wildlife management 3255832 558 565 ach S atwater and J schnell editors deer stack- mcabemccalmMCABL RE AND TR MCCABE 1984 of slings and pole books mechanicsburgMechanicsburg PA anoisanowsarrows an historicalhistoncal retrospection pages 19 72 in LKL K halls editor white tailed deeldeer ecology and received 5 august 1996 management stackpole books Harnsharrisburgburg PA accepted 24 january 1997 MILLERmil 11 K GSG S jnJR AND R KELLOGG 1995 list of northnor th amerleanamerican lecentrecent mammals united states national museum bulletin 205 954 appp great basin naturalist 573 0 1997 appp 281 282 establishment OF THE TUNGID FLEA TUNCATUNGA MONOSITUS siphonaptera PULICIDAE IN THE UNITED STATES

michael W hastriterlhastnter1

key words tunga monositusmono situs siphonaptera distribution

the flea genus tunga containing 9 species pinna two rodents had 2 T monositusmonositus each I1 has a neotropical origin occurring on edentatesEdentates newly attached and the other a fully developed CTT bondari wagner T penetrantpenetranspenetrans linnaeus T neosomenewsome one of the 6 rodents had scarring terashaterasma jordan T travassositravassosi pinto & dreyfus reminiscent of a recent postneosomic infesta- and rodents of the families muridae T caecata tion based on descriptions of the degree of enderlein T callidacalliea li & chin T caecigena engorgement during the feeding processes of jordan & rothschild and cricetidae T libis T monositusmonositus by lavoipierreLavoipierre et al 1979 I1 esti- smit T monositusmonositus barnes & radovsky 1969 mated that 2 of the fleas had been attached for introduced specimens of T penetranspenetrantpenetrans have less than 24 h while the others had attained been reported several times in the united complete neosomy states but established populations have not As a corollary to the presence of T monositusmonositus been documented of particular interest eggs in northern latitudes north ofof31n31n to 37n37 N and adults of T penetrantpenetranspenetrans were reportedly 2 allied species T caecigena and T calcaicalhdacallieacallidalidaiidailda are taken from the remains of a dog removed from found north of 25n25 N and the former extends the cistscasts of basketmakerbasket maker indian excavations as far north as 33n33 N allailil other species of tunga in northeastern arizona wilson 1933 but smit with exception of the widely distributed T 1960 examined 4 of WilsowilsonrrsrCss slide prepara- penetrantpenepenetranstrans are found at latitudes south of these tions and was unable to verify such findings it is reported in jordan 1962 that T caecigena during a recent collection trip to southwest- china and japan is thought to be a univoltine ern utah 11 13 january 1997 I1 collected 7 species collected only during the cold season specimens of T monomonositussitus adjacent to the south- with optimal temperatures between 10 and west boundary of zion national park washing- 16c16 C and T caladacallidacalhdacalliea southern china is found ton county utah elevation 1275 m previously only during winter months november march this species had been reported only at the such evidence would suggest that T monositusmonositus type locality of cape of san quintin 3027n30027n might also be collected in the mild but temper- ilg116iigaiig0 12w baiabaja california mexico by barnes ate utah locality from october through april and radovsky 1969 and on san martin island although the 2 localities are separated by elevation sea level 200 m specimens on san 1435 km and differ in elevation by more than martin island originally reported as T caecata 1000 m they have similar mild xerotic climatic by banks 1964 later proved to be T donosimonosi conditions desert flora and host populations tus according to barnes and radovsky 1969 ege g P maniculatusmamculatus wagner P eremicusemmicusemmiouseteereemmicus and this paper presents the ist record of tunga neotoma sppapp it is therefore not surprising T monositusmonositus established in the continental that T monositusmonositus was collected in the unique united states twenty one rodents were exam- habitat and mild climate of southwestern utah ined and 6 29 harbored this flea positive thorough examination for sessile fleas through- hosts included peromyscus eremicuseremicus baird out the southwest during the winter months 292 Y P crinitis merriam 1 I1 Y 26 and neo- may result in future collections of this peculiar toma lepidacepida thomas 1 I19Y all attachment sites flea perhaps it would be prudent to collect in were restricted to the external base of the ear the locality in which wilson 1933 claims

untelmonteonte L bean life science museum brigham young university provo UT 84602

