Great Basin Naturalist

Volume 52 Number 4 Article 17

12-30-1992

Full Issue, Vol. 52 No. 4

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GREAT BASINB A I1 naturalist

A

VOLUME 52 NO 4 DECEMBER 1992

BRIGHAM YOUNG university GREAT BASIN naturalist editor JAMES R BARNES 290 MLBM brigham young university provo utah 84602

associate editors MICHAEL A BOWERS BRIAN A MAURER blandy experimental farm university of department ofzoology brigham young university virginia box 175 boyce virginia 22620 provo utah 84602 PARRISH J R CALLAHAN JIMMIE R museum of southwestern biology university of bloBIOWESTBIGWESTBIOBIG WEST inc 1063 west 1400 north logan new mexico albuquerque new mexico utah 84321 3140 california mailing address box hemet PAUL T TUELURTUELLER 92546 department of range wildlife and forestry 1000 JEANNE C CHAMBERS university of nevada reno valley road USDA forest service research university of reno nevada 89512 nevada 89512 nevada reno 920 valley road reno ROBERT C WHITMORE JEFFREY R JOHANSEN division of forestry box 6125 west virginia uni- department of biology john carroll university versityversity Morganmorgantowntown west virginia 26506612526506 6125 university heights ohio 44118

PAUL C MMARSHARSH center for environmental studies arizona state university tempe arizona 85287

smith zoology claciaclaytononmjonmM editorial board richard W baumann chairman zoology H duane anyrangeanytangeRange white zoology berranjerran T flinders botany and range science william hess botany and bangerange science all are at brigham young university ex officio editorial board members include clayton S huber dean college of biological and agricultural sciences norman A darais director university publications james R bames 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great basin naturalist the great basin naturalist founded in 1939 is published quarterly by brigham young university unpublished manuscripts that further our biological understanding ofthe great basin and surroundingandurrounding areas in western north america are accepted formor publication subscriptions annual subscriptions to the great basin naturalist for 1992 are 25 for individual subscribers 15 for student and emeritus subscriptions and 40 for institutions outside the united is 12 back issues are in print and states 30 20 and 45 respectively the price ofsingle issues all inerintfrintErint available for sale all matters pertaining to subscriptions back issues or other businesbubinesbusiness shouldouldouid be provo directed to the editor great basin naturalist 290 MLBM brigham young university UT 84602 scholarly exchanges libraries or other organizations interested in obtaining the great basin naturalist through a continuing exchange of scholarly publications should contact the exchange librarian harold B lee library brigham young university provo UT 84602 editorial production staff joanne abel technical editor jan spencer assistant totheto the editor natalie miles production assistant

copyright 1992 by brigham young university ISSN 001736140017 3614 official publication date 30 december 1992 129212 92 7503555tso750 3555 the great basin naturalist PUBLISHED AT PROVO UTAH BY BRIGHAM YOUNG university

ISSN 0017 3614

VOLUME 52 DECEMBER 1992 no 4

great basin naturalist 524 appp 293 299

WINTER NUTRIENT CONTENT AND DEER USE OF GAMBEL OAK TWIGS IN NORTH CENTRAL UTAH

I1 rosemary L pendleton fred J wagstaffWagstafe and bruce L welch

ABSTRACEABSTRACT we examined winter nutritional quality of current year bud and stem tissues from burned and unburned stands of gambel oak quercus gambmmbelngambeliigambellieliieili nutt nutritional analyses were based on the amount of forage consumed by wintering mule deer deer use along the utah valley foothills averaged 6256 25 10710.710 7 cmern of current year growth of the tissues examined post fire bud tissue had the highest nutrient content with a mean of 9519 51 crude protein 0190 19 phosphorus and 34034 0 in vitro digestibility composite values bud stem for unburned stands were slightly higher in crude protein and phosphorus and lower in digestibility than those reported in previous studies nutrient values from burned stands were significantly higher than those of unburned stands for all three measures tannin content of the burned area regrowth was also higher overall forage value of gambel oak to wintering mule deer isis relatively low

key words quercus gambgambeliigambellielii odocoileus hemionus nutrients foraging behavior utilization browse winter

gambel oak quercus gambelligambeliigambelii nutt is a 17 browse species based on time spent browsing valuable year round source of food and cover for and plant weight consumed many wildlife species including deer elk big- although important to wintering mule deer horn sheep small mammals and a variety of in terms of forage availability and palatability birds reynolds et al 1970 harper et al 1985 oak ranks among the bottom in nutritional value tirmenstein 1988 because of its abundance smith 1957 bunderson et al 1986 autrinutri and location oak is an important food source for tionaldional studies report winter oak browse as being wintering mule deer providing up to 75 of the low in essential nutrients and digestibility available winter browse along the wasatch smith 1957 kikufeldifeldafeld et al 1981 meneely and front perry 1980 winter use of oak varies schemnitz 1981 smith and hubbard 1954 with location but it has been reported high in described oak as being well liked but of low some areas along the wasatch front declining forage quality currently little information is in the presence of more palatable rosaceous available on the nutrient content of different shrubs smith 1952 julander 1955 deer use portions of the plant stem or on the selection of in western north america ranges from moder- plant parts by deer ate to heavy throughout the year kufeld et al the effect affireoffireof fire on the nutritional status of 1973 and references therein in winter prefer- oak browse is also of some interest to land man- ence trials smith 1950 and smith and hub- agers fire may provide an effective manage- bard 1954 ranked oak as ath7th or higher out of ment tool for opening the canopy of the more

1 shrub sciences laboratory intermountainIntermount mi researchRe seaichsealch station USDA forest service 735 north 500 eastE ut provoprove utah 84606

293 294 GREAT BASIN naturalist volume 52

primarily of gambel oak and sagebrush arte- misia hitritrlfridentata sspasp vastyanacaseyanavaseyanacasevaseganayana with scattered patches ofcliffroseofcliffroseeilfcliffrose cowaniadowaniaCowania stansburstansburianaiana and bitterbitterbrushbrush pulpuipurshiashia tridentata all three loca- 0 PG tions are heavily used by wintering mule deer 18 A in august 1987 a wildfire burned approximately tu 1270 acres on the southwest facing slopes above 15 orem and lindon utah oak present on the 0 bum showed considerable regrowth two months following the fire two study sites were established at the lindon location one on the bum itself the other in the adiacentadjacentadiaadlacent unburned study were alsoaiso established at zaz1 r vegetation sites ilcvic the pleasant grove and hobble creek canyon locations for a total of four study sites deer utilization was determined by measur- ing the length of marked twigs before and after browsing in november of 1987 679 twigs on the lindon burn site and 660 twigs on the adja- 10 km cent unburned site were marked with colored plastic tape twigs were selected from around the periphery of multiple clones to represent all fig 1 location of four oakbrushoakbrusboakbrush study sites in utalutaiutah county utah PCPG pleasant grove LB lindon burned directional aspects and a variety of heights LU lindonundon unburned HC hobble creek canyon accessible to deer twig lengths were measured from the tape to the end ofthe terminal bud in march 1988 the twigs were measuredreremeasured and the oak thickets to allow greater herbaceous dense number ofcentimeters browsed determined for growth anonymous 1966 dills 1970 hallisey each twig ratio ofbud tissue and twig tissue 1976 et al 1985 deer use the and wood harper consumed by deer was then calculated the of browse has been found to increase species procedure was repeated at the pleasant grove fire in some 1938 hallisey and following horn and hobble creek sites the following year wood 1976 but not all kufeld 1983 cases where 186 were marked and measured at content of some oak species has twigs the nutrient each been reported higher following burning site twenty two samples for nutritional analysis hallisey and wood 1976 meneely and collected at midwintermid winter from 12 burned schemnitz 1981 were 10 unburned oak clones at the lindon loca- of this study was to provide a and the intent portions of each of the burned clones were more accurate assessment of the nutrient con- tion november to ensure availability tent of oak forage consumed by wintering mule fenced in early mid collection material in late janu- deer on the wasatch front specific objectives of midwinterwinter 200 300 stems were removed from each were 1 to determine what portion of cambelgambel ary snow and transported to the oak twigs was used by wintering mule deer in clone packed in collected from all sides utah valley 2 to determine the percent crude laboratory twigs were eliminate protein phosphorus and in vitro digestibility of oftheodtheof the periphery of each clone to pos- terminal buds and stems ofgambelofgambel oak and 3 sible differences due to directional aspect at to compare values obtained from adjacent the laboratory stems from each clone sample burned and unburned stands were divided into a lemiem1 cm terminal bud portion and an adjacent 10 cm stem the proportion of current year growth sampled 11 cm was MATERIALS AND METHODS approximately equal to that removed by winter- ing mule deer where stem lengths measured deer utilization was studied at locations near less than 11 cm total current year growth was lindon utah above pleasant grove utah and used in the analysis twig diameters at I1 and 5 in the mouth of hobble creek canyon fig 1 cm from the tip were also recorded vegetation at these foothill locations consists the ensuing 44 bud and stem tissue samples 199211992 WINTER NUTRIENT CONTENT OF GAMBEL OAK 295

TABLE 1 summary ofdeeiof deer utilization on marked twigs of gambel oak at four study sites in utah county utah

no twigs no twigs percent mean marked browsed browsed utilization cm1cmaacma lindonlmdonamdon bumedbumen 679 194 286 107 t 0044a441 lindon unburned 660 368 558 107 t 024 hobble creek 186 112 602 63 039 pleasant grove 186 157 839819 77 t 033

mean standardstandlstandiird error were ground using liquid nitrogen and stored at percentage data were arcsine transformed soc80 C in vitro digestibility crude protein and and analyzed using the general linear models phosphorus were determined for both bud and GLM routine available on SAS the model stem portions these three measures were con- used was a 2 X 2 factorial design with bubbumburn sidered sufficient to determine overall nutri- treatment burned unburned and tissue type tional quality of oak as they are the nutrients bud twig as main effects clone was used as most commonly deficient in winter diets of the error term for the burn treatment main range animals welch et al 1986 in vitro effect tissue differences were also examined digestibility was assessed using pearsonpearsonss 1970 separately for burned and unburned areas modification ofthe tilley and terry 1963 tech- because of a significant burn treatment X tissue nique this technique while possibly overesti- interaction mating in vivo digestion of cell contents in tannin containing forages robbinsbobbins et al 1987 RESULTS nastisanastis and malechekmaleemaieeMalechekbek 1988 remains the easiest 10.7107 and most accurate of the in vitro techniques deer use at the lindon sites averaged 107 cm for both burned and unburned clones nastisanastis and malechek 1988 and is commonly table 1 individual use varied widely employed in nutritional studies of range forages twig ranging from lsis1.5 to 33 cm although mean use at the two inoculum for the digestion trial was obtained 15 lindon sites was the same the burned area had from a slaughterslaughterhousehouse steer coa injected the a greater proportion of small bites than the inoculum was processed within 45 minutes of unbumedunfumedunbumed area 2 over 24 of the bites removal from the rumen milchunas and baker fig werewere in the 151.5isls 5 cm category at the burned site 1982 studies have shown that inoculainocular obtained as compared to 575.7 in this category at the from domestic can successfully ruminants unburned site also a smaller percentage of of forages to approximate digestibility range marked twigs was browsed in the burned area palmer and cowan 1979 welch et al deer table 1 mean use at the pleasant grove and 1983 phosphorus and crude protein determi- hobble creek sites during the milder 1988 89 plant nations were made at the and soil analysis winter was somewhat less than at the lindon laboratory at brigham young university crude sites averaging 777.7 and 636.3 cm respectively protein was based on kjeldahl nitrogen content table 1 A technicon auto analyzer technicon instru- results from the nutrient analysis of sam- ment corp tarrytown NY was used to deter- pled tissues are given in table 2 main effects mine phosphorus content to simplify from the analysis of variance were all highly comparisons with values reported in the litera- significant post burn sprouts contained more ture composite values for the complete ilem11 cm crude protein and phosphorus and were more sample were calculated as follows composite digestible than unburned samples bud tissue value 10twig10 twig value bud valuelllueIliueli bulk exceeded stem tissue in all three measures the samples made up ofofonecofoneone twig from each sampled interaction term was also highly significant for clone were tested for tannin content twigs crude protein and phosphorus p 0001.0001 and were kept frozen at 80 C until use then p 002002.0020021.00211 respectively running separate anal- ground under liquid nitrogen tannin content yses for burned and unburned areas revealed for each bulk sample was determined at the that the difference between bud and stem plant and soil analysis laboratory using values was greatest for post bumburnburmbumm sprouts creat- hagermanHaehagerman s 1987 radial diffusion method ing the significant interaction term bud and 296 GREAT BASIN naturalist volume 52

25 undon burned lindon unburned

20

15

10

0 0 Q

0 2 4 6 8a 10 12 14 1618 18 20 22 23 2 4 6 a 10 12 14 16 18 20 22 23 4

Q 35 pleasant grove hobblehobbie creek Q

25

20

13 &

10

5

01 120 10 14 1618 18 20 22 23 12 1 23 2 4 6 8 2 4 6 8a 10 12 14 is 18 20 22 twig utilization cm

fig 2 distribution of stem utilization at four oakbrushoakbrush study sites in utah county utah

41 compared TABLE 2 attained significance values from analyses of tent of414.1 mopermgpermg per 100 mg plant tissue variance for nutrient content of gambel oak with 343.4 mg for unburned clones the stem tissue sample derived from burned clones had a source of crude tannin content of 161.6lgig mg per 100 mg plant tissue variation protein phosphorus digestibility compared with 070.7 mg for unburned clones bumburn treatment 00001 00002 00001 tissue type 00001 00001 00001 discussion bumburn X tissue 00001 00021 03519 clone 00001 00228 00015 previous reports on fall winter nutrient con- tent of gambel oak twigs from mature stands range from 464.6 to 575.7 for crude protein from twig values from burned clones differed signif- 0090.09 to 0100.10 for phosphorus smith 1957 for all three variables table 3 bud and icantly kufeld et al 1981 meneely and schemnitz unburned clones differed only twig values from 1981 and from 26626.6 to 40240.2 for in vitro burned and in nitrogen content twigs from digestibility kufeld et al 1981 meneely and unburned clones also differed in appearance schemnitz 1981 similar values have been burned twigs being more slender at I1 cm isls181.8 obtained for other oak species meneely and mm vs 262.6 mm p 0002.0002 and at 5 cm 212.1 mm schemnitz 1981 composite values from vs 292.9 mm p 0001.0001 unburned stands in our study table 2 are sim- burning also had a significant effect p ilar to previous results though slightly higher in 0001.0001 on tannin content the bud tissue sample crude protein and phosphorus and slightly derived from burned clones had a tannin con lower in digestibility the use of different twig 199211992 WINTER NUTRIENT CONTENT OF GAMBEL OAK 297

TABLE 3 means and standard errors for percent dry matter nutrient content of bud and twig samples collected froinflorn burned and unburned oak stands growing near lindon utah letters following means indicate significant differences p 0001 between bud and twig values ahinwhinwithin burn treatment composite values 10twig10 twig value bud valuevaluedvalue1vdluej111 1

crude protein phosphorus digestibility burned stands bud 95 036 a 019 t 0 006800068 a 34034.034 0 059 a twig 75 t 022022bb 013 0 0092000920.0092 b 29829 8 0700.70070bb composite 77 023 014 00087 30230.230 2 066 unburned stands bud 65 009 a 0 12 0 005600056 a 26226.226 2 168 a twig 57 OJOoao010bb oiloii011 0 008600086 a 23523.523 5 115 a composite 58 010 oilolioii011 0 008100081 23723.723 7 116 lengths for nutrient analysis did not affect the tissues sampled had a crude protein content of overall results significantly composite values less than 10 the composite value from based on twig lengths of 777.7 or 636.3 cm mean burned clones was less than 8 which ranks deer utilization at pleasant grove and hobble below that of sagebrush aspen and rosaceous creek differed very little from those based on shrubs smith 1957 kufeld et al 1981 debyle the liemilem11 cm sample in the absence of foliage et al 1989 actual amounts of protein digested buds provide the highest source of nutrients may be somewhat less than predicted by the in during late fall and winter values for leaves of vitro technique tanninsfannins present in summer oak gambel oak are substantially higher in summer and other forages have been found to increase but comparable to bud tissue by midwintermid winter the fecal excretion of protein by domestic live- urness et al 1975 meneely and schemnitz stock robbinsbobbins et al 1987 in mule deer and 1981 welch et al 1983 austin and umess other browsers nitrogen excretion may be 1985 reduced by tannin binding proteins present in burning had a significant influence on the the saliva robbinsbobbins et al 1987 winter digest- nutrient content of oak forage particularly bud ibility of gambel oak is also low when compared tissue values obtained from burned stands to other forages bunderson et al 1986 ranked were relatively higher than those obtained from digestibility of winter oak forage 25th25tb out of 27 unburned stands post bum bud tissue had by species tested our results are similar in digest- far the highest nutritional value of any tissues ibility to that listed in the ranking showing examined increased nutrient content of forage slightly higher digestiblitydigestibility for burned stands and following fire has been reported elsewhere lower digestibility for unburned stands although indications are that such an increase is deer use of oak depends on many factors temporary dills 1970 hallisey and wood 1976 including cover exposure density of oakbrushoakbrush meneely and schemnitz 1981 debyle et al and availability of other forages smith 1952 1989 the evidence to date suggests that julander 1955 kufeld 1983 austin and urness improved phosphorus content of forage due to 1985 fire affects the structure of vegetation fire is fairly short lived usually lasting only one and cover as well as quantity and quality of year nitrogen benefits may last longer depend- forage produced horn 1938 hallisey and ing on the species and season the higher values wood 1976 meneely and schemnitz 1981 reported for burned stands in this study likely kufeld 1983 whether or not deer use an area occurred because sampling took place less than more after burning appears to depend on struc- one year following the burn ture of the oak community and type of vegeta- despite the slightly higher values for crude tion present on adjacent areas as well as protein and phosphorus reported here for intensity and size of burn where oak stands unburned stands and the markedly higher form impenetrable thickets or where little values for recently burned stands the overall understory is available burning has resulted in winter nutritional value of gambel oak remains increased use by deer horn 1938 hallisey and relatively low even post burn bud tissue which wood 1976 in contrast kufeld 1983 found had the highest nutrient content of any of the increased use by elk but not deer following 298 GREAT BASIN naturalist volume 52 burning vegetation at this colorado location acknowledgments consisted of a mixture of mature oak stands sagebrush snowberry symphoricarpos albus thanks are extended to len carpenter chokecherry prunus virginiajavirgivirginiananiana and service- steve monsen and art Tiedetiedemannmaimmarm for sugges- berry amelanchier alnifolia burning elimi- tions on research design to warren clary philip nated big sagebrush plants and decreased urnessumess robert ferguson and an anonymous production of several other important browse reviewer for comments on the manuscript and pleasant grove species partially as a result of abnormally dry to john alienallenailen and the ranger weather conditions district for access to thediedle lindon burn site we found no evidence for increased use on the lindon burned site table 1 the mean literature CITED number of centimeters browsed at the burned site was identical to that of adjacent unburned ANONYMOUS 1966 conversion of thicket covered areas to stands even though twigs sprouts from the productive grazing lands USDA forest service inter- burned stands tended to be longer also a lower mountain region range improvement notes 11 14 URNESSUBNESS 1985 values of four of marked twigs was browsed at the AUSTIN D D AND P J percentage communities for mule deer on ranges with limited burned site the apparently lower use ofburned summer habitat journal of range management 38 twigs by deer despite higher nutrient content 167 171 BUNDERSONBUNDFRSON E D B L WELCH AND D J WEBERWEBEB 1986 may be due to several factors oak stands in the in vitro digestibility ofjuniperusJumperus osteospennaosteosperma torr area form discrete clones rather than large little from 17 utah sites forest science 32 834 840 impenetrable thickets important browse spe- DEBYLE N V P J URNESS AND D L BLANK 1989 burned and unburned aspen commu- cies such as sagebrush and bitterbitterbrushbrush present forage quality in nities USDA forest service intermountamintermountain research on unburned areas were lost as a result of the station ogden utah research paper INT 404 8 appp fire also a lack of cover and increased tannin DILLS G G 1970 effects of prescribed burning on deer content of forage on the burn may have had browse journal of wildlife management 3454034 540545540 545 HAGERMAN A E& 1987 radial diffusion method for deter- some effect on deer preference mining tannin in plant extracts journal of chemical the oakbrushoakbrush zone is critical to wintenwinteringng ecology 13 437 449 WOOD 1976 prescribed fire deer populations along the wasatch front HALLISEY D M AND G W in scrub oak habitat in central pennsylvania journal of although not the most preferred winter food its wildlife management 40 507507516sig516 protective cover and sheer abundance make it HARPER K TFJT F J WAGSTAFFwagstaffandlANDLAND L M KUNZLER 1985 gambel oak widely used smith 1949 smith biology and management ofthe vegetative one ofthe most type a literature review USDA forest service inter- and hubbard 1954 julander 1955 current mountain forest and range experiment station emphasis in the intermountain region is to ogden utah general technical report INT 17931179 31 appp the oakbrushoakbrush zone primarily for wildlife HORN E E 1938 some wildlife forest relationships manage transactions of the north american wildlife confer- winward 1985 several management tools ence 3 376 380 have been suggested including fire harper et JULANDERJULANDEK 0 1955 deer and cattle relations in utah 1 139 al 1985 winward 1985 burning may result in forest science 130 KUFELD R C 1983 responses of elk mule deer cattle and a temporary improvement in nutritional quality vegetation to burning spraying and chaining of as well as opening the canopy sufficiently to gambel oak rangeland colorado division of wildlife publication 34 47 allow establishment of other shrub and forb fort collins technical appp KUFELD R CCMM STEVENS AND D C BOWDEN 1981 species however without some form of follow- winter variation in nutrient and fiber content and in up treatment the proliferation of oak sprouts vitro digestibility of gambel oak quercus gamberigambehigambehi dentata from diversi- ultimately result in denser less useable oak and big sagebrush artemisia tntri may fied sites in colorado journal of range management forage and reduction of understory species 3414934 149 151 harper et al 1985 stevens and davis 1985 KUFELD R C D D WALLMO AND C FEDDEMAFEDDFMA 1973 1985 moreover the loss of fire food ofthe rocky mountain mule deer USDA forest winward service rocky mountain forest and range experi- susceptible browse species such as big sage- ment station fort collins colorado research paper brush mountain mahogany and bitterbitterbrushbrush RM 111 31 appp MENEELYMENFELY S C AND S D SCHEMNITZ 1981 chemical may have serious consequences for wintering composition and in vitro digestibility of deer browse mule deer riggs et al 1990 outweighing any three years after a wildfireaaildfire southwestern naturalist 26 possible benefit 365374365 374 199219921 WINTER NUTRIENT CONTENT OF GAMBEL OAK 299

MILCHUNAS D GcandGANDAND D L BAKER 1982 in vitro diges- based on browsing time and forage consumed journal tion sources of within and between trials variability of range management 8262 265 journal of range management 35 199 203 STEVENS R AND J N DAVIS 1985 opportunities for NASTIS A S AND J C MALECHEK 1988 estimating imuroim1roimprovingng forage production in the gambelgambol oak types digestibility of oak browse diets for goats by in vitro of utah pages 37 41 in K L johnson ed proceed- techniques journal of range management 41 255 ings ofthe third utah shrub ecology workshop college 258 of natural resources utah state university logan PALMER W L AND R L COWAN 1979 comparison of TILLEY J M A AND R A TERRY 1963 A two stage tech- deer and sheep digestive capacities journal ofwildlife nique for in vitro digestion of forage crops journal of management 43 798 801 ttheh british grassland society 18 104 111 PEARSON H A 1970 digestibility trials in vitro tech- tirmenstein D 1988 quercus gambehigambgambeliigambelinehiebieliihilihilf in fischer niques pages 85 92 in H A paulsen E H reid and W C compiler the fire effects information system K W aarderparder eds range and wildlife habitat evalua- data base USDAUS DA forest service intermountain tion a research symposium USDA forest service research station intermountain fire sciences labor- miscellaneous publication 1147 atory missoula montana magnetic tape reels 1600 PERRY W M 1980 oakbrushOakbrush management plan and envi- bpi ASCII with common LISP present ronmentalronmental assessment report for pleasant grove URNESS P J D J NEFFANDNEFF AND R K WATKINS 1975 nutri- ranger district uinta national forest 8 appp mimeo- tive value of mule deer forages on ponderosa pine graphed summer range in arizona USDA forest service REYNOLDS H GGWW P CLARY AND P F FFOLLIOTT 1970 rocky mountain forest and range experiment sta- gambelgambol oak for southwestern wildlife journal of for- tion fort collins colorado research note RM 304 estry 6868545545 547 6ppapp6 appp RIGGS R A PR J URNESS AND K A GONZALEZ 1990 WELCH B L E D MCARTHUR D L NELSON J C effects of domestic goats on deer wintering in utah PEDERSON AND J N DAVIS 1986 hobble creek a oakbrushoakbrush journal of range management 4322943 229299 234 superior selection of low elevation mountain big sage- ROBBINS C T S MOLE A E HAGERMAN AND T A brush USDA forest service intermountain research HANLEY 1987 role of tanninsfannins in defending plants station ogden utah research paper INT 3703701010 appp against ruminants reduction in dry matter digestion WELCH B L J C PEDERSON AND W P CLARY 1983 ecology 68 1606 1615 ability of different rumen inoculainocular to digest range for- SMITH A D 1949 deer forage observations in utah jour- ages journal of wildlife management 47 873 877 nal of wildlife management 13 314 315 WINWARD A H 1985 perspectives on gambel oak man- 1950 feeding deer on browse species during agement on national forests of the intermountain winter journal of range management 3 130 132 region pages 33 35 in K L johnson ed proceed- 1952 food habits of mule deer in utah journal of ingsoftheings of the thirdird utah shrub ecology workshop college wildlife management 16148 155 of natural resources utah state university logan 1957 nutritive value of some browse plants in winter journal of range management 10 162 164 SMITH A D AND R L HUBBARD 1954 preference rat- received 9 march 1992 ings forvinterfor vinterwinter deer forage from northern utah ranges accepted 18 october 1992 creatgleatgreat basin naturalist 524 appp 300 308

BOTANICAL CONTENT OF BLACK TAILED jackrabbitJACKRABBIT DIETS ON semidesert RANGELAND

1 23 2 tchouassiTchouassi bansiwansi rex D pieperpieper23 reldon F beekbeck and leigh W murray4murrayamurray

abstractabstiact botanical content of black tailed jackjackrabbitrabbit diets was deteldeterminedmined by micromieromicrohistologicalniicrohistologicalhistological examination of fecal samples collected from six different vegetation types in southern new mexico on three dates grasses comprised the largest component of jackrabbitthethejackiabbitjackjaekrabbit diets with dropseed species sporobolus sppapp and black grama bouteloua erioerloenoenopodaeriopodapoda the most abundant grasses in the diets leatherweedfeatherweedLeatherweed croton croton pittmpottsiipottmpottsiisli and silverleaf nightshade solanum elaeagnifoliumelaeagmfolium were grasses important forbsfoimol bs on most vegetation types diet composition varied in response to season and vegetation type were important during the summer glowinggrowing season while forbs were selected during their growing season summer or winter spring shrubs were less abundant in the diet than grasses and forbs

key worchwordswonch aticrohistologicalnticrohistological analysis fecal analysis lepus califomicuscalifornicuscalifornicus

black tailed jackrabbitsjackrabbits lepus califomicus similar rangeland the present study represents are widely distributed in western and central a continuation of earlier studies and should add north america they range from canada south- to understanding seasonal and yearly fluctua- ward to the states of sonora and chihuahua tions in diets of black tailed jackrabbitsjackrabbits mexico and from the pacific coast eastward to the great plains hansen and flinders 1969 STUDY AREA because of this wide distribution jackjackrabbitsrabbits potential food sources encounter a variety of study was conducted on the new and young 1980 considerable work the mcadoo mexico state university college ranch about food habits of the black has been conducted on 40 km north of las cruces new mexico the tailed jackjackrabbitsrabbits especially in arizona colo- ranch lies on the jornada plain between the san great plains rado and the arnold 1942 reigel andres mountains and the rio grande at an 1942 lechleitner 1958 sparks 1968 hansen elevation of about 1300 m wood 1969 valen- and flinders 1969 flinders and hansen 1972 tine 1970 the climate of the jornada plain is ureskUresk 1978 fagerstone etetalal 1980 johnsjohnsonon and semiarid with a yearly mean temperature of anderson 1984 these studies show that jaejacjaekjack about 16 C mean monthly temperatures are rabbits are opportunistic feeders varying their highest in june 35350 and lowest in january 13 diets depending on available forage average annual precipitation is 32 cm range in spite of the relatively large number of 929.2 36236.2 cm of which about 50 falls during publications reporting the feeding habits of july august and september paulsen and ares black tailed jackrabbitsjackrabbits few have been con- 1962 ducted in new mexico and the southwest fecal pellets from black tailed jackrabbitsjackrabbits dabo et al 1982 found jackrabbitjackrabbit diets were were collected from six vegetation types habi- composed of many species but only a few spe- tats 1 mesquite prosopis glanduloseglandulosa grass cies of grasses and forbs formed the bulk of the 2 snakeweed Gutiergutierreziagutierrezbarezia sarothsarothraerae 3 diet they found that diets inferred from fecal mixed shrub grass 4 black grama 5 analysis differed among habitats forborbbr jaekjackjackrabbitsforjackrabbitsrabbits creosotecreosotebushbush larrea trithitrndentata and 6 tar- during summer and fall in contrast fatehifatemi et bush flourensiaFlourensia camuacemua these vegetation al 1988 found similar diets among habitats on types are characteristic of desert grassland and

presentaddress for livestock mezamcameroonmezam cilmeroon D pitipiir tiretimetimm lit of animal trltriand hangerange sciences ncwmexiconew mexico state university lascruceslas Cruces Nnewwmexico8w3mexico 88003 Present address sector hezam 2epartinentdcpiii tnicnt ofanfialandol01 animal alliaiilabil range suenesuensciencese new mexico stitestate university las cruces new mexicomexico881do388003 hauthor3authoralithoi to choinwhoinwlioni colcoieol i cspondencecorrespondence lioultlshould be addressedaddi essed 4 departinentdcpaitiiicnt Experimentalofexperimentalof statistics new mexico state university lisUsluseus ciuCilcrucescruicesleestees new mexico 88003

300 199219921 BLACK TAILED jackrabbitJACKRABBIT DIETS 301 desert shrubshmblandsshrublandslands humphrey 1958 major tute 1985 proc catmodcarmod is a program for ana- grass species include black grama bouteloua lyzing relative frequency data by chi square tests eriopodaeriopoda mesa dropseed sporobolus herbage standing crop an estimate of herb- flexuosus flufffluffgrassgrass erioneuronErioneuron pulchellumpulchellum age availability was determined by clipping and threeawns aristida sppapp abundant forbs herbaceous species from ten 050.5os X 10lom m quad include leatherleatherweedfeatherweedweed croton croton pottsiipottsii rats located randomly in each of the two repli- wooly paperpaperflowerflower psilostrophe tagetinaetagetmaetagetinae cations within each vegetational type at the silverleaf nightshade solanum elaeagnifolium time the fecal material was collected herbage and other species shrubs include mesquite was separated by species oven dried 70 QC creosotecreosotebushbush and tarbush and weighed shrub biomass was determined for the major species by dimension analysis as described bvby ludwig et al 1975 preference METHODS indices werewerg calculated as the ratio between the amount each species contributed to the diet divided by the composition in the standing crop jackrabbitrabbit fecal material was collected from Jack krueger 1972 only those preference indices each vegetational type in june august and than 2 are reported in this paper to 1988 sample consisted of 15 20 greater october the indicate those with a relatively high pellets collected randomly on each date and in species degree of preference each of two replications of each vegetational type fresh pellets were identified by their shiny appearance field observations indicated that RESULTS pellets lost their shiny appearance within a week of deposition the pellets were dried and herbage availability ground to pass through a 10lomm mm screen in a wiley mill the ground material was prepared grasses contributed more than half of the as described by bear and hansen 1966 and herbaceous standing crop only on the black holechek 1982 five microscopic slides were grama type fig 1 generally grass composi- prepared from each sample and 20 random tion increased from june to august except on fields were read from each slide holechek and the creosotecreosotebushbush type summer is the major vavra 1981 individual plant species were iden- growth period for the c4ca perennial grass species tified by comparison with known reference in this area pieper and herbel 1982 forbs slides all identifications were made by the contributed more than 50 to the plant stand- senior author with an accuracy of94 calcula- ing crop on the mesquite grass black grama tions of percent composition by weight were and snakeweed types fig 1 shrubs were made following procedures outlined by abundant contributing about 20 of the stand- holechek and gross 1982 ing crop on the creosotecreosotebushbush tarbush and Micromicrohistologicalhistological examination offecal mate- mixed shrub grass types rial has some limitations in diet evaluations composition holechek et al 1982 problems are related to diet differential digestion of different species seasonal changes in jackrabbitjackrabbit diets sidahmedSidahmed et al1 1981 differential detection appeared to be greater than standing crop avail- and recognition under a microscope westoby ability for grasses forbs and shrubs fig 2 et al 1976 and differential particle size reduc- table 1 generally grass content of the diet tion crocker 1959 in spite of these limita- peaked in august and declined until october tions fecal analysis is one of the main methods fig 2 forb content of the diet changed little for quantifying diet composition of wide rang seasonally for pellets collected on the tarbush ing herbherbivoresivores creosotecreosotebushbush and snakeweed types forbs statistical analyses of dietary data were comprised a larger percentage of the diet in based on species counts using a split plot com- june and october than in august on the mes pletely randomized design with vegetational quite grass and black grama types shrubs gen- type as the whole plot and sampling date as the erally contributed less than 25 of the diet split plot differences among types periods and except for pellets collected from shrubby types the interaction were analyzed using a categorical at certain dates eg october on the mesquite modeling procedure proc catmodcarmod SAS instiansti grass type october on the snakeweed type rvolume 52 302 GREAT BASIN naturalist

creosotebushCreosotebush snakeweed type loo100 loo100 0 0 0 0 0 0 ae 70 10 shrubs 7e 70 0 60 forbs 60 CL E E so 0 50 grasses 8 40 E 40 X 8 30 shrubs 30 X 0 20 0 20 forbs 10 10 grasses 0 0 june august october june august october

mixed shrub grasstype tarbush type loo100 loo100 0 0 0 ca 2.2 c0 70 70 0 60 X C0L 60 E 0 so X E so x 0 40 40 C P 0 shrubs 30 shrubs IL 20 forbs Q forbs 10 grasmse 10 gr ssesases 0 june august october june august october

grass black grama type mesquite type 100 loo100 loo 90 X 0 80 r 0 0 2.2 B 70 70 0 0 shrubs CL 60 CML 60 E forbs E 50 0 s0sa 0 U 40 grasses 40 shrubs 30 30 ad2d 20 forbs 10 10 irassesbrasses 0 0 T augustau6ust october june august october june different fig 1 standing crop of grasses forbs and shrubs on vegetation types

01.01 and june and october on the creosotecreosotebushbush and tar- cantly P oi01 among seasons vegetation X date bush types types and the vegetation type interac- table 2 shows the vegetation type x date tion was also significant table 2 dropseed col- interaction was significant P 05os05.05 for several content of the diet was highest in pellets and lowest species this interaction indicates these species lected from the mixed shrub type did not constitute a similar percentage of the from those collected in the tarbush type in diet from june to october on the different veg- some types dropseed content of the diet was snake- etation types highest in june eg mesquite grass and black dietary content of dropdropseedsseeds varied signifisignify weed types while in others eg grama 19921 BLACK talledTAILED jackrabbitJACKRABBIT DIETS 303

creosotebushCreosotebush type 100 snakeweed loo loo100 type 90 90 so 2.2 70 X gmo CL 60 12 shrubs E E 0 so 50 forbs 40 40 grassa 30 shrubs 30 20 forbs 20 1 0 G 10 0 0 june august june october june august october

tarbush type mixed shrub grass loo100 loo100 90 90 0C 80 80 70 shrubs 70 60 forbs 60 E E X 0 50 grasses 0 0 40 40 30 0 shrubs 20 a 20 forbs 01 10 10 malMAI grasses 0 jujune 0 augustagustugust october june augustau ust october

mesquite grass type black grama 100 loo100 type 90- 90 80 0C 80 gogo70 70 60 021 60 E E 0 0 0 50 40 40 30 M shrubs 30 shrubs 0 20 PM forbs 01 we W 0 10 grasses 0 0 june august october june august october

2 fig dietary content of grasses forbs and shrubs in pellets collected from different vegetation types and tarbush it was highest in october types the difference among dates was relatively dropseed pellets content of collected from the small eg mixed shrub type table 1 these creosotecreosotebushbush type was consistent from june inconsistencies contributed to the significant through 1 october table vegetational type X date interaction FP oiol01.01 black grama content of pellets was not dif- table 2 ferent P iolo10.10 among vegetational types but dietary content of fluff9rassfluffgrassfluffgrass and threeawnthree awn was different dates io10.10 among P 10 table 2 in grasses was generally low table 1 however most cases black grama content of the diet fluff9rassfluffgrassfluffgrass contributed more than 22 ofthe diet peaked in august but for some vegetational in june on the black grama type and more than volume 52 oam BASIN ondCID grestGREAT 4 naturalist

1 111 t 10 i oi 00OO lo10 oi to 0 9.5 84 040.4 888.8 050.5 31931.9i 94 151.5 15615.6 18718.7 95mm OO mw 51951.9 00OO 05em 319i october OS 15lm 1cac 00 95OS 00 eh 519 ii 1 0 1 0 0 0 0 m co Su 0 0 rir i 3 0

3 OO CT 1 tarbush C i i 0 00 00 00 1.2 4.8 0.8 222.2 181.8 13.00 3 0.2 9.7 0.1 010.1 505.00 12 480 08 45945.9 zmi 130 e 02em mg97 01 CT39839.8 23823.8 me zmi hm 08em u bb82birbirkyirklq co august 02 97 el01 comrm 0el01 5010 lb 0 0 459 yir lre 2 s 0 oieole 0 CO ci010.1

CO CO 1 CO 00OO aticti aqcq c1ca i 1 1 fig 001 1.6 u 3.8 2.1 87 919.1 454.5 010.1 68 38 512 13213.2 zm16 258oemomm 29 mm38 61ciecle21 22222.2yi 00oo 91ml 45wm 01 co co i i co ler june 688 0 ii i ritokko710 cico cio0 1 l 0 0 m itooi

I-uco CO i 00 1.3 10 OS to t 00 5.8 141.4 0 777.7 mm13 25025.00 lo 9.6 58 f i 7.5 191.9 151.5 96 mm 14lw1 440 gi67t r 75 1 leq168 me 58 1 017.0 T 30.7 CT october 170 bm1 wo lm 307 96 10 lr lf 0 zoemoet 85 lo 15 307 D lo t ri 0.1 1 0 0 0 cl01 u 1 0CO r tf bush3 2 0 U omiOTI os os oo co QO t 1.0 creosotebush oi co 00 ii 2.8 2.4 111.1ii 100 040.4 wm494.9 1.8 111.1 28 24 1 46.7 2.0 494.9 18 10310.3 1 bwmh lmI 467 10 ew 49 go 0 zm1 august 152 21821.8 20 wm 47947.9 lw zem lm 88 M wei 04 S 1 20me 49 himwim r ler al hei 10 t 1 0 0 0 lo1 Creosote 0i 0 i N f a a U 00OO i 1 c1ca n I CO 10 CO 05 0 10 3.8 727.2 0 rir 030.3 292.9 060.6 656.5 mm38 35.1 124 31631.6 0 120012.0 171.7 21 mom 10510.5 em bm 650em rm 351 iblm R 15015.0 l c 326 zem 03 89 06 O 10 cl june 10 C llrzi C lee 0 0 CO 11 CO 1 rl 1 1 CO 0 0 ll i

1

1 D 1 CO OO 1 05 05 10 t 00 01OI M 4.9 0.6td 141.4 292.9 24.1 26326.3 g 1.2 3.7 025.0 wm49 06 mwi 01 bm 465hemCD 241 10710.7 98 mm12 37mg 26226.2 250 49 em lq 89 ber0 october rgCO 06 1 0 oi lb td bee80010 0 0 0 M 0 M 01 01 c1ca n3na 1uau 8

i 1 05 05 1 I CO 10IO snakeweed t- 1 l 10 t 0 1.3 8.1 f U S 0 767.6 090.9 10.9 11111.1 13 55.5 81qz i114 2.0 109 1 l 555 979.7 14214.2 36 020 335mrm 10710.7 im em lem109 or1 10 00 august CO bm CO tle 09 aln lo 1 so CI zei 1 0 97mi be lel l ca 05 0 0 01 r 10 c3 CO 0il u c3ca 1 typeammo 1 CO C 00 u c0ca 1 1 05 M 10 f 4.2 0 5.5 111.1 333.3 545.4 42 45845.8 13013.0 17117.1 i 9.1 0.1 11211.2 55mm 1 em hm CO lom 12312.3 91 12112.1 01 c369rem 1 55 lnml 33CO IOmw hb 10 leezee h-zol juneazzoc 91ml c1ca ez 10 5410 i 0 CI 05 hl 1 1 r i 11 0 co 0 ri ii feces 1 w 5 vegetationQ 11 llU OS CO 0 c1ca oolooi 1.6to 1.4 rabbit3 00 0 10 9.4f 0.60 1.2 01il0.1 010.1ii 51.9 16 14 12312.3 bo 06 12 13213.2 5191 16lm ahlw 94 1 l a 0 17.8 11.4 3003.0 26 mw ee zmI CO 01 Camr 178 1141 me 35635.6 94 06 lb 01 october 1 i I CO30 CO eme 05 al il 1 10 1 jackrabbit 0 lohaoh lo 1 0 0 1 1 0 0 10 y i i CO jack0 shruba w 3 0 XT ca i 1 05 1 10 10 10 10 c0 10 4.7t 8.9 lo10 t t lo 9.5 151.5 151.5 030.3 010.1 353.5mm 26626.6 47 QOmm89 16016.00 u 0.7 95 1 89 amm n mm 1 tailed 3 ami137 07 mom 15lm 15lm em 01 35 to august 24524.5 016116.1 zkiCO eg 575 95 03 el 00 0 1 so O 07 t 0 0 0 ca i mixed 1 c1 T 0 0 1 1 M il 10 i S 1 1 I CO nblack U oo roo oo 00 00 05 8.7 6.4f w 00 10 t- 3.0 0 87 64 17017.00 lo 2.8 5.1 19 300 23 ew 1 0.4 51 31031.0I 070.7 04 28 68 1 I me 1 64 11811.8 12512.5 5181 eq 88M 30CO 310 00 lremie juneaxxeaccoQ 264026.4f 1 ozl 07ag mh 5110ml N 0 O 0 CO 3 N i 0 10 0 inS i 5 CT i- co t co co co 01 io ci 2.3 co io 838.3 494.9 151.5 56.7 19219.2 mr23 25725.7 identified1 ii mm 12912.9 wm 1 567 102amm 545.4 050.5 01 17317.3 14214.2 49 15lm 567 05 bec0ca 10 october mw em ahm 00 0 cl t 1 5410 05 ziet 1 T 1 0 00 lo lir f 5 1 0 10 1 0 0 grass s 0 rl 1 0 i co 00oo co 00oo 00oo h 00oo 10 2.2 151.5io ci t ci 8.0 22 15215.2 I 18818.8 3 1.2 545.4 02.0 78 080 85 40840.8 15lm1 OO species 0 474.7 zm12 40.8 mw 20 me OO bb82 408 amb10 00 120 22222.2 1 408 8000 00 lo august OO CO miml zb hem 5410 c120ca t 1 u mesquite 00 0 0 i i CL 0 1 GO 0 V S il

S CO OO plant S CO CO CO 00OO 00OO b 00 CO c0ca 10 05 3.3 585.8 636.3 zeb102zem i S v 8.6 0.5 11.3 12312.3 mm33 52 mm 59759.7os mrem is 86 05em 29929.9 1130 cdrm 10 58 597 to & junec 20820.8 00me 05 89905 oi lo io 0 0 208808 1 0 0CO 0 0 M i 4 0 10 g i

i i important ii CO CO 4 t U 1 CO CO 10IO 05 10 0 10 tat0 05 CO 10 1.6 2.2 3.5 242.4 011 jaj3 0 9.1 0.5 161.6 zem 737.3 gm757.5 16 22bm 03 35mmCO OO48148.1 19919.9 bh84 23523.5CO 0 91 05 1 28.9 133 16lm bb 00 19905 M october 17617.6 lmze omm289 CO 65 CI 0 t 0591ml 05em 00 tirle t 0 t 1 M S r l 0 00 11 nj rl ll 1 I S of0 grarnagranna

i 00 10 00 05 & CO 10 00 0.3c0ca 2.905 1.800 8.6 2.8 10 2.1 0303.0 414.1 03 29me 18mmi 29829.8 me86 28 21921.9i august 9.5 85 21 clicitbm lq omm oo cobm mm95 27327.3 147847.8 blbaco reeeco hl 03 clilciti 05 00 9505 663863 00 0 co o i 1 oo 0 i 0 oi c black 1000 content S l 1 CO CO I1 c0ca 05 05 CD r t 10 tl CO t CO CO 4.3 6.9 161.6 111.1rl 373.7 0.3 0.1 69 1 mo lo 0.7 838.3 090 535.3 03 01 wp43 mm 56956.9 zm lm1 I COro S 959.5 64 22222.2 07 39639.6 mr 53mr erme 01 hr io69 memco le june 0595mm 0o ci 0ei07 05emm 00 05 10 0 0 f 10 1 3 00oo co botanical r 0 pod0 CQ S flower mallow g volmweed daleaealom graina araina globemallow 1 paperflower shrubs spectaclepod 1 1 grasses 1 s seeds nightshade1 1 grass 1 forbs quite a snakeweed featherweedleatherweedlay acl Spectacle silverleaf tj threeawns latcroton mesquitemcsquite a dropseedsi lamoreamorpaper fluffgrass scarlet globe totalforbs TABLEea S Leather dwarf wooly 1 icl 1 yucca shrubssbrubs grasses 11051 Drop black Mcs species0 ll051Fluff total forbs Total syshs D lill total illltj Ppll laleesj eiE i eiE i ci U ijriiih 199211992 BLACK talletalledTAILETAILEDD jackrabbitJACKRABBIT DIETS 305

fi S CS W 2 B a 2 0 2 2 1 3 2 0 CO 00 S i 10 05 CO 5 05 05 t S 1 i i il s 0 0 CO CO 00 Q 05 00 0 1 to 1 il S a

S aaiaciCTS S CO 2CO S S 1co tfaf2 10 2 2 p4pa I 3 1 s CO GO GO i f oi 00sas 00CO CO S 0o t- so a 1 S 00aqgq 1 D rlr l CO 10 a CO il i1ia 05 CO a 1 S S i 00co C 0i i 00oooq azzz0 2 3 S 6 Q t u3ua 10 s to10 01 l CO 1ly vi w 11 0 w 1 2 2 2 tl 0 0 1 2 & i 00 CO s u r f CO 8 CO rf if iu t SO 1 v 00 CD i 2 V 00oo co to E co 00oo c S 00 t0ta io co 3 OSScoC 12co 3 0 0 osbos t a V CO t 1 1 tt- co co 1 s ya 3 E 2 2 e W W S S 2CO S 2 2 2 2 to10 1 r l CO 0 r- oi co OS a 10 f 1 P rl a co tat0 11 XI CO 1 riq ll UVI CO 1 00 0 s I1 0 CO fl a S S 9 11 1 13 ii 3 0 0 CO clCO 1 t3ta 3 00 M CO a U I1 CO COC 10 I- 3 0 CO 05 CO 0 COC u 00 CO 1 s fr 0 ol01 i 11 1 11q 05 CO 10 00c6ca 00 P CO i 05 M 101.0cct CO i COcqaq tu 0 io0 S 00 IQ 1 s i s 0 a a s3sa 3 3 8 M ertcrt 2 00oo i & i s i 1 0 10 oio 1 0 1 1 i 0 5 fr S il cnCO a s it i 5 S U i w i ai 00 0 t1 u u f 0 2 tl ta co 10 u 00oo 10 i 00 t1 M CO 10 CO i 1 g S 10ID0 S S co ro 01 CO i 2 & s jl Q s r 1 r1ra 0 w 00 1 u s fi 0 a 1 CO 0 CO 5 X tat2 a s 0 or i cna 53 8 0 2 2 0 ii B 3 4 z z a M 0 a 2 a c 10 tat0 e i co Q OS y & 0 wetwee 0 1 to to10 05 0 eft to 11 H v t S i H a CO CO 05 8 CO CO C 3 ft r1ra 11 g s 2 0.0 Q e s s is a a CO 00 i 5 D c 1 a 10 co co 2 2 I1a S 1 0 0 l- S 2 2 oo t 01cs irliri i in 10o 00 co Q 0 i CO 3 CO 10 Q CO 0 tat0 il 11 05 u 1 ic S S C c3ca N lo10

1 4 OJ 3 lsl3s a ji y fi t to e S dl 03 t-toyroyos0 i U M q0qa 101.000oc0 y a y 3 8 il cl0 m 5 Z s 0 a S a s 1 W f 005 COC S u 00CK S t n 0 0 as 0 10 CO 00 y & 3 05 GO 05 S m ns C aczc0 i t CO s 4ai1i 01 1 co 00QO c 0 qqCO CO 0 23 0 3 1 10 00CO 3 s co clico co t 0 11 IQ 0 llQ M W ci S 11 2 2 2 s 00 2 1 1 1 05 005 1 i cl R 0 h0 S 10 00 r 10 t CB nt 0 tmM CO rt 0 0 CO S 0 co 0i 0co 00CO CO ttat2 1 0 s i t f 1 1 5 se S s rf 03 i 1 1 1 a 11 gw 1 0 0 S s SP i 0 5 y 0 h 3 J S U g 61 tat3 A g to10 CO to10 CO 2 in CO 3 s219 cU 8bc 0 ot & 0 1 & 0 C0 v f s 0 wa & Q Q fi Q 0 a a 0 1 3 2 3 ea t u U 0 & 5 u g hi CO 4 4 5 s S c S g 0 3 a 5 g i 43 TO e CO a oi x i L ai XV a x ia0 p n ri S e5ea W S U D D D 3 y S u lu & s & V V 0 0 m i 0 UGO 3 5 cicn Q E cn 306 GREAT BASIN naturalist volume 52

all black tailed jackrabbitjackrabbit diets tabletailleTABLF 3 month and vegetation type when preference index exceeded 202 0 for species in on six vegetational types

vegetational type grass tarbush species blackblaekblabiack grama mesquite grass mixed shrub snakeweed Creosotecreosotebushbush

di opseedlopseeddropseed june june oct aug oct black arainagrainagrama june aug oct flufffluffgrasspluff grass june oct oct june june june aberts buckwheat oct june aug oct aug snakeweed june desertdeseitbaileyabaileyabaileys oct desert holly aug june oct aug oct oct dwaldwarff dalea oct fendlerfendlersFendlerss bladderpod aug oct june june globeinallow aug oct aug june oct june aug ilyrnenopappushymenopappus june leatherweedfeatherweedLeatherweed croton june aug aug oct june aug june aug june rattlesnake weed june oct silverleaf nightshade june aug aug june aug oct june aug Spectaclespectaclepodpod oct oct wooly paperpaperdoweipaperflowerflower oct june oct june june mesquite june aug yucca june aug