281 282 GREAT BASIN naturalist volume 57 to have found tunga penetrantpenetranspenetrans among the tion of all stages journal of medical entomology 6 basketmakerbasket maker indian ruins in arizona 19 36 JORDAN K 1962 notes on tunga caecigena siphonaptera tingidaetungidaeTungidae bulletin of the british museum natural acknowledgments history entomology 1235212 352 364 lavoipierreLAVOI PIERRE MMJM M J FJ RADOVSKY AND PD BUDWISER appreciation is expressed to nancy adams 1979 the feeding process of a tingidtungid flea tunga curator of siphonaptera collection national monositusmono situs siphonaptera tingidaetungidaeTungidae and its relation- museum of natural history washington DC ship to the host inflammatory and repair response journal of medical for kindly entomology 1518715 187 217 providing type specimens smitSMITFGAMEGAM 1960 on two archaeological records offleasof fleas siphonaptera proceedings of the entomological literature CITED society of washington 6226262 262 264 WILSON GEG E 1933 three prehistoric parasites science BANKS RCR 1964 birds and mammals of the voyage of 77560 the gringabringaGrgungainga transactions of the san diego society of natural history 1317713 177 184 received 3 february 1997 barnesBARNCS AMA M AND FJ RADOVSKY 1969 A new tunga accepted 9 april 1997 siphonaptera from the nearctic region with descnpdescripdescript information FOR AUTHORS the great basin naturalist welcomes previously VOUCHER specimensauthorsSPECIMENS authors are encouraged to unpublished manuscripts pertaining to the biologi- designate properly prepare label and deposit cal natural history of western north america high quality voucher specimens and cultures docu- preference will be given to concise manuscripts of menting their research in an established permanent up to 12000 words simple species lists are dis- collection and to cite the repository in publication coucouragedraged references IN THE TEXT are cited by author and SUBMIT manuscripts to richard W baumann date eg martin 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 or illustrations 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 submitted elsewhere authors should lowing formats adhere to the following guidelines manuscripts not so prepared maybemay be returned for revision mack GD and flake 1980 habitat relation- great LD manuscript preparation in general the ships of waterfowl broods on south dakota basin naturalist follows recommendations in stock ponds journal of wildwildlifelifeilfeiloe management por scientific style and format the CBE manualforManumanualalforfor 44695 700 publishers authors editorsandEditoreditorssandand ath6th edition sousa WPWE 1985 disturbance and patch dynamics counecoonccouncilcouncil of biologyyeditorseditors 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- 312 201 0101 FAX 312 201 0214 we do however 3122010101 3122010214 gy of natural disturbance and patch dynamics differ in our treatment of entries in literature cited academic press new york authors may consult vol 51 no 2 of this journal coulson RN and 1984 ento- for specific instructions on format these JA witter forest instruc- mology 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 TABLETABLESS are spaced of the great basin naturalist for changes double on separate sheets and designed to fit the width of either a single column TYPE AND DOUBLE SPACE all materials including use literature cited table headings and figure legends or a page lowerlowercasecaseease letters to indicate foot- avoid hyphenated words at the righthandright hand margins notes OF FIGURES use WordwordperfectPerfect s italics feature for words to be photocopies are submitted initially printed in italics use standard bond 22x28 cm with the manuscript editors may suggest changes leaving 25 cm margins on all sides lettering on figures should be large enough to withstand SUBMIT 3 COPIES of the manuscript 5 copies of reduction to one or two column width fish manuscripts and the original on a 35 inch disk originals must be no larger than 22x2822 X 28 cm utilizing WordwordperfectPerfect 51si5.1 or above number all NOTES if the manuscript would be more appro- pages and assemble each copy separately title priate as a short communication or note follow the page abstract and key words text acknowledg- above instructions but do not include an abstract ments literature cited appendices tables figure A CHARGE of 50 per page is made for articles legends figures published the rate for individual subscribers will TITLE PAGE includes an informative title no longer be 35 per page however manuscripts with com- than 15 words names and addresses of authors a plex tables andor numerous photographs may be running head of fewer than 40 letters and spaces assessed an additional charge reprints may be pur- footnotes to indicate change of address and author chased at the time of publicationofpublication an order form is to whom correspondence should be addressed if sent with the proofs other than the first author FINAL CHECK ABSTRACT states the 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 faxnumberFAX 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 ISSNissn0017001736140017 3614 GREAT clarinriarBASINclarIN naturalist vol 57 no 3 july 1997 CONTENTS articles spider wasps of colorado hymenoptera Pompipompilidaelidde 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 Tltlobolus7lobolusobolus utahensis chamberlin with remarks on T frederickfredericksonisoni 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 JMJ M phelps iliIII111 and JSJ S griffith 238 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 PMEM eckel JA hoy andjcelliottand JC 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 of myofibrogranuloma on serum calcium levels in walleye stizostedion vitreum craig A shoemaker and harry L holloway jr 268 notes helminthshelminthes from the sonoran spotted whiptail cnemidophorus noraesonoraexonoraeso and the western whiptail cnemidophorus tigris sauria teiidae from southern arizona with comments on abbreviateabbreviata terraterrapenispenis nematoda physalopteridae stephen R goldberg charles R bursey and hay cheam 273 range expansion of white tailed deer odocoileus virginvirginianusvirginivirginiamisianusfanusamis into urban and agricultural areas of utah mark FE mcclure john A bissonette michael R conover and dennis D austin 278 establishment of the tingidtungid flea tunga nwwsitusmonositusmonositus siphonaptera pulicidae in the united states michael W hastriter 281 announcing