12 in june and august on the mixed shrub of the diet on the black grama mesquite grass type and in june on the snakeweed type table and snakeweed types during most seasons 1 threeawns contributed less than 9 of the table 1 dietary content of other forbs was diet on all dates and vegetational types inconsistent among vegetational types and dates other grass species made small contribu- table 1 russian thistle salsola ibeibeticaibericaTicapica was tions to the diet plains bristlegrassbristle grass SesetariatatlataTia the only forb species with a nonsignificant P leucopilaleucopila vine mesquite panicum obtusum loio10.10 vegetational type x date interaction table 2 and burroburrograssgrass scleropogon brevifoliusbrevifolius did shrub contentoftbejackrabbitofthejackrabbit dietsdietswaswas also not differ in diets P loio10.10 among vegetational inconsistent among vegetational types and types or dates and the vegetational type X date dates mesquite contributed substantially to the interaction was not significant table 2 diets on most vegetational types between june forb content jackrabbitofjackrabbitof jaekjackrabbit diets varied over and october mesquite constituted more than time and vegetation type for example the con- 24 of the diets on the snakeweed type in tent leatherweedfeatherweedofofleatherweedleatherieatherweed croton differed significantly october but only 1 on the creosotecreosotebushbush type yucca yucca P oi010.001.011 among vegetation types and dates and in august table 1 elata contrib- the vegetational type X date interaction also was uted more than 7 of the diet from the significant P oiol01.01 table 2 its content varied creosotecreosotebushbush type in june but was not found in from about 24 in pellets collected during pellets collected from the snakeweed type on august in the tarbush type to none in the mixed any date table 1 however several shrubby shrub type at the same time Leatherleatherweedfeatherweedweed species did not show a significant P 1010.10 croton appeared to be an important component vegetational type x date interaction crucifixion 199219921 BLACK TAILED jackrabbitJACKRABBIT DIETS 307 thorn koeberlinia spinosa creosotecreosotebushbush zin- species in the diets although preference for nia zinnia aceosalareosaaceosaaceosalosa and ephedra ephedra sppappspp1 mesquite was not high other important shrubs varied considerably over time and space dietary preference the preference index was generally below 2 acknowledgments for most grass species table 3 however jack rabbits apparently preferred black grama on all this is journal article no 1560 of the new dates in the mesquite grass type fluffgrassFluffgrass was mexico agricultural experiment station las preferred during some months on all types cruces except for the mixed shrub grass type the preference index exceeded 2 for fluffgrassfluffgrass in literature CITED june on four of the vegetational types the preference index exceeded 2 for several arnolARNOIARNOLD D J F 1942 forage consumption and preference of forb species table 3 those with a preference experimentally fed arizona and antelope jackjackrabbitsrabbits university of arizona agricultural experiment station index 2 for six exceeding more than combina- bulletin 98 86 appp tions of vegetational type and dates included BEAR G D AND R M HANSEN 1966 food habits desert holly pereciaPepereziarezia nana fendler bladderpod growth and reproduction of white tailed jackrabbitsjackrabbits min lesquerella fendlerifendleri globemallowglobemallow sphaer- southern colorado colorado state university agricul- alcea leatherweedfeatherweedweed and silverleaf tural experiment station technical bulletin 90 59 appp sppapp leather croton CROCKER B H 1959 consumption of forage by black nightshade dwarf dalea dalea nana was pre- tailed jackrabbitsjackrabbits on salt desert ranges ofnew mexico ferred only in october in the black grama type journal of wildlife management 30 304 31311L dabo et al 1982 found dalea was highly pre- DABO S M R D PIEPER R FK BECK AND G M SOUTH WARD 1982 summer and fall diets of black tailed ferred and comprised as much as 65 of the jackrabbitsjackrabbits on semidesertsermdesertdesentdesenn rangeland new mexico diets in the fall on grassland vegetational types state university agricultural experiment station mesquite and yucca showed a preference index research report 476 above 2 for june and august on three vegeta- fagerstone K A G K LAVOIE AND R E GRIFFITH 1980 blaekblack tailedtalled jackrabbitjackrabbit diet andlindaind 3 JR density on tional types table rangeland near agricultural crops journal of range management 33 229 232 FATFHIFATEHI M R D RFPERreperPIEPER AND R F BECK 1988 seasonal discussion food habits of black tailed jackrabbitsjackrabbits lepus californi cus in southern new mexico southwestern naturalist black tailed jackrabbitsjackrabbits in southern new 3336733 367 37070 mexico appear to be opportunistic feeders FLINDERS J TTANDRAND R M HANSEN 1972 diets and hab- itats ofjackrabbitsjackrabbits in northeastern colorado colorado although this study and earlier ones indicate state university range science department science that as many as 30 plant species can be found in series no 12 fort collins colorado 29 appp fecal samples at any one time 5 or 6 species HANSEN R mandlMANDJM AND J T FLINDERS 1969 food habits of generally made up the bulk of the diet forbs north american hares colorado state university range science department science series no 1 fort often contribute a greater proportion of the diet collascollms colorado 18 appp than grasses but the important forb species vary HOLECHECK J L 1982 sample preparation techniques for considerably among locations seasons and mieromicromicrohistologicalmicrobistologicalhistological analysis journal of range manage- years leatherweedleatberweedfeatherweedLeatherweed croton is perhaps the main- ment 35 267 268 HOLECHEK J L AND B D GROSS 1982 evaluation of stay of the diet among the forbs although sev- different calculation procedures foifolfor micromicrohistologicalhistological eral others such as silverleaf nightshade and analysis journal of range management 35 721 723 wooly paperpaperflowerflower psilostrophe tagetinaetagetinae HOLECHEK J Llanamlandmand M VAVRA 1981 the effect of slide contribute substantial amounts to the diets and frequency ofobservation numbers on the precision of micromicrohistologicalhistological analysis journal of range man- dropseed black grama and fluffgrassfluffgrass agement 34 337338337 338 appear to be the major grass species contrary HOLECHEK J L M VAVRA AND R D PIEPERPIFPER 1982 to cattle which utilize black grama mainly botanical composition determination of range herbi- during the dormant season et al 1975 vore diets a review journal ofrange management 35 rosiere 309 315 rodriguez et Aal 1978 jackrabbitsjackrabbits apparently HUMPHREY R R 1958 the desert grassland botanical consume more black grama during the summer review 24 1 64 growing season consequently high jackjackrabbitrabbit JOHNSON R D AND J E ANDERSON 1984 diets of densities could reduce the amount of black black tailed jackrabbitsjackrabbits in relation to population den- sity and vegetation journal of range management 37 79 grama available for cattle later in the year KRUEGER W C 1972 evaluating animal forage prefer mesquite appears to be the main shrubby ence journal of range management 25 471 475 308 GREAT BASIN naturalist volume 52 lechleitnetiLE IILEITNER R R 1958 certain aspects of behavior of SAS 1985 SAS useruserss guide SAS institute inc cary north the black tailed jackjackrabbitsrabbits american midland natu carolina ralistcalistiahst6060 145 155 SIDAHMED A E S C DENHAM J G MORRIS L J KOONG AND S R BADO SEVICH 1981 precision of ludwicLUDWIG J A JJ F REYNOLDS AND P D WIIITSONWHITSON 1975 radoradosevich siesize biorbiomassnass relationships of several Chibuchihuahuanchibuabuanabuan desert micromicrohistologicalhistological estimates of goats from fecal analysis shrubs american midland naturalist 94 451461451 461 and cell wall passage through the gastrointestinal tract proceedings of the western section of the american mcadooM ADOO J K AND J A YOUNG 1980 Jackjackrabbitsrabbits of animal 32 201 204 rangelandsrangelands22 135 138 society science SPARKS D R 1968 of black tailed jackjackrabbitsrabbits on PAULSENPAULSLN H A JR ANANDFD F N ARES 1962 crazinggrazing values diet albAJR sandsandhillbill rangeland in colorado journal ofrange man- and management of black grama and tobosa grasslands 21 203 208 and associated shrub ranges of the southwest USU S agement URESKUKESK D W 1978 diets of black tailed hare in steppe department of agriculture forest service technical vegetation journal of range management 30 439 12701970 56 bulletin 127056ppappp 442 R D AND C H HERBELHCBBEL 1982 herbage dynam- PIIPIR DANDC hebbel VALENTINE K A 1970 influence of grazing intensity on and productivity ofdesert grassland ecosys- ics primary improvement of deteriorated black grama range new new mexico state agricultural tems university mexico state university agricultural experiment sta- 695 43 experiment station bulletin appp tion bulletin 553 BEIGCIREIGEL A 1942 somesorne observations of food locations of WEWESTOBYSTOBY M G R ROSTANDROST A ND J A WEwelsWEISI1 s 1976 problems rabbits in western kansas during periods of stress with estimating herbivore diets by microscopically transactions of the kansas academy of science 45 identifying plant fragments from stomachs journal of 369 375 mammalogy57167mammalogy 5716757 167 172 PIEPERPIFPER S SMITH 1978 RODRIGUERODRIGUEZ J GGRR D AND G WOOD J E 1969 rodent populations and their impact on botanical colcoieolcompositionriposition of cattle diets on desert grass- rangelands new mexico state university Aagricultural land range new mexico state university agricultural experiment station bulletin 555 experimentexpel imentament station research report 363 4 appp ROSIERE R E R F BECK AND J D WALLACE 1975 cattle diets on semidesert grassland botanical compo- received 9 april 1991 sition journal of range management 28 89 93 accepted 15 july 1992 great basin Natnaturalistmalist 524 appp 309 312

SPECIES OF EIMERIA FROM THE THIRTEEN LINED GROUND SQUIRREL spermophilus tridecemlineatus FROM WYOMING

12 1 1 1 robert S seville diane M thomas russell pickeringPickermg and nancy L stanton

abstbacABSTRACTr five species of the coccidian genus eirneriaedmenaetmena E beecheyibeeuieyzbeecheyi prevalencepievalence 179 E t dlofpermophihcallospermophili mominensismorainensis 2861286128.6128 6 E laniwrefisislarinwrensis igi16116 landande bilamellatabilamellatebtlamellata36wereieovcredhoms613361 were recovered frornbrorn 5613 lined ground squirrelssqinii elseis spermophilus tridiceinlineatustrtchceinltneatus collected fromhiom two sites in eastereasternn wyoming two squirrels from one site were also passing an unidentified polypolyspolyspoipolysporocysticsporocysticpoipor ocysticoocystic coccidian infected squirrels were found to harbor from one to threethiee species simultaneously previously these samesarnegame eimenaneirnerian species weiewelewere found infecting sympatric populations of wyomingWyorning ground squirrels sper nwphtlwmophilitsmoph littsilitslitls eleganseldelegansegans and white tailed pianplanprairieie dogs cynomys lencunisienleulenleucurtisneucurtiscuniscurtis at one of the sites it is suggested that the exchange of these generalist paiaparasiteparasite species among co occuiocchioccurringangnng selundsciurid hosts contributes to the consistent prevalence levels reported in wyoming ground squirrels

kenkeykegnenneu morismoniswordsmonds eimeriaelmenaelmeria spermophilus tndecemlineatustridecernlineatus prevalence polyporocijittcpolysporocysticpolysporocystic coccidia

shults et al 1990 reported the occurrence allowed parasite exchange among related hosts of six species of merianeimerianeieimerman parasites protozoa one important factor in maintaining the stabil- apicomplexa in sympatric populations ofwyo- ity ofofeimerianeimerianassemblagesassemblages is exchange of par- ming ground squirrels spermophilus elesanselegantelegans asite species among closely related sympatric elegansdelegans kennicott 1863 and white tailed prai- host species rie dogs cynomys leucums merriam 1890 the purpose of this study was to determine from wyoming stanton et al 1992 conducted which eimerian species are present in wild pop- a study of eimerian species in four wyoming ulatulationsions of 13 lined ground squirrels sper- ground squirrel populations and found that nwmomophilusphilus tridecemlineatus mitchill 1821 and most infected ground squirrels harbored two or to assess the role these hosts play in mainte- stable observed more species and that the eimerian assemblage nance of the eimerian guild in was present across populations and over years wyoming ground squirrels toft 1986 recognized two classes of para- sites micro and macroparasitesmacroparasites macroparamacropana METHODS sites eg helminthshelminthes tend to prodproduceilceileeLice long lasting infections and are endendemicernicernieernle in host in 1991 we sampled 13 lined ground squir- populations while microparasitesmicroparasites protozoa rels from two locations 1 a native short grass bacteria viruses produce short lived infections prairieprairiehayfieldhayfield 10 linkinkm south of laramie wyo- and long lasting immunity resulting in oscilla- ming 4ri2n41oi2n 10533w and 2 a native tions of infection frequency epidemics within short grass prairieprairiehayfieldhayfield 18 kmkinkiuklu south of gil- the host population the stability for intestinal lette wyoming 4414417nTN 105310531wIW protozoaprotozoansns reported by stanton et al 1992 at the laramie site squirrels were live does not support tofttohttommis s prediction regarding trapped using national live traps once a month microparasitesmicroparasites while there have been no from july to september over the four day trap- mechanisms proposed for maintaining stability ping period squirrels were trapped using three in microparasite communities stock and 60 X 42 m trapping grids with traps set every 6 holmes 1987 proposed that species richness in 162 total traps traps were set at 2000 hr of intestinal helminth communities of brebesgrebes and checked each morning by 0800 hr was enhanced by reduced host specificity which at the gillette site six 400 m transects and

department ofzoology and physiology bofbox 3166 university ofwyoming iaaarnielailal amieamlejmie wyoming USA 82071 2 21resent21fixpitpixresentsenisent addressiddiaddi ess ontario ministry of natural resoincesResoincesces wildlife branch box 5000 maple diitaiioontario LAA isais9 canada

309 310 GREAT BASIN naturalist volume 52

tabletailleTABLF 1 total percent infectedandinfected and preprevalencesvalences hosts infected with given speciesspecieshostshosts examined ofofeimenaneimerianmerlan species in gigroundoundaund dwelling sciuridscnindschind hosts at laramieLararniearnle and gillette collection sites in wyoming inf total percent infected with Eftebbeimenaeftneriaedmenanerianeyfa elbeeibe E beecheyibeecheyi eibi E bilamellatebilamellata elca mo E callospernwphilicallospermophili morainensis ellaeilaelia E larimelanmerensislarimerensisrensis and eispbisp E sperinophthsperinvphili sciurid host inf elbeeibe eibi elcaelea nomoi ellaeilaelia eispbisp spennophibisspennophilu9 tndecemlmeatustrideceinfineatus laramie n 41 43943 9 73 24 96 14614.614 6 0 gillette n 15 86786 7 46746.746 7 67 80080.080 0 20020.020 0 0 total n 56 518 179 3616 286 161 0 S elelegantegans 1 laramie n 1007 68068 0 34034 0 110 4304343.00 17017.017 0 50 1 Cynorcynomysnys leucurusleucurus laramie n 18 94094 0 83083 0 1701717.00 22022.022 0 0 0

Pementpetuntigcspementapsadsaps for01 S tkgansdegatogans andcdcmdcC leuclencfuturesurus determined by takinghighesttaking highest of two values for PE calhspennophihcallospannophili ore01 F morainensiamoralnmorainawraffiensisensiAengla fromfioinsluiltsctshaltsults t alil ign1990

with an olympus CH microscope identifica- tion to species in most cases could be accom- plished based on oocyst size and external and internal morphology however for eimeria callo spermospermophilispemwphilispermophilephili henry 1932 and E morainenwrainenesismorainenesisnesis 0 torbett et al 1982 the respective size ranges overlap making identification dependent on internal morphology unfortunately rarely do all oocystisoocysts in a fecal sample sporulate therefore although both species were identified the two are ccombined into a single species complex E callospennophilicallospermophili nwrainensismorainensis comparisons of total percent infected and preprevalencesvalences of each species between the two sites were made using chi square tests number cruncher statisticalStatisticA system version 5035.03 hintze 1990

fig 1 photomicrograph of a polypolysporocysticpoly&porocysticsporocystic coccidian collected from a 13 lined ground squirrel 1250x Nonomarskydomarskymarsky RESULTS interference showing typical sporocyst with residuum clearly visible forty one 13 lined ground squirrels were sampled at the laramie site and 15 at the gil- six 50 m transects in various vegetation types lette site five species of eimeria were found were trapped during the second week in august infecting squirrels in both populations overall 3870 trap nights stations were 15 apart in 518sis51.8 of all squirrels examined were infected each of one victor rat two victor consisting trap with at least one species of eimeria the total mouse traps and one sherman live trap sher- percent infected was significantly higher at the closed man traps remained during daylight gillette 86786786.7 than at the laramie site hours and traps were checked and reset at dawn 43943943.9 P 05os05.05 infected squirrels at gillette and dusk also had higher parasite species richness 1771771.77 all fecal samples collected from animals at speciesspeciesinfectedinfected squirrel than at laramie both sites were placed in 2 potassium dichro- 1171171.17 total percent infected and preprevalencesvalences mate solution at room temperature 25 QC for at by species at each site are presented in table 1 least three weeks to allow oocyst sporulation for overall the eimeria callospermophili species identification oocystisoocysts were isolated by nwrainensismorainensis complex was the most prevalent flotation in saturated sucrose solution specific species found infecting 28628.6 of the 56 hosts gravity 12121.2 and identified at looxIOOX objective examined significantly more hosts were 199211992 elmeriaEIMERIA FROM 13 LINED GROUND SQUIRRELS 311 infected with this species complex at the gillette spurious infections ofinvertebrate origin based than the laramie site 80 vs 969.6gg P 05os05.05 on these reports it is likely that the poly eimeriaelmeria beecheyibeecheyi henry 1932 was the sporocystic coccidian observed in 13 lined second most prevalent species found infecting squirrels is a member of the genus bossiaklossiamossia and 17917.9 ofthe hostsosts examined significantly more possibly of invertebrate origin however iden- hosts were infected at the gillette site 46746746.7 tification to species requires further work vs 737.3 P 05os05.05 including the identification of the primary host eimeria larimerensislatinwrensislatinwlarime rensis vetterling 1964 was the results of this study indicate that while found infecting 16116.1igi of the squirrels exam- the eimeelmeeimerianmernaniianlianllan fauna of 13 lined ground squirrels ined prevalence was higher at the gillette site is very similar to that ofwyoming ground squir- 20 vs 14614.6 but the difference was not rels and white tailed prairie dogs at the lara- significant P 0505.05 mie site there were some differences in the eimeria bilamellatabilamellate was the least common preprevalencesvalences of the different parasites of the species found during the study 36363.6 again five species found infecting 13 lined squirrels prevalence was higher at the gillette site 67gt676.7 all have been reported previously from sympat- vs 242.4 but the difference was not significant ric ground squirrels shults et al 1990 stanton P 05os05.05 et al 1992 and all have been reported from two squirrels at the laramie site were also white tailed prairie dogs in wyoming todd and infected with a subspherical polysporocysticpolysporocystic hammond 1968a 1968b todd et al 1968 coccidian fig 1 with 10 12 sporocysts the shults et al 1990 however at the laramie site number sporozoitesofofsporozoites could not be determined 13 lined squirrels were not as frequently due to the large amount of residuum present in infected and had lower preprevalencesvalences than wyo- the sporocysts mean size for 15 measured ming ground squirrels for all species and lower oor oocystisoocysts was 386238.62 X 302030.20 JL sporocysts were preprevalencesvalences than white tailed prairie dogs forefor E spherical and measured 106510.65 X 1065logs10.65 JLL n beecheyibeecheyi E callospennophilicallospermophili morainensisnwrainensis and 15 and had no steidasheida body both oocystisoocysts and E bilamellatabilamellatebilanwllata values for 13 lined squirrels at sporocysts contained numerous residual bodies the gillette site where no other species of attempts to infect two captive wyoming ground sciurids were present were more similar to squirrels Spennspermophilusophilus eleganseldelegansegans were unsuc- those for wyoming ground squirrels at the lar- cesscessfulful amie site table 1 additionally wyoming ground squirrels hadbad greater species richness than 13 lined squirrels stanton et al 1992 discussion species richness for prairie dogs has not been reported E beeche E bila the occurrence of beecheyiyi results indicate that related sympatric hosts fellatamellatanwllatamenwllata and E morainensis in 13 lined ground can be infected by the same species ofeimeria squirrels constitutes new host records for these which may contribute to the stability of the species in this host polysporocysticPolysporocystic oocystisoocysts have eimerian guild not been previously reported from sciurid rodents levine et al 1955 identified two poly acknowledgments sporocystic species bossiaklossiamossia perplexperplexensens from deer mice peromyscus maniculatus and K this research was supported in part by the variavariabilisbilis from the western big eared bat department of zoology and physiology and the corynorhinus rafmesquii collected at the rafinesciuii office of research university of and grand canyon all wyoming arizona because species of NSF grant BSR 8909887 bossiaklossiamossia previously described were found in invertebrates levine et al 1955 postulated that the two species were parasites of inverte- literature CITED brates eaten by the deer mouse and bat domeydorney DORNEY R S 1965 elmeriaeimeria tuscarorensisttiscarorensts n sp protozoa 1965 reported finding two polysporocysticpolysporocystic elmenEimeneimenidaeeimeriidaceumenidaeidae and rcdescriptions of other coccidia of the oocystisoocysts in feces from a woodchuck marmota woodchuck marnitamarnwta monax journal of protozoology monax from pennsylvania that resembled the 1242312 42342649342642312423426426 descriptions of the two species in the genus LEVINE N DVIVENSD V ivensIVFNS AND F J kruidenier 1955 two new species of bossiamossiaklossia sporozoa adeleidae from a bossiamossiaklossia reported by levine et al 1955 dorney deer mouse and a bat journal of parasitology 41 speculated that the two oocystisoocysts might represent 623 629 52 312 GREAT BASIN naturalist volume

ground squirrel Spennspermophilusophilus shullssiiurrs L M R S SEVILLE N L SIANIONSTANTON AND G E 1932 from the uinta MR journal of protozoology 15 1 8 MENCKENSmr N KI NS JR111 1990 eltEiTeirnenaeitnerialeitnerianerianerla sp apicomplexa armalusannatusarmatus lii of Ennenennenidaeeimeriidaeenneoidae from wyoming aroundground squirrels sper 1968b life cycle and host specificity eimenaeinwriaedmena ayn vetterlingVetterlmg 1964 from the uinta ground inophilus eleganelegant3 and whitwhite diledtailedtalled prairie dogs cinoincyn larimerensislarime rensis journal of protozool- omy leucunisleucurnsleucuniscurns in wyoming great basin naturalist 50 squirrel Spennspermophilusophilus areatusarnetusarmatusarnwtus 327 331 ogy 15 268 275 S HAMMOND AND L C ANDERSONANDFRSON SIANIONSTANTON N L L M SIIULIS M pankerPARKERPARKFR AND R TODD K SJRS jil DMD M LL life of eirneriaedmena SEVILLEsrvnarvn LF 1992 coccidian assemblages in the wyo- 1968 observations of the cycle eimena 1932 in the uinta ground squirrel rninggroundming gi oundaund squirrel spermophilus elegansdelegans jourjournalnalofof bilamellatebilamellata henry in parasitologyPariparlsitology 7832378 323 328 spermophilus armalusarmatusannatus journal of protozoology 15 rela- slockSIOCKSTOCK T M AND J C HOLMFSHOLMES 1988 functional 732 740 with tiontionshipsships and micromicrohabitathabitat distributions of enteric hel tonTOFT C A 1986 communities of species parasitic toh case minthsmintasrninths of brebesgrebes podicipedidae the evidence for lifestyleslife styles pages 445 463 in J diamond and T J new york intelinteractiveactive communities journal of parasitology 74 eds community ecology harper and row 214 227 TODDTOOD K SSJRJR AND D M HAMMOND 1968a life cycle and host specificity of elmenaeimeriaelmeria callospermophilicalloipemwphili henry received 12 february 1992 accepted 18 september 1992 great basin naturalist 524 appp 313313320320390

PLANT AGESIZE distributions IN BLACK SAGEBRUSH ARTEMISIA NOVA EFFECTS ON COMMUNITY STRUCTURE

james A youngyouner I1 and debra E palmquist I1 abstractabstracrabsiracr the demographydernography of black sagebrush artemisia nova nelson was investigated in the buckskin mountains ofwestern nevada to determine patterns of stand renewal in sagebrush communities currently free from wildfiiwildwildfiresfireses biomass sampling was conducted to develop growth classes that reflected apparent age of the shrubs the density of black sagebrush plants was twice that of basin blibizbig sagebrush A tntridentata sspasp dentatatntritridentatatndentatatridentate nutt in adjacent communities on contrasting 2 soils 22222.22 2 versus 111 1 plants per rm black sagebrush accumulated only 75 as much woody biomass as big sagebrush regression equations were developed and tested foiroiforbol predicting total woody biomass current annual growth CAG and leaf weight of black sagebrush plants apparent age classes were developed both for the black sagebrush plants and the sub canopy mounds on which they grew discriminant analysis was used to test this classification system plant succession apparently controlled by nitrate content of the surface soil appeared to eliminate the successful establishment of black sagebrush seedlings on the mounds after the shrubs die the mounds eventually deflate we propose that mounds reform around shrub seedlings but because seedling establishment is so rare in these communities this could not be verified

key words bionblonbiomasslasstass shrub succession desert soil formationsoilfonnation soil nitrate black sagebrush artemisia nova

black sagebrush ATteattenisiaattemisiaartemisiamisiamisla nova nelson is west central nevada is characterized by black one of the dwarf sagebrush species which col- sagebrushdesertsagebrush desert needlegrassneedle grass stipa speciosaspeciosospeciosa lectively constitute about half the sagebrush trin & duprrupr plant communities the buck- vegetation in nevada beetle 1960 black sage- skin mountains are located 100 km southeast of brush plays a dominant role in a number ofplantofplantpiant reno nevada in the rain shadow of both the communities in the great basin zamora and sierra nevada and pinenut mountains this is a tueller 1973 rarely does black sagebrush portion of the carson desert in which billings share dominance with another species of arte- 1945 suggested that atriplex dominated salt misia in the section tridentate of the genus desert shrub vegetation occurred because of algeaigeATteattenisiaattemisiaartemisiamisiamisla black sagebrush is perhaps the spe- atmospheric drought rather than occurrence of cies most adapted to andaridanidannd environments black soluble salts in the soil ifwe compare the black sagebrush is closely associated with shadshadscalescale sagebrush communities of the buckskin moun- aftlplexatriplex confertifoliaconferticonfiettifoliafoliapolia torr & frem wats tains with those described in the regional study dominated landscapes blaisdell and hohngrenbohngrenholmgrenHohngren conducted by zamora and tueller 1973 we 1984 the browse of black sagebrush is highly find that the highest elevation north facing preferred by domestic sheep ovis atiesarlesaries slope communities of the buckskin mountains pronghompronghom antilocarpa americana and sage correspond to the most andaridarndannd communities pre- grouse Centrocentrocereuscereus phasianusorophasianusoro from the viously described from this we assume the 1890s until the late 1950s black sagebrush plant black sagebrush communities in this study rep- communities in the carson desert of nevada resent an andaridanidannd extension of this type were a vital part ofwinter range for the domestic only recently have occasional wildfireswildfires of range sheep industry years of excessive brows- any extent occurred in black sagebrush commu- ing by sheep actually shaped the outline ofblack nities in western nevada the fires that have sagebrush shrub canopiescanopies zamora and tuellertheller occurred have been associated with the recent 1973 reported they had difficulty in finding spread of the alien annual cheatcheatgrassgrass bromus relic communities in high range condition tectoriumtectorumtectorum L into these arid environments vegetation of the buckskin mountains of young and tipton 1990 apparently for much

usdaUSDAUSUA agtilultmdlreseuchseniceagriculturalagficultural besereseareliResearcilareli service 920vdlle920990 valleyvailey road benorenobe no levadinevadinevada 89512

313 314 GREAT BASIN naturalist volume 52 of the twentieth century these communities located along the western flank of the buckskin have not been subject to wildfireswildfires because of mountains the five stands located on the same lack of herbaceous vegetation to carry the fire croppingoutcroppingout of mickey pass tuff were separated because ofthe lack oftrees to produce fire scars by small canyons where the westerly tilted ash it is difficult to determine whether these sites flows were broken by faulting all sites were were subject to periodic burning under pristine west facing and located in a band along the conditions this is in sharp contrast to basin big mountainside at 1720 1780 inm elevation sagebrush communities where periodic cata- A starting point was located on aerial photo- strophic stand renewal by burning from wild graphs in each stand and 10 plots each 10 m2ma fires has been common the lack ofcatastrophic in area were located randomly along line tran- stand renewal in black sagebrush communities sects parallel to the slope A total of 50 plots should be reflected in the agesizeage size class struc- were established 5 stands X 10 plots per stand ture of the communities in each plot the following were determined a our purpose was to determine the agesizeage size shrub density by species b crown cover of distribution of black sagebrush plants to deter- shrubs ocular estimate c shrub height d mine community structure area of mound and interspacesinterspaces and e herba- ceous cover ocular estimate mound cover MATERIALS AND METHODS refers to the slightly raised areas beneath shrub canopiescanopies where subaerially deposited soil and studies were conducted from 1984 through saltation deposits accumulate buckskin located about 1988 in the mountains at each plot location the herbaceous vegeta- 100 kmkin southeast of reno nevada geo- the tion frequencyfrequencywas was sampled with 100 step points of this mountain range have been logic features arranged in 4 lines of 25 points each following and orielonnel 1979 described in detail by hudson the procedures of evans and love 1957 the and soils of the range have been vegetation herbaceous vegetation was resampled annually mapped and related to the geologic map of the the same starting point but by placing area lugaski and young 1988 the plant com- using the transects and down the slope 25 black munitiesmunities used in this study were located on the up brush mounds were located in each stand guild mine member of the mickey pass tuff saliesatiesalzesatiebrushsagebrush shrubs rooted on each mound were mea- this geologic unit consists of crystal rich mod- the sured for a height b maximum and mini- erately to poorly welded ash flow tuff proffett diameter c stem number as black and proffett 1976 it has been proposed that mum crown sagebrush ages the cambium splits forming the soils a developed in place b developed stemmed plants and d stem diame- from subaerially deposited material from long multiple at soil surface diameter of the group of distance or c developed from a ter the transportation aerial of the plant was combination of residual and subaerially depos- split stems the portion subdivided clipping into the following cate- ited material unpublished research ARS- by a coarse stems 252.595gs cm or larger in USDA bulk of the profile is an argillic gametergorieslieslles the Tatameter b fine stems 0250.25 to 2.424 cm in diam- horizon about 50 cm thick which consists of diametermeter 24 c current annual growth and d leaves 50 or more clay textured material it is pro- eter material was dried at 80 degrees C for 24 posed that this clay horizon is a relic of a soil that the hours and weighed developed on the site and whose original surface shrub was horizon has been removed by erosion the after the aerial portion of the the was col- important point is that the clay horizon which removed the litter beneath canopy 2 is intermittently exposed on the soil surface lected and screened through a mm screen developed under different environmental con- the material too coarse to pass through the ditions from the current surface horizon the screen was saved dried and weighed the max- the mound current surface soil consists of a relatively imum and minimum diameters of mound recently deposited layer apparently from sub- were measured and the height of the horizon aerial deposition that is largely confined to was determined by digging to the clay rooted on the mini mounds beneath the canopiescanopies of the black the number of perennial grasses of sagebrush plants soil is classified as a fine mound was counted by species and the cover the mound iridic montmorillonitic typic paleargidPaleargid cheatgrasscheatgrass was estimated ocularly per spatial structure ofthe black sagebrush com- A series of agesizeage size classes was established munitiesmunities was determined by sampling five stands for the black sagebrush plants sampled these 1992 19921 BLACK SAGEBRUSH demography 315

TABLE 1 mean plus standard for 2 2 error SE shrub density per m percent projected canopy cover frequency lom10 m plot withinmathin stands N 10 and constancy among stands N 5

species density SE cover SE 2 frequencyFiequency SE constancy SE maMm2

ATtenartemisialisiatisia nova 22 040 22 24 100 0 100 0 chrysothamnus viscidi viscidiflorusflorus 07 010 2 04 40 8 80 8 ephreda nevadensis ethreda 03 008 1 04 64 10 100 0 tetradymia glabrataglabrategla tla brata 02 004 32 5 60 4 eriogonum all11 microthecum 01 0040 04 5 1 20 2 eriogonum umbellatumumbellatum a 02 000 8 1 20 1

indicates less than 1 average cosercovercoveneoven

classes were based on the size growth form RESULTS AND discussion percentage dead canopy and apparent age of the plants the classes were a seedling b community competition young plant c mature plant d patriarch e senescent and f dead the plant communities of the buckskin mountains dominated by black soil samples from the surface 5 cm were sagebrush are low in diversity table 1 green rabbitbrushrabbitbrusbrabbitbrush taken a next to the shrub stem b at the chrysothamnus viscidiflorusviscidiflorus hook nutt canopy edge and c 10 cm beyond the edge of occurs in patches in the community nevada the shrub canopy these samples were dried ephedra ephedra nevadensis wats is rather screened and shipped to a commercial labora- evenly distributed through the black sagebrush for tory nitrate nitrogen analysis communities but at a low density littleleaf A two way analysis of variance and post hoc horsebrushhorsebrush tetradymia glabrataglabrateglabrata gray is a rel- duneanduncanduncanss multiple range test were performed atively infrequent component of the communi- to analyze differences in soil nitrate concentra- ties the two species of eriogonum are tion between sagebrush agesizeage size classes and semiwoody species that also occurred in the sample location A series ofofstepwisestepwise regressions most andaridarndannd black sagebrush communities that was performed utilizing the general linear zamora and tueller 1973 reported model wherein a subset ofvariables was chosen squirreltail elymus hystrix scribn and that would best predict plant weight annual cheatgrasscheatgrass are the most frequent herbaceous growth of plant weight and leaf weight of species table 2 the relative frequency of the two black sagebrush separate step up regressions species reverses from year to year depend- were performed for plant and mound character- ing on available moisture for plant growth Cheat istics neter and wasserman 1974 the inde- cheatgrassgrass is abundant only in years with ade- quate moisture during pendent variables that were significant the spring the density squirreltailofsquirreltailof plants remains contributors to predicting agesizeage size classes for relatively constant in dry years squirreltail is virtually the these two groups were chosen as discriminant only herbaceous species in these communities variables to be used in two separate discriminant analyses the agesizeage size classes of the sagebrush biomass plants were used as the grouping structure in along the western the discriminant analysis the plant character- margin of the buckskin mountains black and basin istic variables selected as significant contribu- big sagebrush com- munitiesmunities occur side by side on sharply tors to classification into agesizeage size classes were contrast- ing soils the basin big sagebrush communities a weight ofcoarse stems b number of stems have been burned in wildfireswildfires based on historic c plant height and d plant diameter the records and fire scar analysis young et al 1989 mound characteristics ranked in order of our analysis of black sagebrush communities is importance used were a litter cover b litter essentially based on aboveground woody mate- weight c soil nitrate concentration and d rial accumulation of the dominant shrub we cheat cheatgrassgrass cover on the mound had previously conducted a study ofthe biomass volume 52 316 GREAT BASIN naturalist

of herbaceous for an average of four years sampling TABI 1 2 mean plus standardstan daiddard error SE for frequency species 1989 no april precipitation and a year with above average moisture averageaveldeeelageeidge precipitation 175 mm for a dry spring available foifor plant growth 1986 225 mm precipitation based on 5000 sample points per year

frequencycy

average dry 198919 Q wetweilweijwetj986gsg986.9861986

four years SE spring SE spring SE

g11ownGKOWIIIII1 1ilomfilomkomsllkomolls11cil11II IsS piPERENNIALkl NNIAI nasscrassHASSGRASS 70 6 05 elymus hystrixlntrilutrilutfilnlantri tri 39 41 69 8 stipaspedosastipestipa ipeciovi 3 03 07 stipastipd tliurhenanathurberthurberianathurbenianalana poa secunda 1 08 onzojms0qyzopsis hyiracnoideshyirmnoicles 1 02

annualgrassANNUAIANNUAL crassGRASS 14 76 38 broniusbioniusbroiratsBroi ratsruts tectorunitectoriontestortectortectorionuniunt 44 66 28

PERENNIALpi HI NNIAI 1 OKB castilleja chroncliroiiwachrontosatosa 1 02 1 02 phaemlleasphaerllceasphaeralcea parvifohaparvifoliaparvifolia 2 03 phlox hoodiihoodei 3 03

ANNUAIANNUAL IOKBFORB 2 08 erodiumodlumodium cicittanuitnrihmcicittaritonriton 5 08 li 5 descurainia pinnatapignatapnmata 07 deuiminia 10 sisymbrium iltiswnumaltissimuin 5 09 08

lndiandi t s than 1 arawa raw off basin big sagebrush adjacent to the western wide difference between the two communities edge of the buckskin mountains young et al is apparently due to the higher woody biomass 1989 this allowed comparison of the produc- of more mature basin big sagebrush plants tion of biomass of basin and black sagebrush woody biomass of black sagebrush was best from the same area the basin big sagebrush predicted by the equation community had a sandy loam surface soil and a a Y 9879.87 1211.21 XI 11121.1212 x2xa 088oss0.88 x3xa greater soil depth haplargids derived from meta volcanic sources big sagebrush ages were where Y total woody biomass grams XI clumped at 55 60 4cm540 45 and 10 15 years old fine stems x2xa coarse stems and x3xa root 2 the general aspect of the two communities crown R 96gg96.96 for this determination yearly t is strikingly different with the maximum heighelgheight growth increment was predicted by the equation of the black sagebrush being 60 cm and that of 026 0.16olg t xa the big sagebrush over I1 m in contrast to the Y 169616.96 0260.26 XI 016 x2 central woody stems ofthe big sagebrush plants 0730.73 xax3 0140.14014x4x4xa multi stemmed black sagebrush plants appear where Y current growth XI fine stems x2xa height the two com despite the difference in coarse stems x3xa plant height andandx4x4xa root have similar biomass because of the 2 munitiesmunities crown R 5757.57 for this determination despite density of plants in the black sagebrush higher the inclusion of a fourth variable our third community there is more coarse and fine equation predicted leaf weight woody material in the basin big sagebrush com- munity table 3 Y 235323.53 0290.29 XI 0930.93 x2xa iffeifweif we assume both populations are the same 020.2oe 0 x3xa 0350.35 x4xa age assumption is necessary because actual age Y leaf weight XI coarse stems x2xa of black sagebrush plants could not be esti- where xa fine stems and x4xa plant mated the rate of woody biomass accumula- height of plant x3 four variables in the equation tion was 13213.2 gm2gmyeargmgmeguyearyear and 64564.5 gm2yeargnygnyearguyearear for density these the data black and basin big sagebrush respectively the accounted for 64 of the variability in 1992 BLACK SAGEBRUSH demography 317

30

20- 0 0 0a

A 10 A C A A A A A U A A AA A A AAA A A A C C0

0

0 wa 88 9sssi&9 0 OS

10 000ooo 0

20 1

4 3 1 2 0 T1 2 3 4 canonical fncanc 2

1 plot fig of black sagebrush group membership based on plant D characteristic discriminant equations where mature A patriarch M senescent and 0 dead 0 young

agesizeAge Size classes material in the canopy to 30 and complete the selected variables for both plant andI1 separation of the stems the separated stems andana formed mound characteristics were important contrib- U shaped flutes with the open end of the U toward utors in distinguishing between agesizeage size classes the former center of the stem it was not possible to and were good indicators of group composition establish the maximum age of the class because fig 1 very few misclassifications occurred by the center of the stem was missing use of the resulting discriminant functions the individual section had at least 40 growth rings the bulk of the black sagebrush stands was senescent plants formed the composed of mature plants 20 60 cm tall with next appar- ently older agesizeage size class in this class canopiescanopies 20 50 cineinelncm in diameter table 4 this at least 50 of the canopy was dead black is a wide range in height and canopy size but older sage- brush plants do not get taller probably the mature agesizeage size class was distinguished from because they have no central stem to support the young plants by the presence of up to 10 dead canopy the diameter of the crowns does increase material in the canopy and the beginning ofthe there is a marked increase in woodywoody biomass separation of the stem into individual cambium between the patriarch and senescent classes bundles the patriarch class was distinguished seedlings and young plants constituted only from the mature class by an increase in dead 6 of the black sagebrush populations tabletabiemablemabie 4 volume 52 318 GREAT BASIN naturalist 2 nova and A 2 SE and oven dry biomass gm of artemisia TABLETABLF 3 mean density stemsmstemsm plus standard error dentata from a previous study young et al 1989 dentatatntmentata subsp dentatatntritridentatetndentatatridentata data for A ttltntrityl dentata subsp tntri 20 biomass per m total coarse fine CAG leaves species density SE 2 g SE g SE g m SE g SE g SE 1550 240 90 520 60 180 45 130 40 artemisiaArtemi wi nova 22 04 750 tridentatetndentatatridentata 420 artemisia dentatattltntrltri 970 110 170 40 130 50 2120 subsp tntrltrityldentata 11 03 850 100

breakdown of characteristics for individual agesizeage size classes demographic TABLF 4 artemisia nova crown and biomass table related to age for younger plants but once stems separate ages black sagebrush communities by growth classes classes are are not based on annual rings characteristics crown characteristics biomass dead coarse fine percentage age branchletsbranchlessbranch lets stems stems CAG leave stem Age size height diameter density of stand years agesize g g g g number class cm cm

1 2 5 0 0 15 5 10 1 seedling 5 5 30 5 5 30SO 28 64 30 40 1 young 10 20 5 10 50 0 30 50 plant 140 120 80 60 multiple 60 20 60 10 50 80 10 mature 420 140 100 multiple 17 40 patriarch 20 60 20 80 60 30 560 640 60 30 multiple 12 20 60 20 100 30 60 980 senescent 910 320 0 0 multiple 5 dead 20 60 20 100 0 100

mound was disturbed the dark seedlings being very rare the separation surface of the with by a grayish shade mounds seedling and young plants was based color was replaced between reach their maximum height with this the occurrence of coarse woody biomass in appear to on black sagebrush mounds of latter class young plants had entire stems growth stage of the had perennial grasses associated with no evidence of division ofthe cambium mature plants with the sub canopy area the most frequent mound types perennial grass was squirreltail accumulations increased with the class had a litter each black sagebrush agesizeage size riarchdiarch agesizeage size class but height ofthe mound mound the patpatriarch corresponding type of sub canopy did not increase apparently trapping of sub- entire study was only seedling found in the aerial deposition material and saltation particles between mounds obvi- located in an interspace must be related to growth stage of black sage- not a valid sample but the ously one seedling is brush plants in terms of crown architecture critical factor in the dynam- lack of seedlings is a subaerially deposited particles are obviously communities studied the first detect- ics of the very unstable and subject to redeposition ifthey occurred under young plants only able mound fall in the largelylargrely bare interspace among shrub the sub canopy area under black sage- 5 10 of mounds young and evans 1986 if litter accu- plants in the young plant agesizeage size class brush mumulationlation increases on patriarch mounds why covered with litter table 5 the litter was was do they not trap these secondary erosion prod- composed of fragments ofblack sagebrush leaves ucts and the mound keep growing in height agesizeage size class the cover in the mature plant canopy structure changes with the patriarch and the weight of litter increased table oflitter size class with increasing bare stems and mounds were easily distinguished by agesizeage 5 the spreading but not taller plants it would appear height and surface soil color and texture both that aerial dynamics of the crown of black sage- surface of the mounds appeared darker in the brush plants influence mound height color and the reddish tinge to the clay surface with the senescent agesizeage size class adivergence soils of the interspace was not apparent if the 1992 BLACK SAGEBRUSH demography 319

TABLE 5 mound characteristics in relation to agesizeage size classes of artemisia nownomnouanova illustrates change with agesizeaeeage size classes of shrubs that mound characteristics

mound litter black sagebrush dlandiandiameterleterieter perennial CheatCaeatgrasscheatgrass growth classes height max mm cover grass min depth weight grass density cover cm cm cm number cm g per mound W samples seedling 0 0 0 0 0 0 young plant 0 0 1 2 5 60 30 5 10 mature 05 40 0 2 6 5 15 80 40 40 60 1 patriarch 15 480 20 15 76 10 15 100 60 80 2 3 senescent 690 28 12 21 10 15 100 60 80 2 3 dead 720 21 60 15 10 15 100 60 80 5 25 970 64 5 6

TABLE 6 mean nitrate level mgkgm kg of soil at the stem canopy edge and outside the canopy of black in relation to maturity classes budbuckskinbug skin mountains nevada a sagebrush plants

location agesizeAge size class stem canopy outside ppm agesizeagesizcAge size ppm ppm class mean 0 young plant 47 b 47h 434 3 hi 41 hi d mature 666.6gg g 44d44 patriarch 669 55g55 g 40j404.0 j 54c54 c 10510 5 d 12012.012 obb senescent 80soee 10210.210 2 a 13213 2 a 113 c dead 113c 70f70 f losios10510 5 a 84c84 c 7.070 b 70f70 f 71f71 f 7.575 b mean location a 75b75 87a87 80b80 b 60gocgoec means followed bytheby the 1 same letterietter are not significantly different at the 01 level means of probabilityprobabilityasasis determined by s of location followed by the same letter areure not duncan multiple range test means aieale significantly different at the 01 level ofprobability as ofagesizeof agesizeage size classes followed by the same letter determined byduncanby duncandunean s multiple range test aiealeare not significantly different at the 01 level ofpiobabilityasof probability as determineddetel mined by duncanduneanDinicanseanss multiple range test

in herbaceous species composition on the beneath the canopiescanopies of desert shrubs has mounds occurs table 5 some been mounds documented by the research ofoanN E become densely covered with ofn west and cheatgrasscheatgrass as workerscoworkersco charley and west black sagebrush plants become 1977 west and senescent and skujins 1977 west 1979 others support colonies of squirreltail nitrate levels of sur- face soils dropped significantly P oi0101.01 after the black sagebrush plants die litter once the black sagebrush plants died nitrate levels weight continues to increase and litter changes in surface soils at the edge of shrub in appearance litter under dead plants is mounds com- increased with apparent posed of stringy bark fragments and individual increasing age of black sagebrush plants and mounds black sagebrush leaves cannot be distinguished these areas cor- in the litter respond to the micro topoedaphic situation described as coppice benches by eckert et al 1989 for soil nitrate levels shrub mounds in big sagebrush com- munitiesmunities apparently the increase in soil nitrate surface soil nitrate levels were higher results from leaching from the mounds once beneath shrub canopiescanopies than in the interspace black sagebrush plants are dead and grasses table 6 levels were highest next to shrub dominate the mound soil nitrate levels decrease stems nitrate levels beneath the canopy rose as agesizeage size classes of black sagebrush indicated mounds and black sagebrush older plants and mounds this is in itself an community structure indication that agesizeage size classes actually do we did not find grass dominated mounds or reflect increasing age development the of ver- grass dominated mounds with black sagebrush tical and horizontal patterns in soil nitrogen seedlings we did note the remains of mounds attributed to the localization of litter fall that appeared to be eroding away apparently volume 52 320 GREAT BASIN naturalist

desert ecosystems of utah soil biol- formed and eroded in dominated semisemidesert mounds are dynamically ogy and biochemistry 9 357 365 death WARD W H relation to the establishment and eventual ECECKFRIKERr R E JR F F PFFFRSONPETERSONpererson M K EJR role of sagebrush plants failure to find BLACKBURN AND J L sre111fnssrfphfns 1989 the osblackofblackof black the of semiarid mounds may be a function of soil surface morphology in the function grass dominated agricultural experiment station livestock sheep feral rangelands nevada herbivoreherbivoryherbivory by domestic TB 89 01 university of nevada reno tailed jack bulletin horses equus cacaballascaballusballus and black EVANS R A AND R M LOVFLOVE 1957 the step point rabbits lepus californicuscaliforcaliforornicusnicus grass dominated method of sampling A practical tool in range research 10 208 212 fail to since grasses cannot journal of range management mounds may persist 1979 geologic map of of leaffall andcanopy HUDSON D MMANDWMAND W M ORIFLORIEL maintain mounds because buckskins range nevada map 64 nevada bureau black sage- the structure differences compared with of mines and geology mackay school of mines uni- brush plants the only patchy vegetation versity of nevada reno 1988 utilization of encountered in the communities was groups of LUGASKI T P AND J A YOUNG mapper data and aerial photog- plants perhaps rabbitrabbitbrushbrush LANDSAT therthermaticthermoticmatic rabbitrabbitbrushbrush raphy for mapping the geology soils and vegetation increases after relatively short lived squirreltail assemblages of the buckskin mountain ranges plants die or are reduced by grazing in an nevada american congress on surveying and map- photogrammetry 1988 adjacent big sagebrush community we pre- ping and american society for three episodes of seedling 218 227 viously determined W WASSFRMANWASSERMAN 1974 applied linear sta- and 57 years before neiernelerNFIFRNETER J AND establishment at 12 42 tististicaltical models richard D irwininvin inc clus- PROFFETT 1976 1985 young et al 1989 plant ages were proffetpropfetPROFFF 11r J M JR AND B H proffePROFFF 11 stratig- tered around these apparent establishment raphy of the tertiary ashflow tuffs in the yerington periods of district nevada nevada bureau of mines and geology dates the clusters may represent university of or report 27 mackay school of mines desirable climate for seedling establishment nevada reno 28 appp of a single season when establishment occurred weslwest N E 1979 formation distribution and function 659 they may also represent variability in growth plant litter in desert ecosystems pages 647 in classes we R A perry and D W goodall eds aadlandaridlandandlandfridland ecosys- ring deposition or recognition the and management vol I1 too broad to tems structure functioning constructed in this study are much international biological Proprogrammaprogrammegramme 16 cambridge establish- pinpoint this type of episodic stand university press londonL don in for black sagebrush perhaps black sage- westWFSI N E AND J SKUJINS 1977 the nitrogen cycle in ment semidesertdesert ecosystems brush communities not renewed catastrophically north american coldwatercold water semi renewal at such ecologiaecologicEcologia Planplantarumplantariumtarum 12 45 53 by wildwildfiresfires only require stand EVANS 1986 erosion and deposi- dead YOUNG J AAANDRAND R A low levels 5 of the stand standing tion of fine sediments from playasolayas journal of andaridarld plants that our one seedling sampled is suffi- environments 10 103 115 EVANS D E PALMQUISIPALMQUIST 1989 cient for community regeneration YOUNG J A R A AND big sagebrush artemisia tntritrldentata seed production weed science 37 47475353 of cheatgrasscbeatgrasscheat grass CITED YOUNG J AaandftipionAND F tipton 1990 invasion literature pages into and environments of the lahontan basin bulletin 368 E D mcarthur E M romney S D smith 1 of 37 41 bl 1 11 I A A 1960 A study sagebrush in ll compilers proceedings of the sym- agriculturalillcil experiment station university of wyoming and P T tueller on cheatgrassCheatgrass invasion shrub die off and lallaramieLai ainiediniealnie posium of the carson aspects of shrub biology and management 1 W D 1945 the plant associations other bibilLLINGSINS INT 276 forest service desert region western nevada butler university general technical report botanical studies 7 89 123 USDA ogden utah B AND P T TUELLERTUFLLFR 1973 artemisia BLAISOIII J PPANURAND R C HOLMCKFNHOLMGREN 1984 managing ZAMORA rangelandselands salt desert shrub ranges arbuscula A longilongilobalongilohalobalohaioba and A nova habitat types in intelintermountainmountain lang 225 242 general technical report INT 163 forest service northern nevada great basin naturalist 33 USDA intermountain forest and range experiment station ogden utah 52 appp 1991 of received 21 october CHARLEY J L AND N E wislwisir 1977 micro patterns 1992 cliakl wis shrub accepted 10 september nitrogen mineralization activity in soils of some great basin naturalist 524 appp 32127321391 32727