THE GREAT BASIN SYMPOSIUM ON GLACIAL AND postglacial DRAINAGE

GREAT BASIN AQUATIC SYSTEM HISTORY shyseySSS

A T I1 G fifty years of geologic biologic and hydroclimaticHydro climatic progress in late cenozoic aquatic system history

17 21 september 1997 university park hotel salt lake city utah

the purpose of the 1997 symposium is to delineate the development of great basin drainage and its relation to the evolution and distribution of late cenozoic and recent biota using the evidence of tectonics stratigraphy geomorphology biogeography evolutionary biology paleoanthropology paleopaleohydrologyhydrology and paleoclimatology speakers discussions field excursions and poster sessions will highlight the symposium

for more information contact GBASH 8015815809801581801 5815809581 5809 2174 annex 801 5813165581 3165 FAX conferences conferadmindceutahedu university of utah wwwdceutahedugraphicsconfconfhtm salt lake city utah 84112 LEARNING FROM THE LAND scientific INQUIRY FOR PLANNING AND MANAGING THE GRAND STAIRCASE ESCALANTE NATIONAL MONUMENT

3 7 november 1997 southern utah university ttmem i aaytttewteW riajrmiry ai ATajyft y T cedar city utah s1airaaasMM J

the utah state advisory council on science and itnfrva xt JLr S AN ann41at technology and the US bureau of land management jiliiwjbra are pleased to assemble scientists planners educators and the public in a symposium designed to share knowledge and identify needs that will help determine the management approach for the grand staistalstaircasercasearcase escalescalanteante national monument N A T j 0 N laA ii jlofl 4 ii kloekroeMhronf N yrU affyffX the 2 day symposium will t tff hlteati of lawtlmmafitrtwi focus on the natural history of the monument especially geology biology archaeology and paleontology

assess data research and documentation of previous and current investigators

establish an interactive network of individuals who have a continuing involvement in studies of the area

determine additional needed scientific endeavors pertaining to the resources of the monument

transfer knowledge gained to the monument planning team and others

discipline specific and general poster sessions will be held with a panel discussion addressing the impacts of the formal establishment of the monument and future scientific research several field trips are scheduled sunday and monday prior to the symposium for more information please contact dr suzanne winters Ms marietta eaton utah state science advisor grand stastaircasei rcasearcase escaescalanteI1 ante national monument 8015381038801538801 5381038538 1038 cedar city UT 84720 email suzwintersstateutus 8018655114801865801 8655114865 5114 FAX 801 5381547538 1547 email meatonutblmgov