MUSHROOM consumption MYCOPHAGY BY NORTH AMERICAN CERVIDS

2 karen L launchbaugh and philip J urness

ABSIRACABSTRACT I1 native mushrooms play an important though often underestimated role in deeldeer elk and north america mushrooms are often caribou diets in noted as an unusual or anomalous food in the diets ofcervidscervidsids diets in the late summer of cerv yet they often dominate and fall in forested areas ofwestern north americaArnericaherlea and U S mushrooms throughout the year in the southeastern are particularly high in protein 16 19 phos phosphorusphoiphol us average 0750 75 and potassium average 2 mushroom production is generally fall aveiage2 also greatest in therefore they aiealeare a highly nutritious food native forages may in late season when other marginally meet basal nutrient requirements of ungulaungulatesungulatedtes

key words caribou cervid deer diet elk mycophagy mushroom nutrition ruminantTuminyuminant wildlife scientists have longiong ionaionolono recognized that desties and insects miller and halls 1969 certain highly nutritious bonus foods fogel fre- and trappe 1978 martin 1979 mushrooms quently contribute significantly to animal wel- have long been recognized as an important fare though their contribution com- to the diet ponent of small mammal diets and may be small acorns fogel eg mushrooms and trappe 1978 however mushrooms are seldom mesquite beans seeking by out these high considered a significant component of cervid quality but generally scarce or ephemeral foods diets even though they have been anecdotally herbivoresherbivores can balance nutrients against lower recorded as a preferred food item discount- quality forages that are more abundant native ing mushrooms as an important dietary compo- mushrooms have often been recorded as a nent maystem from a misunderstanding oftheir bonus food in the diets of deer elk and cari- nutritive value the purposes of this review are bou in 1 north america however their contribu- to assess the contribution of mushrooms to tion to cervid nutrition is not commonly cervid diets 2 summarize the known literature understood on the nutritive value of mushrooms to anguungu the term mushroom refers to the fleshy lates and 3 assess the implications ofmycoph fruiting body sporocarp of many species of agy to habitat selection and nutritional ecology fungi mushrooms are technically not plants they belong to the kingdom mycenae mycetae under the contribution OF MUSHROOMS TO five kingdom classification system whittaker DEER ELK AND CARIBOU DIETS 1969 the primary mushroom producing fungi are in the group called basidiomycetes but mushroom consumption by deer many mushrooms eaten by wildlife including many studies moreis are ascomycetes mushroom have recorded mushrooms in produc- diets of both mule tion is triggered when species specific odocoileus hemionus and require- white tailed odocoileus ments of minimum temperature and moisture virginianusvirginianus deer table 1 diet composition conditions are met smith and weber 1980 estimates range from a trace to a majority of the diet mushroom consumption mycophagy has on the upper limit 71271.2 mushrooms on a fresh been recorded for many wildlife species in in weight basis were recorded in fallfalifailfiallbhail deer diets in north america mushrooms are eaten by anguungu alabama kirkpatrick et al 1969 65.8658 lates eg deer and elk small mammals 658 in eg august diets in arizona hungerford 1970 and squirrels and armadillos as well as birds tur 59559.5 in august diets in montana lovaas 1958

f range science department 2present utah state university logan utah 84322523084322 5230 plesent addressaddless range and wildlifeWild lifie management department texas tech university lubbock texas 79409212579409 2125

321 volume 52 322 GREAT BASIN naturalist over seasonseasonlseasonal elk and caribou diets in north america averaged TABLITABLE 1 proportion of mushrooms in deer

0ofif diet species sourced b summer fall winter kindkm iofdataof data state or province vegetation type spring odocoileus hemionus mule deer 03 absobs bites 31 colorado sprucefirpine forest vol 21 00 121 00 00 rum montana sprucefirpmesprucefirpine forest absobs mass 10 meadow 70 utah dry mountain 150 absobs mass 10 utah mature conifer forest absobs mass 10 stagnated conifer forest 140 utah 54 93 absobs mass 4 utah conifer foreforestoakstoak woodland absobs time 16 forest 164 arizona mixed conifer 10 rum vol 20 chaparral oak woodland california 00 130 40 rum vol 8 british columbia conifer forest 00 white tailed deer odocoileus virginianusvirginianus rum mass 26 forest 137 67 91 new brunswick coniferconiferdeciduousdeciduous 00 absobs mass 9 forest 00 00 450 maine pine hemlock 08 45 absobs time 14 clearcutclear cut forest 16 02 pennsylvania 198 84 62 rum vol 12 USU S oak hickory pine forest 21 southeastern 156 86 49 rum vol 12 USU S mixed pine forest 04 12 southeastern 164 54 32 rum vol southeastern USU S southern evergreen forest 06 19 00 400 25 00 rum vol virginia eastern deciduous forest rum vol 19 forest 00 106 70 00 north carolina oak hickory pine 107 rum vol 19 forest 02 334 26 south carolina mixed pine 90 138 rum vol 19 southern evergreen forest 00 97 19 g0georgiagar ia 104 267 132 rum vol fiorflor forest 14 1 floridai E southern evergreen 92 rum vol I111I florida sousouthernthem evergreen forest 252 rum vol I111 I1 scrub oak forest 19 florida pine 00 712 05 174 rum vol alabama southern pine hardwood forest rum vol I1 hardwood forest 73 48 08 alabama southern pine 05 absobs bites 28 bluestem range 05 15 35 louisiana pine 04 19 07 absobs bites 29 louisiana pine hardwood forest absobs bites 29 forest 010 1 21 02 louisiana clearcutclear cut 10 70 rum mass 25 mixed hardwood forest 30 340 texas pine 00 00 43 10 rum rel fret 30 oklahoma oak savannah rum vol 22 forest 20 wisconsin northern hardwood rum vol 2 hardwood forest 10 00 minnesota northern 21 00 rum vol 15 dakota forest 00 40 south pine 07 05 05 rum vol 23 south dakota pine forest

elk cervus elaphusclaphus 10 rum vol 3 deciduous forest virginia eastern 53 rum mass 17 saskatchewan pine forest rum mass 17 forest 42 saskatchewan mixed 83 absobs mass 7 dn meameadowdow 42 7 utah dry mountzmountain 187 557 absobs mass mature conifer forest 7 utah 184 554 absobs mass utah stagnated forest 03 absobs time 13 california pacific rain forest rangifer tarandus 5 caribou 00 250 120 00 rum vol newfoundland conifer forest 04 rum vol 24 canada conifer forest 18 northern 12 rum vol northern canada boreal forest 20 absobs vol 6 foresttundratundra 00 120 100 alaska spruce forest 450 rum vol 27 alaska sprucespruce forest

1 availableiv ul ible A ddashisliasli listed ilsitsis in diet ineansindansme ins no ddatait weiewere 1 according to aldnclialdrichaidrich 1963 1 or01 vegetationveget ttionstion arealreire cordingic generalGLIILI descriptiondlsliiptlons livengivegiven by authors beget data hurnrum rinnenumenrunnen1 contents 5 tidlid 1972 6 Boertje 1984 obsabs scrvationalobseivthonil ditl mdand pittonpattonputton 1938 obeale44bldle4bealeBeale and darby 1991 bergerlid5bergenidBerger 1959 2 Aldous mdand smith 1938 ab3baldvvin3b ildwm 1962 al1eally 1971 keskey tot refelreielefcrenccsLULLS IAiadains1adimsdains 1961 irlowarlow mdand hooper 1971971 13harper1311arper131113 Harperarper 04iiely key enles 10descllarnp et illolloulil 1979 11flarlollhirlow 12llarlow1211 1977 8 Cowanowmcwm 1945 ac99cCrawfordi iwtoid 1982 lodesllnmp alil 1951 21lovaas2121lovais1958Lovaas 1958 22mlc22 McCafferyiffery 7colinsl9777 Collins illuli 1919kirkpitricketKirkpatrick et iliillcliil 1969 20leopoldet 17iinnt979unt 1979 18kels1818kelsallkeisKelsallKels 1968 1986 et alil 1990 151115iiill1511illilliii andmdnurisl943harrisnarris 1943 06iiingeford16iiungerfoidl9701970171119701711unt 1974 27skoog27 Skoog 1968 28thill28rhill28 rhillThill mdand martinM irtmiram 29tlrill29rhillet 1967 25short25 Short 1971 2626skinner26skmnerSkinner andfelfermd lelfei ltet alil 1974 23sclienck23slliciilketet iliillil 1972 2424slottei24scotterScotter 30van3vmvilcdcbreedevreede 1987 31wallino31willmoltet al1 9721972 19921 MYCOPHAGY BY CERVIDS 323

late summer and fall are generally the sea- in caribou r 1 rangifer tarandus diets sons ofor greatest mushroom 1 when consumption prob- mushrooms are available they may ably because mushroom production is constitute generally 10 25 of caribou diets but they may greatest then though mushroom biomass average pro- as much as 45 table 1 and have duction is seldom recorded in diet studies been sev- recorded as high as 84 in one eral authors note that mushroom individual production is skoog 1968 even in 11 triggered by fall winter reindeer unerr- rains tevis 1952 hungerford ingly detect snow 1970 umessurness covered frozen mushrooms 1985 consuming them mushroom greedily karaevbaraev 1968 the species most consumed by boertje 1981 reported deer are not that most genera of precisely known because species mushrooms are taken without are seldom recorded hesitation by car- in diet surveys and prefer- ibou mushrooms of the genera boletus ence studies have not been conducted copri- in addi- nus lactarius lycoperdon morchella and tion species identification is rare because most russula have been listed as major dietary com- wildlife researchers are not acquainted with ponents karaevbaraev 1968 skoog 1968 boertje 1981 common mushroom species and professional taxonomic help is difficult to obtain cowan NUTRITIVE VALUE 1945 in most field studies mushrooms are OF MUSHROOMS categorized into groups such as field mush- many rooms mixed mushrooms or simply authors state that deer elk and cari- fungi bou however when listed species of the following strongly prefer mushrooms and in some cases genera are consistently taken by deer amanita actually travel from site to site seeking mushrooms hungerford 1970 ahnAnnarmillariaillaria healy 1971 the obvious question is why miller and halls 1969 boletus cowan 1945 what nutritional benefits do cervids gain from fungi some hungerford 1970 beale and darby 1991 authors consider mushrooms nearly devoid clavariaclauciauClavarlaariaaTla dixon 1934 clitocybe cowan 1945 of nutrition while others suggest they beale and darby 1991 cortinariusCotti nariusnafius compare favorably with soybeans or spin- hunger- ach ford 1970 morchella cowan 1945 lactarius crisan and sands 1978 miller and halls 1969 lentinusLenjentinustinus dixon 1934 little is known about the true nutritive value polyporus skinner and telfer 1974 russula of mushrooms since few comprehensive studies have cowan 1945 miller and halls 1969 hunger- been conducted crisan and sands 1978 conducted ford 1970 and suillusquillus miller and halls 1969 a thorough literature review on the nutritive value of wild mushrooms to mushroom consumption by elk monogastrics eg humans several range and wildlife scientists have collected and elk cervuswervus elaphusclaphus diet studies rarely analyzed mushrooms prominent in record fungi as a component an extensive liter- ruminant diets but the nutritional procedures used ature review of elk food habits in 1973 did by most range not and wildlife scientists were mention mushrooms as a recorded food designed to analyzeanalanai e item grasses and forbs when these lyerye kufeld 1973 however at least four studies procedures are aappliedPpliedpiled to mushrooms the results have recorded mushrooms as a component of are often ineinclneincorrectlyorrectlycorrectly interpreted because mushrooms elk diets table 1 composition estimates range are much different from vascular plants from a trace to as high as 75 on a dry weight in their chemical composition information basis collins et al 1978 As with deer mush- further on room the nutritive value ofmushrooms can be gained consumption is greatest during seasons of from greatest availability late research on mycophagy by insects and summer and fall small mammals it seems reasonable the following discussion is a to assume that mush- summary and room species sought interpretation ofnutrition studies by deer would also be to assess acceptable to elk the value of mushrooms to ruminant though evidence is lacking animals collins 1977 listed species ofAof leuria boletus and russula as important and highly preferred moisture content of mushrooms dietary components over 80 of the fresh weight of most mush- mushroom consumption by caribou rooms is water table 2 this large water pro- portion requires that the mushrooms have often consumer eat large been recorded as volumes to obtain nutritional benefit very palatable and highly sought although dietary items high water content rarely restricts intake the volume 52 324 GREAT BASIN naturalist

of wild mushrooms table 2 nutritivenuti itiveetive value and digestibility N free phos- digesti- initial crude source ash fat extract fiber calcium phorus bility composite samples based on moisture protein 316 20890820gos8 crude 7 available 348 81 48 5 species 90 50 480 150 crude species available 230 056 4 215 66 39 542 13813 8 crude 009 species available 839 010 042 2 species in cattle diets summer 220 055 2 250 010 species in cattleeattleeattiec11 tletie diets fall 046 588 1 221 008 species avavailableai able winterwinterwintelter 894 647 1 231 007 047 species avalavailableableabie spring 876 053 566 1 290 005 species available summer 872 004 053 599 1 species available fall 859 248 808 6 species in deer diets 889 213 775 6 895 241 species in elk diets 317 NDF 003 070 900 3 diets summe i 347 317ndf species in caribou suminer 315 NDF 003 071 900 3 diets fall 353 315ndf species inilliii caribou 299ndf299 NDF 003 079 910 3 species in caribou diets winter 400 ofot ticsliarsticali weight iiiitialil i sture which isis expressed isas frs it s ai ofayol01 dry iiinlatteriatterittel except initieniti inoistuic 1968 6pilksln6palleseii 1979 all daiadatad il esprssedcipicssid ofhy 313oeiije 1981 40isanisan andmdsindssands 1978 5kllsill5kclsall alil 1984 213jgxtad indand diliympkdalryinple 1968 4bocrtjc 404nsm keyKI tto reficldinusremes JBlbluicblairjblairlair ett 2busstid 7syiilioist7syrjala qitait 1986

from the microbial degradation of addition of water to the rumen per se has little comes fibrous cell walls however fungal cell walls are effect on intake because it is easily absorbed or different from those of higher plants the removed van soest 1982 mushrooms may in much component of fungal cell walls is fact be an ilimportant source of water for some primary whereas plant cell walls are mostly cellu- mammals fogel andandtrappetrappe 1978 chitinchitinwhereas lose crisan and sands 1978 martin 1979 polymer linked mushrooms as an energy source chitin is a N acetylglucosamine with p 14 bonds similar to cellulose unlike the mushrooms like true plants contain lipids fiber of higher plants chitin contains a signifi- or fats nonstructural carbohydrates and fiber cant proportion of nonprotein nitrogen as aann d that are all used as energy sources by ruminants amino sugar A P3 flucanglucan with P3133 13 linkages anand mushrooms ranges of cell wallwaliwaiiwwall the average gross energy of rigpigp3163 16 branches also forms a part the from 300 to 400 kealskeais per 100 grams dry weight martin 1979 additionally lignin and pectin with fleshy fungal tissue compares favorably are not known to occur in fungi many fruits and vegetables but is less rich in energy than seeds or nuts martin 1979 protein content of mushrooms the fat content of edible mushrooms ranges S investigators used the term vegetable fromfrom I1 to as high as 20 Ccrisanri san and andssands early describe mushrooms because analysis 1978 on average however mushrooms con- meat to native mushrooms contain 20 2 6 fat fat component of fungal revealed that tain the matter as protein peck 1895 includes free fatty acids mono di and 50 of their dry tissue studies on mushroom protein con- triglycerides stesterolsrols sterol esters and phos more recent suggest that mushrooms probably rarely phopholipidslipids tent reach 50 protein by dry weight however on a dry weight basis mushrooms are pri- relatively speaking mushrooms are an excellent corncomposedposed of nonstructural carbohy- marily source there is extreme variation in drates nitrogen free extract table 212 A large protein protein content from a low of about 4 to as variety ofcompoundsofcompounds make up the carbohydrate of high as 44 depending on species stage components including penpentosestoses methyl pen growth and environmental conditions crisan toses hexoseshexoses disacwaridesdisaccharides amino sugars and and sands 1978 by comparison fresh cut sugar alcohols and sugar acids crisan 16 19 the most promi- alfalfa medicago salivasativadativa is generally sands 1978 by comparison 1978 nonstructural carbohydrates in green protein jurgens nent usually calculated by multi- fructofructosanssans sugars dextrin and starch crude protein is plants are determined by kjeldahl trlica 1977 plying total nitrogen analysis by 6256.25 this correction factor is based in plants most energy available to ruminants 199211992 MYCOPHAGY BY CERVIDS 325

on the assumptions that most proteins contain and phosphorus averaging about 0.75 16 nitrogen that 075 these proteins are com- change 1980 crisan and sands 1978 pletely digestible and that martin amounts of nonpro- 1979 both mineral levels exceed maintenance tein nitrogen in the cell are negligible since a requirements of most weaned ungulaungulatesungulatedtes based substantial amount ofnitrogen in mushrooms is on sheep and cattle in chitin and other requirements jurgens nonprotein compounds such 1978 mushrooms also contain calcium but at as urea and nucleic acids crisan and sands lower concentration than phosphorus or potas- 1978 suggested a correction term based on the sium however calcium concentration averages assumption that only 70 of the nitrogen in 0140.14 which would not meet calcium require- mushrooms is in the form of digestible protein ments of weaned deer ullrey et al 1973 cal- 6256.25 4384.38 70n this correction term of cium is often in excess of ruminant needs in 4384.38 may be conservative when considering other forages while phosphorus is more com- the use of mushrooms by ruminants and com- monly inadequate paring mushrooms to other forage eaten by ruminants only 60 70 of the nitrogen in digestibility of mushrooms fungal tissue is in the form of protein moore the degradation of fungal cell walls requires landecker 1982 however this estimate is sim- chitchitinaseinase and P3133 13 and rigPpig3163 16 glucanases ilar to the proportion of nitrogen in proteins in martin 1979 chitin is degradabledearadeoradable in the forage plants 60 80 van soest 1982 fur rumen because of chitchitinaseinase activity by rumen therthermorethedmoremore nonprotein nitrogen such as urea is micmicrobesrobes although there may be an adaptation readily converted to ammonia by rumen period necessary to obtain adequate levels of micmicrobesrobes and is either used for microbial growth chitinasechitinase activity cheekechecke 1991 the ability of or absorbed across the rumen wall nitrogen the rumen microbesmicrobes to degrade the P3 fraction of chitin is unavailable glucans in to monogastrics fungal cell walls is unknown but is probably converted to microbial protein the in vitro digestibility of mushrooms is in the rumen in fact chitinous nitrogen may be very high relative to other ungulate forages more available to ruminants than the cell wall table 2 and may exceed 90 in some easescasescakeakcas es nitrogen ofhigherplants due to the lackofligninlaeklack oflignin in fungi consequently identification of mushrooms in fecal analysis israleisrareis rare boertje 1981 vitamin and mineral composition of mushrooms implications OF MYCOPHAGY BY DEER AND ELK mushrooms are a good source of several vitamins including the B complex and vitamin C to conclude this discussion it is fair change 1980 crisan and sands 1978 to ask how- what difference does it make if deer elk ever these are not essential vitamins for rumi- or caribou eat mushrooms not nants because they can be synthesized or mycophagy by by rumen cervids may be for micmicrobesrobes van soest 1982 important several reasons additionally mush first mushrooms rooms are basically devoid undoubtedly make an of vitamins A and D important though which are sporadic contribution to essential dietary components for cervid ruminants nutrition in mushroom rich environ- ments mushrooms are highly preferred and mushrooms accumulate minerals from the nutritious foods for cervcervidsids particularly in late soil and plant material on which they grow summer and fall in forested areas of western therefore mushromushroomsorns probably contain all the north america and throughout the year in the minerals present in their growth substrate southeast mushrooms may be a particularly crisan and sands 1978 stating average minmin- important protein and phosphorusandphosphorus source in late eral concentrations may be misleading because season when many forages yield only enough mineral concentration variesvaniesvanesvannes greatly depending digestible dry matter to meet basal energy on species and soil fertility example for though requirements short 1975 blair et al 1984 potassium level averages 2 in 24 species from therefore even a few bites of mushromushroomsoinsorns by an several locations it varies from oisols0.18 to 4.848 018 48 herbivore may contribute substantially to crisan and sands 1978 meet ing the nutritional requirements and helping the most abundant to minerals in mushrooms balance nutrients obtained from other forages are potassium averaging about 2 dry weight of quite different composition volume 52 326 GREAT BASIN naturalist

mountains utah performance report for may attract herbherbivoresivores type uinta second mushrooms W 105 R11 2 14 publication no that might federal aid project to mature and stagnated forest areas 771877 18 utah division ofwildlife resources otherwise go unused as foraging areas rasmus- COLLINS W B P J URNESS AND D D auslinAUSIINAUSTIN 1978 elk sen 1941 collins et al 1978 warren and mys diets and activities on different lodgepole pine habitat 4279942 799 810 1991 additionally mushrooms may segments journal ofwildlife management terud relationship of the food dietary supplement when COWAN I1 M 1945 the ecological become an important columbian black tailed deer odocoileus seek densely forested of the herbherbivoresivores are forced to henihenthemionusionuslonus columbianuscolumbianus richardson in the coast areas for protection from biting insects or pred- forest region of southern vancouver island british ators bergerud 1972 mushroom production columbia ecological monographs 15 111 139 seasonal food selection and digest- is usually greatest in dense forested areas in CRAWFORD H S 1982 is tailed deer in central maine because mushrooms do not require sun- ibility by tame white in part journal of wildlife management 46 974 982 for growth value pages light CRISAN E mandVANDVANDAA SANDS 1978 nutritional press finally fungi play an important symbiotic 137 168 inm edible mushrooms academic new role mycorrhizal relationships with several york in 1979 conifer including ponderosa pine DESCIIAMP J AAPP J URNESS AND D D AUSHNAUSTIN species from lodgepole habi- 1984 the spores of fungi are summer diets of mule deer pine kotter since of wildlife management 43 154 161 destroyed in the rumen cerbiherbi tats journal apparently not in DIXON S 1934 A study of the life history and food habits to J game vores may serve as vectors for fungal spores of mule deer in california california fish and initiate mycorrhizal associations fogel and 2031520 315 354 trappe 1978 FOGELFOGFL R AND J M TRAPPE 1978 fungus consumption mycophagy by small animals northwest science 52 1 31 literature CITED harlowrfhanlowHARLOW RFR F 1961 fall and winter foods of florida white tailed deer quarterly journal of the florida academy 38 ADAMS W H 1959 chaccolocco deer range analysis and of science 24 19 by management implications proceedings of the south HARLOW R FFANDRAND R G HOOPER 1971 forages eaten eastern association of game and fish commissioners deer in the southeast proceedings oftheodtheofthe southeastern 13 21 34 association ofgameof game and fish commissioners 251825 18 46 habits of of ALDOUS S E AND C F SMIIIISMITH 1938 food harberHARPFRHARPER J A 1962 daytime feeding habits roosevelt minnesota deer as determined by tomachstomach analysis elk on boyes frameprairieprame california journal of wildlife transactions of the north american wildlife confer- management 26 97 100 ence 3 756 757 HEALY W M 1971 forage preferences of tame deer in a of american ALDRKALDRICH II11 J W 1963 geographic orientation northwest pennsylvania clear cutting journal ofofwildwild- 27 529 tetraonidae journal ofwildlifeofwildfife management life management 35 717 723 545 HILL R R AND D HARRIS 1943 food preferences of C P pallonPAIIONPATTON 1938 A preliminary BALDWIN W P AND black hills deer journal of wildlife management 7 the food habits of elk in virginia transactions study of 233 234 of the north american wildlife conference 3 747 hungerford C R 1970 response of kaibab mule deer 755 to management of summer range journal of wildlife BFALI- D M AND N W DARBY 1991 diet composition management 34 152 162 of mule deer in mountain brush habitat of southwest 1 summer autumn and winter diets of utah publication no 911491 14 utah division of HUNHUNT H M 1979 ern canadian field naturalist 93 wildlife resources elk in saskatchewan BERGERUD A T 1972 food habits of newfoundland car 282 287 brrgfrud animal feeding and nutrition ken ibouabou journal of wildlife management 36 913 923 JURGENS M H 1978 company dubuque iowa A J AND A V DALRYMPLEDALRYMPLF 1968 beefheifers dillhuntdallhuntdall Hunt publishing bjugstadBJUCSIAD resources P S on ozark ranges bulletin no 870 missouri agricul- KARAEV G I1 1968 reindeer fodder in tural experiment station Zhigunzhigunovov ed reindeer husbandry USU S department yield BLAIR R M R ALONIZAL ONI AND H F momusMORRIS 1984 of commerce springfield virginia of caribou nutrient composition and ruminant digestibility KELSALL J P 1968 the migratory barren ground fleshy fungi in southern forests journal of wildlife of canada queen s printer ottawa ontario canada management 48 1344 1352 kirkpatrick R L J P FONIFNOIFONTENOT AND R F HARLOW denall LJ bondjrbonujrboeiitje R D 1981 nutritional ecology of the denah 1969 seasonal changes in rumen chemical compo- thesis caribou herd unpublished master s university nents as related to forages consumed by white tailed of alaska fairbanks deer oftheodtheof the southeast transactions oftheodtheofthe north amer- diets of the denahdenall caribou herd 1984 seasonal ican wildlife and natural resources conference 34 165 alaska arctic 37 161 229 238 CHANGE S 1980 mushrooms as human food bioscience chanceCIIANCcnance r T KOTTFR M M 1984 formation of ponderosa pine 401 kotter 3039930 399401399 ectomycorrhizae after inoculation with feces of tassel 1991 applied animal nutrition macmillan CIIEEKE P R eared squirrels Mycmycologiamycologicologia 76 758 760 publishing co inc new york elk elk KUFELDKUFFLD R C 1973 foods eaten by rocky mountain COLLINS WBW B 1977 diet composition andandactivitiesactivities of journalrnalanal of range management 26 106log106113113 on different habitat segments in the lodgepole pine jou 199211992 MYCOPHAGY BY CERVIDS 327

LEOPOLD A S T RINEY R mccain ANDLAND L TEVIS 1951 SMITH A H AND N S WEBER the jawbone deer herd game 1980 the mushroom bulletin no 4 depart- hunter s field ment of guide university of michigan press ann natural resources division offish and game arbor commisioncommissionCommision SYRJALA QVIST L 1986 improvement LOVAAS A L 1958 mule deer food habits of natural pasture and range use utilization by little belt mountains montana sheep in 0 gudmundsson ed grazing journal of wildlife research management 22 275 282 at northern latitudes plenum PresspressnatoNATO new york MARTIN M M 1979 biochemical implications of insect TFVIS L 1952 autumn foods mycophagy biological review 54154 1 21 of chipmunks and golden MCCAFFERYMcCAFF mantled ground squirrels ebyFRY K R J TRANETZKI AND J PIECHURAPlEC HURA 1974 in the northern sierra RJ andaandj huba nevada journalal summer foods of deer in northern wisconsin journ of mammalogy 33 198 205 journal THILL of wildlife management 388153821538 215 219 R E AND A MARTIN 1986 deer and cattle diet overlap on MILLER H A ANDANDLL K HALLS 1969 fleshy fungi com- louisiana pine bluestem range journal of monly eaten by southern wildlife USDA forest ser- wildlife management 50 707 713 THILL vice research paper SO 49 R E H F MORRIS JR AND A T HARRFLHARREL 1990 MOORmooreMOORF E LANDECKERLANDECKFR E 1982 the fundamentals of the nutritional quality of deer diets from southern pine fungi prentice hall inc englewood cliffs new hardwood forests american midland naturalist 124 jersey 413 417 PALLE pallesenPALLESFNS F N J D 1979 nutritive value of mule deer and elk TRLICA M J 1977 distribution and utilization of carbohy- diets and forages on lodgepole pine summer range in drate reservesreserves in range plants in R E sosebee ed utah unpublished mastermaster s thesis utah state univer- rangeland plant physiology society for range man- sity logan agement denver colorado PFCK C H 1895 report of the new york state botanist ULLRFY D E FTET AL 1973 calcium requirements of page 113 reported mlin L B mendel 1989 the chem- weaned white tailed deer fawns journal of wildlife ical composition and nutritive value of some edible management 37 187 194 american fungi american journal of physiology 1 URNESSURNFSS P J 1985 managing lodgepole 225 235 pine ecosystems for game and range values in lodgepole the RASMUSSENRASMUSSFN D I1 1941 pine biotic communities of the kaibab species and its management symposium plateau arizona proceedings ecological monographs 11 229 275 cooperative extension service washington SCHENCK T E R state uni- L LINDER AND A H richardson versityversity pullman 1972 food habits of deer in the black part 11 in hills II VAN SOSOESTEST PPJJ 1982 nutritional ecology of the southern black hills bulletin no ruminant 606 south dakota comellcornell university press ithaca new york agricultural experiment station vanvreedeVANVRFEDFVAN VREEDE G T 1987 ofwhite SCOITERSCOTTER G W 1967 seasonal diets ofwhite tailedtalled deer the winter diets of barren ground in south caribou northern central oklahoma unpublished master s in canada canadian field naturalist thesis 81 33 39 texas tech university lubbock WALLMO 0 C W L RFGELIN AND D W SHOSHORTRT H L 1971 forage digestibility and diet of deer RFKREICHERThebl on 1972 forage use by mule deer southern upland range journal of wildlife manage- relative to logging in ment 35 698 706 colorado journal of wildlife management 36 1025 1033 1975 nutrition of southern deer in different sea- WARREN J T AND I1 MYSIFRUDMYSTERUD sons journal of wildlife management 39 321 329 tandlTANDI 1991 fungi in the diet of domestic sheep SKINNERSKINNFR WRANDEW R AND E S TFLFER 1974 spring summer rangelands 13 168 171 WHITTAKER R H 1969 new and fall foods of deer in new brunswick journal of concepts of kingdoms of wildlife management 38 210 214 organisms science 163 150 160 SKOOG R 0 1968 ecology of the caribou in alaska unpublished doctoral dissertation university of cali- fornia berkeley received 15 june 1992 accepted 25 september 1992 gigreateat basin naturalist 524 appp 328 334 terrestrial vertebrates OF THE MONO LAKE ISLANDS california

3 4 1 2 and S hallhailhalihall14 michael L morrisonmornson wiilwilliamWilliainlain M blockblock2blocke joseph R jehl jr linnea

between the two islands caobapaohapaoba and legitnegit in mono lake 1 we compared vertebrate populations major ABSTRACT colonization mice mainland to further elucidate the mechanisms underlying island deer california and the adjacent deer were voles microtus montanusinonfanustanus were captured on paohapacha but only mice peromysciisperoinyscus maniculatusmamculatus and montane pachapaoha were captured on the mainland overall rodentrodentabundanceabundance on captuicaptureded on legitnegit in contrast eight species ofrodents captai than on pachapaoha or the mainland adult deer mice and the mainland was similar but on legitnegitncgit it was about three times greater externalnalnai body characteristics than mainland mice coyotes caniscams fromti om paohapacha were significantly P 05 smaller in most extel islands and the mainland no amphibians or reptiles or oflagomorphslagomorphaoflagomorphs were observed on the lcitranlatrambatram i and one 01 two species means low numbers on the mainland rafting and human transport are probable weiwelweree found on the islands both occunmccunoccurreded in on the islands may have modified historic predator prey of colonizationcolo mationmatlon foiforooiooloor micernicemleeanice and voles the occurrence of coyotes relationships and thus the population of rodents and lagomorphslagomorpha

peromyscus inaniculatusmamculatus mieMicmicrottismicrotusrottis montanus land bridge kelikellkeokeifkey words mono lake islands colonizationcolonbationkationmation ISLANDS island animal populations have attracted MONO BASIN AND much scientific interest because they serve as the hydrologic drainage basin natural experiments for the study of coloniza- mono basin is mono basin is surrounded by the tion dispersal extinction competition and for lake the nevada to the west and the great basin other biological processes macarthur and sierra to the north east and south mono lake wilson 1967 because islands are small and ranges estimated at 500000 years of age is one of the populations inhabiting them are more isolated oldest lakes in north america because no vulnerable to stochastic events than their main- water naturally flows out of the basin and land counterparts because of longtermlong term evaporation coupled with island zoogeogra- most previous studies of water diversion the lake s salinity is about 252.5 island occu- phy have emphasized patterns of times that of the ocean in october 1986 the pancy morphology and genetics of restricted surface area of the lake was about 177 kmkm2 subsets of the islands fauna reviewed by mono basin ecosystem study committee peltonen and hanski 1991 our goals were to 1987 compare island and mainland vertebrates of there are two major islands in the lake mono lake and the surrounding mono basin pachapaoha island at about 77tt7.7 km2 and legitnegit island california in light of natural and human influinflux at only about 131.3 km2 fig 1 pachapaoha formed encedencee processes this area was of interest from volcanic activity and an uplift of lake sedi- 1850 because no thorough surveys had been con- ment some time between 1723 and AD ducted on the islands of this large saline lake legitnegit formed as a result of a series of eruptions mono basin ecosys- and because ofpossible changes in local ecology beginning about 200 AD 1987 service associated with falling lake levels from water tem study committee US forest diversion for human consumption 1989

94720 and white mountainMonnmountain beseresearchRese arcilareli station 300030 E 1 univeisityotof Californiacalif olnia betbelberkeleykelcy california vitemite dfpiilinditoldepartnnt of 1 oitsliyindand Resresoniekesoiiiceresonreoureonre management university linolineneslicetstidsted bishop californiaCali fonnalonnafonua 93514 T pe arizonaanona 85287 2 stv4 and hangerange eperiinentexperiinent station foiestforest sciences lab tempe USDAUSDAIforestOCSIoost seaservicesem kockxrocky mountainMonnmoun tain forestfoiestainl 92109 as road san diego california IIiI1 hibbsliis marinemaline researelireseaiclibeseReseareliarellareil centelcenter 1700 south slioicssliores 3sS a world reseaiclires rell institute ililiiills 85721 1 ot anonaamona tucson aronaarizonaanona 4 present addless sclioolschool ofot renewable natinalnatnialnathinal Kesoresourcesnices university ofarizonatucson

328 199211992 MONO LAKE ISLANDS vertebrates 329

N

legitnegit 1

0 marsh

pachapaoha I1

0 2 4 km 1 mono fig lake and the two major islands paohapacha and small location of legitnegit islets are not shown the boxes indicate the general the 1991 study plots stippling indicates the marsh on paohapacha redrawn from various USU S forest service maps

beginning in 1941 the major streams enter- hoffmann unpublished report captured ing mono lake were diverted and no their water small mammals on pachapaoha in one night of effort was transported to los angeles california this J H harris personal communication diversion lowered the lake cap- level about 15 m by tured deer mice peromyscus maniculatus 1981 to the modern on historic low and also legitnegit during several days of in the decreased the trapping early lake volume by about 50 mono 1980s one of us JRJ has made repeated basin ecosystem visits study committee 1987 annually to the islands since 1980 making visual botkin al et 1988 although diversions have observations but not trapping all other been halted a continuing drought through at accounts of the islands mammal fauna are from least 1992 that began in 1986 has prevented any recollections ofearly settlers and local residents significant rise in the lake level eg fletcher 1987 personal communication pachapaoha and legitnegit islands are located along an with JRJ axis running perpendicular from the northern shore of mono lake with pachapaoha the farthest STUDY AREAS away and about 1 km from legitnegit since its formation legitnegit has been separated from the mainland by 0 to 3 km mono basin ecosys- pachapaoha island can be divided into two general tem vegetative zones a small study committee 1987 figs 131.3 and 616.1gi about 2 ha spring fed marsh however no mainland connection with legitnegit along the southeastern shore and the existed since the formation of pachapaoha until the remaining nonmarshnonmarsh vegetation vegetation in the marsh late 1970s the next most land is composed of rush juncus fusus recent bridge bullrush ef apparently occurred about 500 years before scirpus americanusametlcanusamericanosamericanus saltgrasssaltgrass dis- tichlis spicataspicatespicata foxtail present during our study in 1990 91 negit hordeumjubatum and legit bassia bassia hyssopifolia Non was separated from the mainland by several nonmarshmarsh areas are dominated by greasewood hundred meters of mudmudflatsflats and a few meters of sarcobatus ver shallow micumiculatuslatus and hopsagehousage craylagrayiagrabia spinosaspinosci water this area is referred to herein as sage- brush artemisia dentatatritridentatatridentate is present but the land bridge grasses rare and herbaceous plants are scarce and we know ofonly two previous small mammal concentrated in the marsh and one small about trapping efforts on the islands in 1975 W M 0.303 03 ha grassland site located upslope about 300 m 330 GREAT BASIN naturalist volume 52 from the marsh the grassland area is domi- land populations of deer mice were compared nated by exotic cheatgrasscheatgrass bromus tectoriumtectorumtectorum using ttestsnests zar 1984126 131 legitnegit island lacks any marsh vegetation and has in 1991 within each plot on pachapaoha described no permanent freshwater the upland is similar above 18 large sherman live traps were placed to pachapaoha except for more cover by sagebrush at 10lom m spacings 1 I1 row of 6 traps along each dominant vegetation on the mainland plots is long axis of a plot each plot was trapped for a sagebrush rabbitbrushrabbitbrush chrysothamnus nause total of 54 trap nights and days ie traps were 0susobusosus bitterbitterbrushbrush purshia hitrithitrndentata and scat- left open constantly for 3 days and checked both tered individuals of greasewood curlleafcurlcurlleanleaf during the morning and in late afternoon traps mahogany Cercocercocarpuscarpus ledifoliusledifolius and desert were baited and animals handled as in 1990 peach prunus andersoandersoniinii vegetation in the mainland and pachapaoha traps were run 7 may 24 basin was detailed by burch et al 1977 soils june trap lines were run on legitnegit 4 5 august are a loose mixture of sand gravel ash and silt as described for 1990 data are reported here as loeffler 1977 the number of new individuals ie excluding in 1990 trap lines were established to deter- recaptures captured per 100 trap nights we mine species composition and approximate dis- assume that this measure of capture success is tributions of small mammals on pachapaoha and an adequate index of actual population abun- legitnegit specific trap locations were based on dance indices of abundance were compared ease of boat landing and proximity to the next using chi square goodness offit zar 198440 43 nearest trapping location adjacent trap lines surveys were at least 200 m apart other in 1991 we systematically established 10 during 1991 one 42421 L igal1 gal can was fixed study plots 50 X 20 m on pachapaoha island placed near the center of each trapping plot and 5 on the adjacent mainland to compare cans were placed on all mainland plots and on mammals on the island and mainland island six pachapaoha island plots each was coveredadthcovered with a plots were placed in the marsh 3 plots and dry wooden board raised 2 3 cm above the can shrub vegetation 7 plots all mainland plots traps were run 4 17 days three additional were located to the north and northeast of black traps were placed in the marsh on the southeast point on the northwest shore of mono lake side of pachapaoha island this being the most likely this location was selected because its vegeta- location for shrews soricidae thus six traps tion resembles the dominant vegetation on were placed in the marsh all mainland pitfalls pachapaoha island and represents a likely source for were opened 9 12 june island traps were terrestrial animals opened 7 mayamay 4 june A im2ima1 m2ma area in an open location near the METHODS center of each plot was selected to determine the presence of medium to larger sized mam- small mammal live trapping mals travelinatraveling across the plot the soil in a track plot was smoothed by hand and moistened with all traps used during this study were largelarprelampre wwaterter fine grained sand or soil was added as 76767.6 x 898.9 x 22922.9 cm 3 x 353.5 x 9 inch needed A can of chicken flavored cat food was sherman live traps in 1990 trapping was done secured at the center of each track plot each on pachapaoha island on 27 29 april and 23 25 checked daily for three days for evi- august negit island on 27 29 april and the plot was legit dence wildlife use one halfofthe studyplots mainland on 4 7 september trap spacings of 47 pachapaoha and three mainland study plots were ranged from 10 to 20 m and were based onn on used availability of vegetative cover traps were hour baited with rolledrolledoatsoatskats and peanut butter and time constraint surveys of one person each conducted in all study plots checked each for 1 3 days depending duration were morning and general weather conditions and thus access to the the species date time location upon each observation were islands mainland trapping in 1990 was vegetation type for restricted to a marsh on the northern shore of recorded the lake captures were identified to species museum records sex and age and were measured marked and released at the trap location measurements we obtained records for all vertebrates col- between sexes and between island and main lected in mono basin from the los angeles 199211992 MONO LAKE ISLANDS vertebrates 331

tableTABLF 1 index of abundance no100no loo100 trap nights for small mammals captured on study plots on paohapacha island n 10 a1 plots and adjacent mainland n 5 plots and on legitnegit island trap lines mono basin california 1991

paohapacha island trap nights total total total legitnegit total species marsh nonnonmarshmarsh island island mainland 108 324 432 120 342 peromyscus maniculatusmamculatus 176 130 141 625 64 male 83 90 88 325 41 female 93 40 53 300 23 microtus nwntanustrwntanus 56 09 21 03 perognathus parasparms 67 dipodomys panamintinuspanammtmuspanamintinus 53 D micronsmicrops 18 peromyscus bochiboyhiboylii 03 Eutaeutamiusmius minimus 03 Sperawsperawphilusspermophtlusspermophilusphilus beecheyibeecheyi 03 total 232 139 162 625 213

il ciuclucin squaqu tre analysism ilysisclysis illallaliailulluil compuisonssons between totaltnttot il marshmilmitmut sh mdand total nonnonmarshjonmnonmmarshirsharsh on paohapachaP oh 1 P 05 between paohapachaP id d totaltnttot il islandisi ind indand legitnegit islandisilsi iniind P 001 between totaltoitot paohalmd ahwli OL naohpaoh 1 ind total m imluidmainland P 05 indand between total mainlandm uni ind indandund legitnegit P 01 county museum of natural history small mammals on pachapaoha was similar to that on LACMNH and the museum of vertebrate the mainland but on legitnegit it was almost three zoology university of california berkeley times greater than that on pachapaoha PF ooi001001.001 or MVZ although no records were available for the mainland P oi0101.01 table I1 the islands data from the basin were summa- abundance ofdeer mice approximately dou- rized to supplement published accounts of bled P oi0101.01 on pachapaoha between april early mainland vertebrate surveys voucher speci- breeding and august end of breeding 1990 mens were deposited at the MVZ subadult males accounted for 67 of this increase table 2 while subadult females RESULTS accounted for only 6 total male and female abundance was about equal in april the small mammal trapping number of males caught increased by 63 and females only by 35 in august although the and montane volesvoiess only deer mice solevolevoie difference was not significant P 1.1 pachapaoha microtus montanus were captured on male and female abundances of deer mice island most voles were captured in the marsh were similar on negit in april 1990 no compa- also slightly abundant legit deer mice were more rable august data were available total abun- there than in dry shrub plots but these differ- dance on legitnegit in april was 48 higher P ences were not significant P 1.1 the sex ratio 05os05.05 than that on pachapaoha table 2 of deer mice was skewed toward males in the male deer mice from pachapaoha weighed dry shrub but was about even in the marsh adult less and had significantly shorter 1 significantly table tails feet and tailbody length ratios than main- only deer mice were captured on negit legit land animals body and ear lengths were not island mouse abundance was about 454.5 times different table 3 adult females from pachapaoha higher on than on pachapaoha P 0505.05 and legitnegit were significantly less heavy than mainland ani- sex ratios were about even table 1 mals and had smaller but not significantly dif- eight species of small mammals were cap- ferent average measurements for other tured on the mainland plots in 1991 great characters comparisons with negit mice were basin pocket mice perognathus porous deer legit perdus not possible because an insufficient number of mice and panamint kangaroo rats dipodomys animals were measured panamintinuspanamint inus had similar relative abundances and were the only species with abundances 5 other surveys individualsindividuals100100 trap nights except for the great basin kangaroo rat dipodomys micronsmicropsmicrops ISLANDS the six pitfalls in the pachapaoha all species were captured rarely all at 030.3 ani marsh were run for 13 days 78 trap days and mals100mals 100 trap nights overall abundance of captured 777.7 voles100voles 100 pitfall days the three 332 GREAT BASIN naturalist volume 52

ta131&iiiiriii 1 2 abundance nolnoInolootiapnoi0000 trap nights of peroinyscuspcromtscus tails were also seen adjacent to the plots on nianiculahismiflftu tdrtii on daoliapaohapaolia and legitnegitncgit islands mono lake cali- fornia 1990 several occasions daoliapaoliapaoha island legitnegit island discussion apil august andlapdl4pnl only two species of small mammal deer gralgrai nights 290 160 74 mouse and montane vole were trapped on malemaiemalc pachapaoha on adult 76 88 135 and one species deer mouse legitnegit subadult 10 15011ISO1501 1 00 compared with eight species including deer juvenile 07 13 00 mice and montane voles on the adjacent total 93 25011250 135 mainland visual and track surveys found the felnfeinfecialefenialetielietle cottontail coyote adultaditit 59 81 108 jackrabbitjackrabbit and on legitnegit subadult 38 50 41 island and the mainland all but the cottontail juvenileuvcnile 00 19 00 were evident on pachapaoha in contrast at least 20 total 9793 150 149 species of small mamimammalsnalsmalsmais have been observed overall 190 400 365 around the shores of mono lake harris 1982 cidli squareaqilsqil ilalinilolinlysismmlysismm lysis P1 01 P 001 llathallaohaP iokiio1i 1 apilapii vs angustnn t andindplolildaoliapaolia 1984 in addition weasels mustela sppapp apnlvsapril vs nit aprilapi I1 badger taxidea taxus bobcat lynx rufus mountain lion felisfeilseelisfellsfelis concolor black bear ursus and mule odocoileus pitfalls inm the dry shrub were run for 17 days 51 americanusamericanosameric anus deer hemionus around mono pitfall days no animals were captured track occur lake harris 1982 furthermore lion remains have been plots were run foiformol 3 days on pachapaoha resulting in reported from an islet near and from the a total effort of 15 track plot days one set of ported legitnegit vicinity of the negit mainland land bridge caniscams latranslalatranotrans tracks wasfoundwas foundhoundmound on a plot legit coyote mono lake unpublished observa- the marsh and one set ofunidentified rodent committee in tion presence of montane voles on paohapacha tiackstracks likely deer mouse was found on a dry the was associateda4thassociated with the marsh and grass vegeta- shrub plot coyote tracks and scat were seen tion that is absent on negit current lake both islands were especiallyy legit the throughout they especiaespecial level has allowed the pachapaoha marsh to expand end of pachapaoha includ- evident on the southeast onto an exposed lake shelf thus increasing marsh black tailed jacijackiabbitsjackjackrabbitsrabbits ing the lepus potential vole habitat the environment may be califorcahformcuscalifornicusnicus weiewere uncommon but were seen unsuitable on the islands for persistence of the tails occasionally on both islands Cottoncottontails larger carnivorescarnivores and deer but appears suitable sylvilagusSylvilagus sppapp were seen rarely on legitnegit butut on caobapaohapaoba because of water and rodents for were not evident on pachapaoha rabbit pellets were weasels and possibly badgers conspicuous on the islands indicating that the animals can colonize islands by swimming populations had been greater at a previous time rafting using ice bridges being inadvertent pas- no heipshelpsheaps were observed on either island during sengers on waterwatercraftcraft calhoun and greenbaum any survey or in any yearly island visit by JRJ 1991 intentional or unintentional releases or since 1980 scattered individuals of sagebrush by flyingg all but flying may apply to the animals lizard scelopomssceloponts gracious were seen whilei e discussed herein the lack of historic quantita- walking on and near the mainland study plots tive data however prevents determination of MAINLAND the five pitfalls were run for the methods and dates of arrival of anianimalsnials 4 days 20 pitfall days four sagebrush lizards on the mono lake islands however hoffman weiewelewere captured 20 hardshards100lizards100lizardsloo100 pitfall days unpublished report set 76 sherman traps for track plots were run for a total of3 days on threethithl ee one night 24 may in 1974 in various locations study plots with one set of black tailed jackrab including in and around the same marsh and bit two sets of kangaroo rat species unknown grassland areas we trapped he caught no ani- and one set of unidentified small rodent tracks mals but did locate a rodent faexfaed although observed thus there were four separate ailiaillariiani- hoffman s efforts were minimal his dataitdata at least mals in 9 track plotpiot days coyote tracks were indicate the presence of rodents prior to 1974 seen on the plots and coyotes weiewere heard calling although the earliest historic accounts of adjacent to plots numerous rabbit and kanga- local native americans date to the early 1860s roo rat tracks were present on all plots cotton jehl et al 1984 1988 various peoples are 1992 MONO LAKE ISLANDS vertebrates 333

tableTABLFTABLY 3 characteristics ofadultofadult peroinystwsPC omysws maniculatusmamculatus captured on paohapacha island mono lake and adjacentaqjacent mainland during 1990 and 1991

adult male adult female

paohapacha mainland pachapaoliapaoha mainland

characteristic X SD X siosloSDSID X SD X SD

massmassg1g 172 211 188 129 181 229 199 2802.802 SO body length mm 814 513 814 1872 87 79479.479 4 6506 50 81181.1siisilsli81 1 318 tail length mm 645 432 671 534 664 685 691 634 foot minmm 200 107 209 095 200 091 206 108 ear wrimmm 174 108 174 145 174 104 179 138 tailbodyTail body 079 006 082 007 084 007 085 007

samplesizesampieSampleS imekimpk sizesiwesie 50 mdividufildiidllal ilsuis eeacheuchh areai e i except toitolfolfor inassmgnassiss 1 sit sarnpleS unaltunplt sizesiwelicllesic 15 individualsll11 eachL tellteil IILIareawrea except fortor massm iss iai1 xcludesexcludestxcludes phegnpregnantpregn nt funluntunfeinalesreinalesfelfeinalesllesliesiles naohpaohP ioitoitolaoi i re 12 mainlandin iinlbinl ind H 1113 P 05 P 01 ftestteftesttet

thought to have visited the basin for a much does rafting due to flooding events confound- longer period fiefietcherfletcherFletehermeher 1987 western ing the present situation is the land bridge or immigrants began making trips to the islands by near land bridge movement across the land the 1860s jehljem etetalal 1984198419881988 fiefletcherfietcherFleteher 1987 bridge to legitnegit followed by swimming or raft- A chicken callusgallus gallus and domestic lago- ing to pachapaoha is likely more probable now than morph ranch was established on pachapaoha in the historically late 1870s a domestic goat capra sp ranch the absence of lizards on the islands is per- was initiated in the 1890s fiepiefletcherFlepletcherpietcherteherweher 1987 and plexingplexing however as there appears to be ample a mineral salts and health spa venture was habitat on the islands and species on the main- attempted in the 1940s lagomorphslagomorpha raised land are potentially good colonizers gensusensu case commercially were apparently european hares 1975 1983 however mainmalnmainlandlaird populations lepus sp but there is no evidence that these are small as the elevation of the mono basin is hares remained on daoliapaohapaolia after the early 1920s at the upper end ofthe normal range for reptiles when the commercial operation ceased A few in the sierra nevada summarized from storer goats survived on pachapaoha until at least 1975 and usinger 1968 tiletllethereforeTiierefore their chance of Hofhoffmanfinanhinan unpublished report but were extir- arrival and persistence is low pated by 1980 snakes pituophis melanoinelanoleucusmelanoleucusleucus and thus human movements onto the islands thamnophis elegansdeleganselegans and amphibians bufo were frequent and rodents such as deer mice boreas hyla regilla scaphiopus hammondhammondiiii S and voles could have been inadvertently trans- interintermontanusmontanus are found around mono lake ported in the grain hay and other items taken MVZ specimens personal observation but to support activities on the islands we do not they are scarce locally personal observation know if native lagomorphslagomorpha were tranportedtrantransportedported to there are no historic records of snakes or the islands by humans amphibians on either island and we saw no there is debate in the literature over the evidence ofeither during our visits As discussed abilities of peromyscus microtus and other above for lizards it appears that the chance of small mammals to colonize islands by swimming arrival and persistence of snakes and amphibi- or rafting because they are not well adapted for ans is low exposure to water redfield 1976 calhoun and the mono lake islands parallel other islands greenbaum 1991 peltonen and hanski 1991 in having a greater population abundance espe- we have no direct way of quantifying the rela- cially legitnegit and a simple species composition tive probabilities of inadvertent human trans- relative to the mainland larger relative abun- port versus rafting however the known and dances may be because few predators are pres- frequent history ofhuman visitation and habita- ent and the lack of noraviannonaviannonavian food competitors tion for commercial purposes during this cen- as has been postulated for other island rodent tury results in a higher frequency of occurrence populations eg halpin and sullivan 1978 and less harsh means of possible transport than the few rodent species absence oflizards and 334 GREAT BASIN naturalist volume 52

reduced birdlaird species richness hall et al in CASFCASEgasegage T J 1975 species numbers density compensation preparation on the islands may result in density and colonizing ability of lizards on islands in the gulf of california 56 3 18 gensu al ecology compensation sensu macarthur et 1972 1983 niche overlap and the assembly of island case 1975 by the islands peromyscus popula- lizard communities oikosbikos 41 427433427 433 tions pletcherFLPICHFRFLETCHER T C 1987 paiute prospector pioneer a his- in contrast to island tory of the bodie mono lake area in the nineteenth biogeographic theory press redfield 1976 sullivan 1977 deer mice are century artemisia lee vining california 123 appp HALPIN Z T ANDAN D T P SULLIVAN 1978 social interactions smaller on the islands than on the mainland in island populations of the deer mouse peromyscus although a founder effect gensusensu kilpatrick maniculatusmamculatus journal of mammalogy 59 395 401 1981 calhoun and greenbaum 1991 could HARRIS J H 1982 mammals ofthe mono lake tioga pass have resulted in smaller individuals on the region kutsavi books lee vining california 55 appp 1984 an islands experimental analysis of desert rodent than on the mainland there is likely foraging ecology ecology 65 1579 1584 some combination of ecological factors on the JEIIL J R litJR111 D E BABB AND D M POWFR 1984 mono lake islands that has either resulted in history of the california gull colony at mono lake maintenance of small body size or has directed california colonial waterbirdsWaterbirds 7 94 104 1988 on the of data with selection toward smaller body size study interpretation historical in our reference to the california gull colony at mono lake the sex ratio of deer mice appears to be male california colonial waterbirdsWaterbirds 11 322 327 biased although more intensive trapping both kilpatrick C W 1981 genetic structure of insular pop- within and between years would be necessary ulatulationsions pages 28 59 in M H smith and J joule eds for confirmation because of mammalian population genetics university ofgeorgia potential trapping press athens biases associated with dispersing young males loefflerlo10LOFFFLFRF FF LE R R M 1977 geology and hydrology pages 663838 in D W winkler ed an ecological study of mono acknowledgments lake california university of california institute of ecology publication no 12 davis california maca R paul macamilurrtiiub HHANDEAND E 0 WILSON 1967 the theory we thank aigner jean block martin of island biogeography princeton university press morton and paul stapp for assistance with princeton new jersey 203 appp fieldwork edward beedy for helping with study macartiiubmacamiiub R HHJJ M DIAMOND AND J R karnKARRKABB design john harris and several anonymous ref- 1972 density compensation in island faunalfaunas ecology 5333053 330 342 erees for reviewing earlier drafts and lorraine MMONOONO BASIN ecosystemE OSYSTFM STUDY COMMITTEECOMMITTFF 1987 the merkle for manuscript preparation data col- mono basin ecosystem effects of changing lake level lection was supported by consultantsubconsultantsub contract national academy press washington DCD C 272 appp to jones and stokes associates consultant to PFLTONFN A AND I1 HANSKI 1991 patterns of island and power occupancy explained by colonization and extinction los angeles department of water rates in shrews ecology 72 1698 1708 and california state water resources control RFDFIFLDREDFIELD J A 1976 distribution abundance size and board and the white mountain research sta- genetic variation of peromyscus maniculatusmamculatus on the tion university of california los angeles gulf islands of british columbia canadian journal of zoology 54 463 474 storerSIORFRSTOREB T IL1 AND R L USINGERUSINGFR 1968 sierra nevada literature CITED natural history university of california press berke- ley 374 appp bolkinBOIKINBOTKIN DDWW S broeckerBROFCKFBBKOECKPK L G EVERETT J SISHAPIROIAPIRO SULLIVAN T P 1977 demography and dispersal in island AND J A WIENSWIFNS 1988 the future of mono lake and mainland populations ofthe deer mouse peromys- university of california water resources center cus maniculatusmamculatus ecology 58 964 978 report no 68 berkeley california 29 appp USU S FOREST SERVICESFRVICF 1989 mono basin national forest BURCII J BBJJ ROBBINSBOBBINS AND T wainwright 1977 scenic area comprehensive management plan USDA pages 114 142 in D W winkler ed an ecological forestfoiest service inyo national forest bishop califor- study of mono lake california university of califor- nia 95 appp nia institute of ecology publication no 12 davis zanZARZAB J H 1984 Biostatistical analysis and2nd edition pren california tice hallhalihailhafl englewood cliffs new jersey 718 appp CALHOUN S WwandlwandiAND 1 F GREENBAUMGRFFNBAUM 1991 evolution- ary implications of genetic variation among insular populations of peromyscus maniculatusmamculatus and peromys- received 19 february 1992 cus oreasareas journal of mammalogy 72 248 262 accepted I1 october 1992 great basin naturalist 524 appppp335335 343

VASCULAR FLORA OF KANE LAKE CIRQUE PIONEER MOUNTAINS IDAHO

robert K moseley1andbandand susan bernatas 12

ABSTRACT kane lake cirque lies in the western pioneer mountains of south central idaho an inventory of the high elevation flolafloiaflora of the cirque revealed the presence of 180 vascular taxa representing 95 genera and 30 families five alpine taxa are here documented from idaho for the first time carex incurvtfomwincurvifonnis mack drabafladnizensisdraba fladnizensis silfenwilfen potentilla aveanivmveaa L ranunculus gerdusgehdusgelidus kar & kir and ranunculus pygtnaeuspygnweus wahlenewahlenb kane lake cirque also contains populations of four additional alpine taxa considered to be of conservation concern in idaho erigeronengeron humilis graham parnassiaPamassta kotzekotzebueikotzebuezbuelbuei cham Sarisamisamlsaxifragafraga adscendentadscendens L and saxifraga cemuacernuacergua L

key words idaho pioneer mountains kane lake cirque alpine vascularfloravascular floraglora staterecordsstate records rareranerarefloraflora

studies of alpine flora have been numerous autionbution and abundance of rare plants and habi- throughout the north american cordillera but tats in the basin for future management only recently have investigations of this kind been undertaken in idaho floristic studies ini- STUDY AREA tiated by douglass henderson of the university of idaho herbarium in the mid 1970s were the pioneer mountains rise abruptly from first to systematically explore the alpine zone of the the northern edge of the snake plain in idaho through numerous collections he and river central idaho between the big lost and big coworkersco documented idaho s alpine flora to workers wood rivers these mountains form a large be in many respects henderson 1978 unique complex block about 60 km long and 50 km al 1981 bruns- brunsfeld 1981 henderson et wide oriented northwest to southeast topogra- al al 1983 lackshewitzLacklackschewitzshewitz feld et 1983 caicco et phy varies from sharp horns serrate ridges and 1985 et al 1983 hartman and constance broad upland surfaces in the alpine zone to all took the nearly these investigations place in steep sided valleys and rounded ridges in the large basin and range like massifsmassios in east cen- foothills elevations range from 1900 m in the tral idaho with few extending into the western valleys on the western slope to 3658 m at the pioneer mountains of south central idaho summit of hyndman peak rare plant inventories initiated by the US the pioneer mountains are composed of forest service were the first to point out the tertiary challis Volcanivolcanicscs consisting of inter- phytogeographic importance of the cirquescinques in I1 bedded lava and tuffaceous units which lie the western pioneer mountains in general and unconformablyuncomformably over a core of precambrian kane lake cirque in particular caicco and metamorphic and paleozoic sedimentary units henderson 1981 brunsfeld et al 1983 bar- during the formation of the cretaceous idaho bara erttefsertters collections in 1977 and our collections batholith small satellite intrusive bodies were in 1987 further highlighted the significance of emplacedemplaced in the western pioneer mountains kane lake cirque because ofincreasing recre- tertiary and quaternary block faulting is ational use of the kane lake area sensitivity of believed to be the cause ofthe subsequent upliftcliftplift the habitats and preliminary nature of the flo- and present relief dover 1981 the gelmorgeomor ristic inventory we undertook this study in phophologiclogic setting has been greatly influenced by cooperation with the challis national forest to quaternary glacial and fluvial activity most provide them with adequate data on the distri streams in glaciated valleys are underunderfitunderwitfit and

1 comeivtionconsemition data center idaoidaho department of fish and camegame box25bosbox 25 boisebowe idaho 83707 2presentpieient address science application international corporation 405 S ath8th suite 201 boise idaho 83702

335 336 GREAT BASIN naturalist volume 52 uplands display classic alpine type glaciated can be attributed to several factors including features including cols aretesareces homshorns and the north facing orientation of the cirque a cirquescinques evenson et al 1982 massive headwall on the south and high peaks the pioneer mountains lie in a transition on the east south and west sides of the basin zone between the maritime climate of northern these features contribute collectively to a heavy and western idaho and the continental climate snow accumulaccumulationation in the winter and its reten- of southeastern idaho and are affected by two tion throughout the summer late lying snow basic storm patterns from november through and an impermeable substrate augmented by march most precipitation comes from low summer thunderthundershowersshowers appear to provide altitude cyclonic storms that move eastward from plentiful moisture to nearly all habitats through- the pacific ocean during may and june most out the growing season except the south facing precipitation results from high altitude convec- slopes north of kane lake tional storms moving northward from the gulf habitats in the cirque can be divided into of mexico and california coast the combina- two distinct groups subalpine and alpine sub- tion of maritime and continental influence cre- alpine vegetation is restricted to areas ates two wet seasons winter and late spring immediately adjacent to kane lake and gener- respectively caicco 1983 no climatic data are ally does not exceed 2850 m alpine habitats pioneer available from high elevations in the cover most ofthe area and are generally sparsely mountains however moseley 1985 estimates vegetated although small areas with continuous annual at 2835 in the mean precipitation in vegetative cover occur along streams and rivu- southern of the to be 813 mm part range lets throughout the basin and contain much of the mountainous regions of the throughout the plant species diversity of the alpine zone world the altitude of upper tretreelinefreelineeline has long plant associations of the study area are not been observed to coincide with the 10 C iso- included published classifications therm of the warmest month griggs 1937 in vegetation of the and are subjectively characterized daubenmire 1954wardle1954 wardlewardie 1974 extrapolation region below oftemperatures from valley stations in the vicin- ity to timberline 3000 in using an adiabatic subalpine communities lapse rate of 0620.62ogg clooc100 inm baker 1944 sub- stantiates this observation for the pioneer coniferous woodland open stands of mountains pinus albicaulisalbicautisalbialblcanliscauliscautiscantis engelm with lesser amounts of abies lasiocarpalasiocampa hook nutt and picea engelmanniaengelmanniiengelmannii parry occupy the level bench north description OF KANE LAKE CIRQUE of kane lake the relatively xeric understory is characterized byvaccinium scopariusscoparium leiberg kane lake cirque encompasses approxi- poa nervosenervosa hook vasey and senecio the mately 567 ha at the head of kane creek in streptanthifolius greene pioneer mountains 21 kmknikhi northeast of western meadows interspersed in low ketchum county idaho upland custer 4344n within the coniferous woodland are boise meridian the lying areas 1146w t5ntan r20e meadows dominated by gramingraminoidsoids and forbs is characterized by permanent snow cirque characteristic of these sites include fields glacially scoured bedrock gneiss and species morainal festuca idahoensis elmer danthonia intenneintermeintermew quartz diorite and unstable talus and greene carex although several small ponds are seatscat- dia vasey erigeron simplex deposits potentilla tered throughout the basin 53 ha kane lake is micropteramlcroptera mack and diversifoliadiversifolia the only large body of water elevations of the lehm isolated study area range from 2800 in to 3648 in the tree islands and krumfolzkrummolzkrummolz tree only appreciable soil development is in deposi- islands consisting of small upright abies tional areas such as along streams and rivulets lasiocarpalasiocampa with an understory dominated by around ponds and lakes and in the coniferous phyllodoce sppapp occur as high as 3050 in on woodland on the north side of kane lake benches south ofthe lake these relatively moist vegetation in the cirque reflects a moister sites are surrounded by bedrock or alpine regime than has been noted at high elevations meadow communities areas of krummholz are elsewhere in idaho brunsfeld 1981 caicco rare in the study area small isolated patches 1983 moseley 1985 this mesic environment consisting of low growing A lasiocarpalasiocampa and 199219921 KANE LAKE CIRQUE VASCULAR FLORA 337 pinus albicaulisalbicaulis occur as high as 3230 m on the METHODS steep south facing slopes north of kane lake lakeside meadows surrounding kane the checklist is based on 263 collections lake and radiating out along inlet and outlet made mostly by the authors in july and august streams are meadows that have soil saturated to 1987 and july 1991 other collectors include the surface and are high in organic matter barbara ertter who visited kane lake in july carex scopulorum holm is dominant here along 1977 and steven caicco who collected in the with scattered forbs such as erigeron per- cirque during july 1981 and august 1982 A egrinus pursh greene senecio cymba nearly complete set of specimens is deposited at larilarioidesoides buekbuck and Cengentianagentianafianaflana lycosacaiticalifcalycosaca cosa griseegriseb the university of idaho herbarium ID with low shrubs including salix planiplanifoliafolia pursh duplicates distributed widely ertterertterss collec- and ledum glandulosum nutt occur occasion- tions are deposited at the albertson college of ally in these meadows idaho CIC collections thought to be new records for alpine communities idaho were confirmed by experts in a particular taxon andor from a search of up to 59 local meadows this mesic community is limited regional and national herbaria moseley 1989 in extent and generallyrenerallyraily occurs in isolated range extension data for these state record taxa patches around seeps or as along stringers were determined from herbarium records and streams and rivulets deschampsia cespitosecespitosa from the atlases and data bases maintained by L is by far the dominant here beaucbeauv species the idaho conservation data center and mon- with a high diversity of forbs and other tana natural heritageb program on the location gramingraminoidsoids occurring in low cover distribution numbers and condition of rare cliffs and ledges this is the most common plant populations in their respective states community in the cirque most cliffs and ledges jenkins 1986 the idaho conservation data are north facing and wet to mesic with draba center data base was also consulted concerning lonchocarpa redbrydb being the most constant spe- the current distribution of additional rare spe- cies alonga4thalong with several species ofsaxifraga the cies in idaho xeric counterparts occur only on the south facing slopes northeast of kane and have lake RESULTS AND few vascular plants discussion and scree common at upper eleva- talus vascular flora of kane lake cirque con- tions in the cirque this relatively mesic commu- the sists of 180 species representing 95 genera in 30 is characterized by a of nity unique suite species families of pteridophytes gymnospermsgymnospermy and able to withstand constantly shifting substrates angioangiospermssperms of these 53 species 29 are species characteristic of material greater than restricted to subalpine communities in the 5 talus include cm in diameter eulseahulsea alaidaalgida cirque while 58 species 33 are restricted to gray & while and senecio fremonfiremonfremontiifremontiafiremontiitiitil T G alpine habitats the remaining 69 species 38 saxifraga cemuacamuacernuacergua L luzula spicataspicate L DC transcend the subalpine alpine boundary and and androsace septentrioseptentfionalisseptentrionalisnallsnalisnails L characterize occur in both types of communitiesofcommunities our collec- small diameter material scree tions of five species from the study area repre- fellfielddellfieldFellfield dry fellfieldfelldellfieldfield habitats are rare sent their first documented occurrence in occurring only in small pockets on bedrock slabs idaho in addition four other arctic alpine spe- on the cirque floor east of kane lake species cies are known from idaho from only a few typical of this poorly developed community occurrences and are considered rare in the state include potentilla brevifoliabnevifoliabrevifolia nutt juncus moseley and groves 1990 only one alien drummonddrummondiidrumnwndiiii E meyer and sibbaldiaSibbaldia pro taxon taraxacum officinaofficinalele weber was found cumbentcumbens L carex elynoidesely noides holm also occurs in the study area in this community but does not develop into the extensive turfscurfs that are found elsewhere in cen- taxa new to idaho tral idaho caicco 1983 all ridges surrounding carex incurviformis mack this species the basin typical sites for fellfieldfelldellfieldfield communities occurs in two areas of the north american cor- elsewhere in the state are sharp aretesareces with no dilleradillera var incurviformis known from the well developed vegetation rocky mountains of british columbia alberta 338 GREAT BASIN naturalist volume 52

montana and now idaho and var danaensis the rocky mountain crest to colorado benson stacey hermann occurring in the southern 1948 its presence in the kane lake cirque rocky mountains of colorado and the sierra represents a disjunction of about 200 km south- nevada and white mountains of california J west from the next nearest known populations mastrogiuseppeMastrogiuseppe washington state university in the pioneer mountains beaverhead county personal communication 1991 the popula- montana hitchcock and muhlick 12899 WS tion in kane creek is disjunct south from the ranunculus pygmaeus is relatively common in next closest known population in deer lodge the kane laketakedake cirque occurring in moist county montana by about 260 km lacks- exposed soil along creeks on ledges and slopes chewitz 3938 MONTU lesica and shelly and occasionally in cracks in cliffs 1991 we found one small population in the additional rare species kane lake cirque occurring in a steeply slop- ing meadow on seepy ledges at 3350 m at the erigeron humilis grahm this circumpo- southern end of the cirque lar species was not known from idaho until drabafladnizensisdraba fladnizensis wiffen A widespread henderson et al 1981 reported it from the circumpolar species draba fladnizensis is leahilemhi and lost river ranges eight occur- sparsely distributed in north america from the rences are now known from the state with the arctic south through the rocky mountains to kane lake cirque populations being the only utah and colorado hitchcock 1941 As ones known outside the two ranges mentioned withcarexwithCarex incuroiformisincurviformis the kane lake cirque above unpublished data on file at the idaho population is disjunct south from the next clos- conservation data center boise erigeron est known population in the storm lake area of humilis is relatively common in moist the pintlarpintler range deer lodge county mon- deschampsia cespitosacespitose meadows throughout tana by about 260 km lackschewitz 6120 the lower portion of the cirque MONTU several very small populations occur parnassia koftebucikotzebueikotzebueibuel cham this species on ledges and in rocky areas south of kane lake was also not known from idaho until recently including spray zones ofwaterfalls bare stream when brunsfeld et al 1983 reported it from gravels and on steep rocky slopes near seeps the lost river range and pioneer mountains potentilla nivea L this circumpolar spe- four occurrences are now known from the state cies occurs in arctic and alpine regions of north unpublished data on file at the idaho conser- america being previously known in western vation data center boise it is relatively north america from alaska south along the common on moist ledges and in sloping main crest ofthe rocky mountains to montana deschampsia cespitosacespitose meadows throughout wyoming colorado and utah and east to the lower portion of the cirque nevada hitchcock and cronquist 1973 the saxifraga adscendentadscendens L the north amer- kane lake cirque population is disjunct from ican representative ofthis addewide ranging species the nearest montana populations by perhaps var oregonensis raf breit occurs throughout 280 km A small population of about a dozen the rocky mountains and northern cascade plants was seen in a moist sloping meadow at range hitchcock and cronquist 1973 in the top of the waterfalls south of kane lake at idaho it is known from nine sites in the white 2950 m cloud peaks pioneer mountains and lost ranunculus gelidusgelious kar & kir A north river range unpublished data on file at the american endemic this species is distributed idaho conservation data center boise kane across the arctic southward in the rocky moun- lake cirque populations occur throughout the tains to colorado benson 1948 the very small area on moist scree sand and gravel often population in kane creek cirque represents a along streams disjunction southwestward of about 350 m from saxifraga cernuacergua L seven small popula- the Bearbeartoothtooth plateau stillwater county mon- tions of this circumboreal species are known tana stickney 4 MRC lesica and shelly 1991 from idaho unpublished data on file at the in the study area it occurs in a stringer of idaho conservation data center boise at deschampsia cespitosacespitosecespttosa along the northeastern kane lake cirque it is widely scattered in small tributary of kane lake at about 3170 m populations from moist subalpine ledges north ranunculus pygmaeus wahlenewahlenb this of kane lake at 2800 m to ledges and cracks on buttercup is circumpolar occurring south along the headwall at 3400 m 199211992 KANE LAKE CIRQUE VASCULAR FLORA 339

ANNOTATED CHECKLIST OF jepsbeps common in moist sandy soil in alpine zone 1186 VASCULAR PLANTS 2336 antennaria dimorphadimorpha nutt T & G rare in dry forest opening north of kane lake 2393 the checklist is arranged by division and antennaria microphyllamicrophylla redbrydb common on dry sub- class in agnoliophytamagnoliophytaM then alphabetically by alpine and alpine slopes 2244 family genus and species within these major antennaria umbrinellaumbrinella redbrydb common in dry to moist subalpine and alpine meadows 2308 2312 groupingsaroupings nomenclature generally follows arnica latifolialatifolia bong var gracilis redbrydb cronacronq hitchccckhitchcock and cronquist 1973 exceptions common in deep soil of subalpine and alpine slopes and being salix brunsfeld and johnson 1985 boulder fields 2246 carex incurvifonnisincurviformis and C scopulorum var arnica mollis hook common in moist subalpine and bracteosebracteosa hermann 1970 and eilogonumenogonum lower alpine meadowsandmeadowlandmeadowmeadowssandand boulder fields 117723192343 artemisia michammichauxianamichawmichawcianamichauxianalanarianatianaciana bess uncommon in moist capistratumcapistratum reveal 1989 unless otherwise unstable lockyrocky drainage bottoms subalpine and lower noted the collection numbers are the authors alpine oneszones 2363 artemisia tridentata nutt rare on dry subalpine DIVISION lycopodiophyta slopes north of kane lake 2261 selaginellaceae aster alpigenusalpigenus T & G gray var haydeniihaydenii porter cronq in forest north of kane common crona dry openings in selaginella densa redbrydb in subalpine and lake 2419 alpine zones moist to dry slopes and ledges and stabilized aster follafoliafoliaceusfoliaceousceus var apricus gray common in scree 2245 asterfoliaceus lindl africus moist alpine meadows eastcast of kane lake 1175 DIVISION polypodiophyta aster stenomeresstenomeres gray dry lockyrocky ledges in forest openings north of kane lake 2235 polypodiaceae chaenactis alpina gray jones uncommon in sub- cryptogramma crispaarispa L BR br uncommon in moist alpine and alpine dry sandy scree 2361 subalpine and alpine talus circumboreal 2292 cirsium tweedyi redbrydb petr common in moist cystopteris frafragilisfragulisgilis L beah common among meadows and on ledges in alpine zone 2378 rocks in moist alpine sites circumboreal 2294 erigeron acris L var dabilisdebilis gray common in pellaea breweribrewere DC eat uncommon in subalpine moist sandy soil subalpine and alpine zones 1183 2287 and alpine zones stabilized scree rocky ledges and boulder 2344 fields 2293 erigeron asperuginusasperuginus eat gray dry slopes and woodsia scopulinascopulina DC eat common on rocks in ledges common in subalpine and uncommon in lower subalpine zone 2252 alpine zones 225023502250 23509350 erigeron composituscompocompositoussitus pursh var glabratusglabratus macoun DIVISION PINOPHYTA common on dry subalpine and alpine ledges 2265 cupressaceae erigeron coultericoulterscoulteri porter rare in alpine meadowmeadows along creek east of kane lake 1174 funijunijuniperusperus communiscommoniscommunis L var montana ait rare on dry erigeron humilis graham locally common in moist ledges of lower alpine zone and in krummholz 2355 alpine meadows 2274 2410 caicco 284 erigeron peregrinus pursh greene sspasp pinapmaceaepomaceaeceae ceab calliancallianthemttscallianthemusthemus greene cronacronq var scaposus T & G abies lasiocarpalasiocampa hook nutt common in woodland cronacronq common in rnoistmoist to wet subalpine rneameadowsdows and krurnmholzknimmholz communities 2392 around kane lake 2306 2367 picea engelmanniiengelengelmanniamannii parry common in woodland and erigeron simplex greene common inm subalpine and krummholz communtiescommcommunitiesunties 2391 alpine zones moist meadows and slopes 118822701188 22709270 2338 pinus albialbicauliscaulis engelm common in woodland and caicco 476 ertter 2108 krummholz communities 2263 haplopappus lyallii gray uncommon on dry alpine ledges 2402 DIVISION magnoliophyta haplopappus macronemamacronema gray uncommon on dry CLASS magnoliopsida subalpine knoll within forest north of kane lake not collected apiaceae haplopappus suffruticoussuffruticowssuffruticosus nutt gray uncom- lomatium idahidahoenseoense math & const rare in dis- mon on dry subalpine knoll within forest north of kane turbed micromicrositessites in moist subalpine meadows north of lake not collected kdneldkekane lake 2256 hieracium gracile hook uncommon in dry forest osmorhiza chilenchilensissis H & A uncommon in deep soil openings north of kane lake 2305 of forest understory 2376b huheahulseaeulsea alaidaalgida gray common in alpine talus 2403 microserisMicroseris hutansnutans geyer shultz uncommon asteraceae bip in subalpine meadows north of kane lake 2385 achillea millefoliummillefolium L sspasp lanulanulosalosa nutt piper senecio cymbalarioides buek common in moist sub- var alpicola redbrydb garrett common on dry subalpine alpine and alpine meadows 1173 and alpine slopes 2262 seneciofremontiisenecio fremonfremontiifremontiapremontiitil T & G varfremontii common in agoserisAgoseris aurantiaca hook greene uncommon in alpine talus 2400 subalpine meadows north of kane lake 2387 senecio streptanthifolius greene common in sub- antennaria alpina L gaertn var media greene alpine zone dry slopes and forest understory 2255 340 GREAT BASIN naturalist volume 52

solidago multiradiatamultiradiatemultiradiata ait var scopulorum gray crassulaceae dry rockyi subalpine and alpine ledges 2258 sedum lanceolatumlanceolatum torr var lanceolatumlanceolafumlanceolafumtum Ledeb DC common in alpine taraxacum lyratum ledealedeb common on moist to dry subalpine and alpine slopes and 1185 2289 one moist meadows and slopes liss11852289 ledges2240ledges 2240 taraxacum officinaofficinalele weber alien rare in subalpine meadows northnoinol th of kane lake 2388 ericaceae boiagmaceaeboraginaccaeboraginaceae kalmia microphyllamicrophylla hook heller common in to wet subalpine and alpine meadows 2304 G common along moist mertensiaMertenfiasiasla ciliata torr don ledum glandulosum nutt var glandulosum subalpine and lower alpine rivulets 2268 common in moist subalpine forest and meadows around 2303 bibrassicaceaeasmcaceac kane lake phyllodoce empetriformiseiwpetriformis sw D don common on arabis sp immature and unidentifiable to species moist subalpine and alpine slopes 2300 uncommon inm dry to moist forestfoiest openings north of kane phyllodoce glandulifera hook cov common on lake 2375 moist subalpine and alpine slopes 2302 arabisarabia lemmoniilemmoitiilemmoniinil wats var lemmolemmoniinii common on x phyllodoce interintermediaintermedialmedia hook camp common on dry unstable alpine slopes 231323562313 23562336 moist subalpine and alpine slopes 2301 arabis micromicrophyllaphylla nutt var micromicrophyllaphylla vaccinium scopariusscoparium leiberg common in dry sites common on subalpine ledges and slopes north of kane in understory of forest and krummholz 2237 lake 2248 arabis micromicrophyllamicrophylluphylla nutt var saximontana rollinsRolluis fabaceae uncommon soil ofalpineof alpine onezonewone 2374 in moist astragalus alpinufalpinusaepinus L circumboreal common in unable to identify possibly a new taxon draba sp moistoistolst meadows throughout cirque subalpine and alpine baiebalerare seen only in one small steeply sloping moist meadow raie oneszones 2318 2373 caiccocaicco474474 3353 east of kane lake 2412 at in cosmus rare in cracks of moist circumpolar rare on dis astragalus eu robins raierale draba fladnizeniisfladnizentis silfenwilfen cliff stream alpine zone 2396 of slopes and along near tintuhtunthibedtnibedbed baiebalebare soil miciomiciositesrnicrositessitessltes steepalpinesteep alpine astragalus kentrophyta gray var implexus canby rivulets 1107 common on exposed dry alpine slopes and var lonchocarpa barneby draba lonchocarpa redbrydb ledges 2352 ertter 2102 common on011 moist ledges and slopes Conunon throughout cirque oiloll trifolium lonlongineslongipesgipes nutt var pedunculatum redbrydb alpi ne oneronezonemone 1106 2314 ertter 2106 alpine hitchchitche uncommon in deep soil along subalpine oligospermia var oligospermaoligospermia rare on draba oligospermaoligoiperma hook streamsheambdnkstreambankbank north of kane lake 2380 dry alpine slopes and ledges 2357 2362 ertter 2102 draba paysonpaysoniiii macbrmacar var treleasii schulz gentianaceae hitchchitche uncommon in dry sandyalpinesandy alpine soil 2405 erysimum asperumesperum nutt DC rare in dry subalpinesubalsubatpine frasera speciosaspeciosospeciosa dougl uncommon in dry subalpine talus noinolnorthtiitil of kane lake 2377 talus north of kane lake 2415 snielowikiasmelowskia calycina steph CA mey var amer- gentianaCengentiana calycosa griseegriseb var asepala maguire icana regel & herd drury & rollins uncommonUnLommoncommon on hitchchitche common in moist subalpine and low alpine mead- diydrydaiy exposed alpine slopes 2349 ertter 2107 ows 1172 Cengentianagentiana proprostrataprostratestrata haenke rare seen only in moist caryophyllaceae steeply sloping meadow above ponds east of kane lake alpine zone 2408 arenaria aculeata wats dry sandy slopes common in subalpine and laietaietalerare in alpine onezone 2236 2351 grossulariaceae arenaria concongestsgesta nutt uncommon on dry alpine slopes eastcast of kane lake 1187 ribes cemuumcernuum dougl var inebriantinelnemeinebriansmebriansbrians lindl arenaria obtusiloba redbrydb fern dry exposed hitchchitche uncommon in subalpine and alpine zones dry slopes and ledges ninioncocommon in alpine and uncommon in ledges and boulder fields 2409 subalpine zonewoneone 2354 ribes hendersohendersoniiftendersoniihendersonminiinil hitchc rare and local in dry boul- arenaria rubella wahlenbwahlene JE smith circlrcn der field east of kane lake alpine zone 2416 cliluborealcnmboieal uncommon on moist to dry alpine ledges 2424 ribes lacustrelacustre pers poir uncommon along sub- cerastium berringianum cham & schlecht alpine creek near outlet of kane lakeL ike 2398 common in alpine onezone throughout chenqueque moist slopes ribes montigenum mcclatchie common in boulder meadows and ledges 2321 2413 fields and dry forest understory subalpine zone 2310 sagina sagisaglsaginoideswginoidesnoides L britt circurnborealciicurnboieal uncom- I1 mon in moist alpine meadows 117922601179 2260 lydrophyllaceaeflydropbyllaceaehydrophyllaceae loeklockroek silene dougladouglasiisii hook var dougladouglasiisii dry rocky phaceliaphacella hastatahastaba dougl var alemaaleinaalpina redbrydb cronacroriqcronq subalpine and lower alpine zone ledges uncommon in uncommon in moist to dryalpinedry alpine talus 2406 2364 silene repensrapens pers var auaustraleaustraliestrale hitchchitche & mag onagraceae rarerairal e among i ocksrocks of boulder field east of kane lake caicco 286 epilobium alpinuntalpinunt L var alpialpinumnum common on stellaria lonlongipeslonginesgipes goldie var altoaltocauliscaulis hulten moist unstable subalpine and alpine slopes circumboreal hitchehitchc uncommon in moist sandy sites and scree in alpineinalpine 2333 meadows 11802327 epilobium angustifolium L rare in dry forest open- stellaria umbellateumbellataumbellata turezturczburczburez rare inm wet to moist gravelsgi avelsaveis ing north of kanaskekanaakekane lake 2418 alongalpinealonaionalong alpinegalpine i ivuletsrivulets 232623472326 2347 epilobium glaberrimum barbey varvan fastigiatum 199219921 KANE LAKE CIRQUE VASCULAR FLORA 341

nutt trel uncommon in moist meadow along stream rosaceae east of kane lake alpine 2258 potentilla brevi LoL cilly oenothera andina nutt rare in disturbed micromicrositessites brevifoliafolia nutt loclocally common on dry in dry subalpine meadows north of kane lake 2264 alpine outcrops at 3050 in southwest of kane lake 2399 potentilla diversifoliadiversifolia lehm var diversifoliadiversifolia polemoniaceae common throughout cirque in moist subalpine and alpine meadows 2283230723722283 2307 2372 phlox pulpulvinatepulvinatavinata wherry cronacronq common on dry potentillafruticosapotentilla fruticosefruticosa L pentaphylloidesflotibundapentaphylloides florlflorifloribundabunda exposed alpine slopes 2348 pursh lavelove common on moist ledges and in boulder polemonium viscosumviscosum nutt common throughout fields of subalpine and alpine zones circumboreal 1171 cirque in talus and unstable sites on ledges alpine 2311 2317 ertter 2104 potentilla glanduloseglandulosa lindlindllindi var pseudorupestris redbrydb breit local on dry subalpine ledges 2366 polygonaceae potentilla nivea L circumpolar rare in alpine zone seen only in moist sloping meadow at headbead of waterfall eriogonum caespitosurncaespitosum nutt uncommon on dry south of kane lake 2379 subalpine knoll north of kane lake 2359 rubus idakusidaeusidaews L var gracigracilipeslipes jones common in eriogonum capistratumcapistratum rev var capistratumcapistratum subalpine boulder fields 2414 locally common in subalpine and alpine zones rocky dry locky sibbaldiaSibbaldia procumbentprocumbens L common in alpine and sub- slopes and ledges 2360 alpine zones on moist sandy slopes and ledges cir- eriogonum ovalifolium nutt var deprdepressumessum blank cmcumeumcumborealboalboreal 2288 common in subalpineandsubalpine and alpine zones dry unstable slopes and ledges 2249 salicaceae oxyria digenadigyna L hill circumboreal common oxiria pall throughout cirque on moist lockyrocky slopes alpine zone 2286 salix arcticaarctisca var petraeapetraca andressandrossanddess common polygonum instortoidesbistortoidesbistortoidesoldes pursh common in moist to throughout cirque in moist subalpine and alpine sites 2276 wet subalpine and alpine meadows 2267 2342 2376a 2397 ertter 2100 salix polygonum kellogg ii greene uncommon in dry sp only vegetative specimens obtained but forest openingsopenineseninas 2390 appears to be S eastwoodiae cockerell ex heller S brunsfeld polygonum viviparumviviparum L common in moist alpine university of idaho personal communication meadows 2395 caicco 477 1991 uncommon in wet subalpine meadow adjacent to the north shoiesholeshore of kane lake 2386 portulacaceae salix aivalisnivalis hook var nivalis raierare on moist slopes min subalpine and alpine zones 2337 claytonia megarmegarhizahiza gray parry var megarmegarhizahiza salix planifoliaplanifolia purshpur&h uncommon min subalpine uncommon in alpine talus 2404 meadow west of kane lake 2266 lewisia pygmaeapygmaean gray robins var pygmaeapygmaean salix tweedyi bebb ball raieralerare only one lobustrobust plant common in dry subalpine andalpineand alpine sites 2271 2369 seen at base of small cascade at 2865 in west of kane lake subalpine zone 2421 pi imulaceaeprimuliceaeprimulaceae saxifragaceaesaxifi fagaceaeagaceae androsace septentrioseptentrionalisnalis L common on dry sandy alpine slopes circumboreal 2353 2422 heuchera cylindric a dougl var alpina wats dodecatheon pulchellumpulchellum RA merrill var common in subalpine and lower alpine zones on dry ledges raf and watsoniiwatsowatsonianii tidestromTidwidestromestrom hitchc common in moist subal- outcrops and moderately stabilized scree 2239 pine and alpine meadows 2341 lithophragmaLithophragma bulbiferabulbifera redbrydb L glabragladra nutt uncommon on moist subalpine slopes 2370 ranunculaceae mitella penpentandrapentandriatandra hook uncommon in lower alpine and subalpine zones moist ineameadowsdows and slopes 2330 anemone parviparvifloraflora michamichx rare in moist alpine massiaparnassiaPa fimbriafimbriatafimbriateta common var fimbriafimbriatefimbriatata meadow south of kane lake 2339 locally common in moist subalpine and alpine meadows aquilegia formosa fisch common in moist sloping 1176 caicco 283 meadows subalpine and lower alpine zones 2269 caicco massiaparnassiaPa kotzebueikotzebuei chamchaincharn var kotzekotzebueibuei uncom- 472 mon and local in gently to steeply sloping alpine meadows caltha leptosepalaleptosepala DC var leptosepalaleptosepala common and on ledges 2285228523282328 caicco 280 throughout cirque in alpine and subalpine zones wet mead- saxifraga adscendentadscendens L var oregonensis raf ows along streams and around lakes and ponds 1181 2298 breit uncommon in moist sloping meadows and talus and delphinium depauperatum nutt uncommon in dry along rivulets in alpine zone 2331 caicco 282 subalpine meadows north of kane lake 2257 saxifraga arbutaarguta D don S odontoloma piper ranunculus eschscholtziieschscholtzia schlecht var esch common along streams and rivulets in subalpine and lower scholtzscholtziiii common throughout cirque on moist subalpine alpine zones 1178 2335 and alpine slopes 2284 ertter 2109 saxifraga cernuacergua L uncommon and widely scattered ranunculus geliousgelidus kar & kir rare seen only in in moist scree and sloping meadows and on ledges of alpine moist alpine meadow at about 3170 in along stream east of and subalpine zones circumboreal 2420 kane lake 1182 saxifraga dabilisdebilis engelm common on moist and ranunculus pygmaeus wahlenbwahlene circumpolar protected alpine ledges and slopes 110811092324 caicco locally common in moist to wet sites in alpine zone along 281 ertter 2105 rivulets ledges and cracks on rock face 1110 2315 2346 saxifraga occidentalisoccidentoccidentalistalis wats var occidentoccidentalisoccidentalistalis ranunculus verecundverecundusus robins rare in moist alpine common in moist subalpine and alpine meadows 2242 boulder field at 3050 in southwest of kane lake 2345 22772316232924012277 23162329 2401 volume 52 342 GREAT BASIN naturalist

lihaceaelinaceae saxifragaSaxlsamlsamifragapraga oppositioppositifoliafolia L common on moist alpine cliff faces circumboreal 2358 allium brandebrandegeibrandegeegei wats rare in dry forest opening north of kane lake 2394 scrophulariaceae allium brevistylum wats common in moist steeply and lower alpine zones 2334 castilleja miniataminiate dougl common in moist to wet sloping meadows in subalpine euryeurycarpuscarpus wats rare in dry areas of subalpine and low alpine meadows 2299 calochortus begel var caespitosus greene subalpine meadow north of kane lake 2384 mimulus tilingtilingiiii regel pursh common in moist sloping grant common along alpine streams and rivulets 1169 Zigazigadenuszygadenusdenus elegansdelegans meadows 2340 caicco 478 fenPenPempemtemonpenstemonpentstemontemonstemon procerus dougl varformosusvaa formosusformosus A nels alpine cronacronq common in subilaubilsubalpmeme zone and uncommon in poaceae alpine zone on dry rocky ledgeslagesl3ges 2259 veronica wormskjoldii roem & schult common in agropyron scribnersscribneriscribneri vasey elymus scribneriscribnersscribneri moist to wet alpine and subalpine meadows 1184 2295 vasey jones uncommon on dry unstable alpine and subalpine slopessiopessl s22722272 violaceae agrostis humilis vasey uncommon on moist sandy greene ham alpine ledges 1196119623682368 viola aduncaaldunca sm var bellidifolia harr agrostis variavariabilisbilis rydb uncommon in moist alpine zones on meadowsdows redb common in subalpine and alpine moist inea meadows 1194 and slopes 2290 2371 calamagrostis purpurascens R br uncommon on macloskeymacloskeyii common in viola macloskeymacloskeyii lloyd var dry rocky subalpine and alpine ledges 2365 wet subalpine meadow adjacent to kane lake 2309 danthonia interintermediaintermedialmedia vasey locally common inm sub- alpine meadows north of kane lake 2389 CLASS liliopsida deschampsia cespitosacespitose L beaucbeauv var cespitosecespitosa cyperaceae common throughout cirque in subalpine and alpine moist meadows where it is often dominant circumboreal 1192 carex atrataacrata L var erectaejecta boott rare in moist soil 2325 caicco 481 of boulder field north ofkane lake subalpine ertter 2110 festuca idahoensis elmer var idahoensis uncom- carex capillariscapillaris L circumboreal uncommon in mon in dry forest openings north of kane lake 2241 moist steeply sloping meadow south of kane lake lower festuca ovina L var brevifoliabrevifolia R br wats F alpine ronezonemone 2332 brachyphylla schultschuit & Schulschultschuittl uncommon in alpine carex elyelynoidesnoides holm uncommon on exposed alpine zone moist to dry meadows and ledges 2273 2407 ledges east of kane lake 2425 oryzopsis exigiaexigua thurbshurb common in dry subalpine carex haydenianahaydeniana olney common in moist subalpine sites north of kane lake 2251 andalpineand alpine meadows 227824312278 2431 phleum alpinum L common in wet to moist subalpine carex incurviformis mack cf var incurviformis and alpine meadows circumboreal 1191 2281 rare seen only in one small steeply sloping moist meadow foaroapoa alpina L common in wet to moist subalpine and at 3353 in east of kane lake 2411 alpine meadows 2296 carex microptera mack uncommon in moist sub- poa cusiccusickiikii vasey var cusiccusickiikii uncommon in moist alpine meadows north of kane lake 2383 to dry subalpine meadows 2296 carex nova bailey common in moist alpine meadows poa cusiccusickiikii vasey var emilisepilis sorisorlsorlimScriscribnlm hitchehitchc 2291 2428 caicco 475 uncommon in moist subalpine meadows 2382 carex phaeocephala piper widely scattered in dry poa gracilgracillimalima vasey uncommon on dry ledges in alpine sites 1190 2430 ertter 2110a forest openings 2238 carex propropositaposita mack common on moist subalpine poa incurveincurva scribn & will uncommon in dry sub- and alpine slopes 2279 alpine meadows 2381 carex rossiirossai boott uncommon in dry aleasareas of foiestforest poa interior redbrydb uncommon on dry alpine slopes understory 2247 and in scree 2426 carex scirposcirpoideaidea michamichx var pseudoscirpoidea poa nervosenervosa hook vasey var wheeleriwheelerewheeleri vasey redbrydb cronacronq common on moist sandy subalpine and hitchc common on dry ledges and in forest understory alpine slopes 1189 2254 2432 caicco 473 north of kane lake 2234 carex scopulorum holm var bracteosebracteosa hermann poa rupicola nash uncommon on dry rockyroeky alpine common in wet meadows along creeks and around kane slopes 2427 lake in subalpine and alpine zones 2282 sitanionSitanion hystrix nutt smith var hystrix elymus carex subnigricans stacey uncommon in moist elymoideselymoides raf Sweswezeyzeyl uncommon in subalpine and alpine and subalpine meadows 2429 alpine zones on dry rocky ledges and slopes 2243 trisetum spicatumspicatum L richter circumboreal juncaceae common in alpine and subalpine zones moist meadows and ledges2280ledges 2280 juncus drummonddrummondiiii E meyer var drummondii common in moist to dry sandy soil of subalpinesubalpineandand alpine slopes 2253225322972297 acknowledgments juncus mertensianusmertensianus bong common in moist alpine meadows 1195 luzula parviparvifloraflora earhehrh desvdeev common in moist this study was supported by the challis subalpine and low alpine meadows 2320 national forest and we greatly appreciate the luzula spicatespicata L DC common on moist unstable especially cindy circumboreal 1197 help of forest personnel slopes of subalpine and alpine zones and steve spencer we 2323 caicco 480 ertter 2103 haggas dave reeder 199211992 KANE LAKE CIRQUE VASCULAR FLORA 343

thank joy mastrogiuseppeMastrogiuseppe washington state GRIGGS BR F 1937 timberlinesTimber lines as an indicator of climate university for rapidly identifying our carex col- trends science 85 251 255 lections steve brunsfeld HARTMANHAIHMAN R LANDLL AND L CONSTANCE 1985 two new spe- university of idaho cies ofcymopterusofcymopterus umbelliferae from western north for identifying our salix arthur cronquist new america bnttoniabrittaniabrittoniaBrittonia 378837 88 95 york botanical garden foridentifyinganwiflfor identifying antenna HENDERSON D 1978 notes on the flora of easieastcast central ria dindimorphadimorpha and barbara ertter university of idaho madronoMadroflo 25 172174172 174 california for identifying potentilla nivea bar- HENDERSON D S BRUNSFFLDBRUNSFELD AND P BRUNSFFLDBRUNSFELD 1981 noteworthy collections from idaho erigeronengeron bara ertter and steven caicco graciously humilis hymenopappus filiafiffilifoliusfihfoltusfolius var idahensisidahoensis opened their collecting books for our examina- carex rupestrismpestnsrupenupestris astragalus admsainnisbinnisamms abissianassiabassiamissi gentiana tion the manuscript benefited greatly from propinquepropinquapropmquapropinqua and papaver kluanensekluanense Madromadronofiobiobho 288828 88 90 reviews by douglass henderson barbara HERMANN F J 1970 manual of caricescabrices of the rocky mountains and colorado basin agricultural hand- ertter and an anonymous reviewer book no 374 USU S department of agriculture forest service washington DCD C 397 appp HITCHCOCK C L 1941 A revision ofthe drabasarabas ofwestern literature CITED north america university of washington publications in botany 111lii11 1 132 HITCHCOCK C L AND CRONOUISTCRONQUISM BAKERBAKEH FFSS 1944 mountain climates ofthe western united LANDAA 1973 flora of the pacific northwest university of press states ecological monographs 1422314 223 254 washington seattle 730 appp BENSON L 1948 A treatise on the north american JENKINS R E 1986 applications and of biological ranunculi american midland naturalist 40140 1 264 use survey data pages 141 152 in K C kim and L BRUNSFFLD S J 1981 alpine flora of east central idaho in knut- son eds foundations for a national biological unpublished thesis university of idaho moscow 205 appp survey association of systematics collections BRUNSFELD S S CAICCO AND D HENDERSON 1983 lawrence kansas noteworthy collections of idaho parnassiaPa kotkotzebuetkotzebuet massia LACK K madrono 306430 64 lackshewitzlackschewitzSHEWITZ KDD HENDERSONHFNDFRSON AND S BRUNSFELDBRUNSt ELD 1983 noteworthy collections of idaho and montana BRUNSFELDBRUNSFFLD S J AND F D JOHNSON 1985 field guide erigeronengeron radiradicatuscatus madronoMadroilo 606460 64 65 to willowsviivillows of east central idaho bulletin no 39 6465 LESICA P AND S SHELLY 1991 forest wildlife and range experiment station uni- PANDJJ sensitive threatened and endangered vascular of montana occa- versity of idaho moscow 95 appp plants sional publication no 1 montana natural CAICCO S L 1983 alpine vegetation of the copper basin heritage program helena 88 area south central idaho unpublished thesis univer- appp MOSELEY R K 1985 synecologicalsynechological sity of idaho moscow 99 appp ecologicalSyn relationships of alpine spike fescue grasslands in east central idaho CAICCO SSANDDAND D M HEHFNDFRSONNDERSON 1981 A of the in unpub- henderson survey lished moscow rare plants of the challis national forest lost river thesis university of idaho 70 appp 1989 the 1989 search district west side with recommendations and man- results of of regional herba- ria for location agement implications university of idaho herbarium information pertaining to idaho s rare flora the fourth search university of idaho moscow 45 appp generation natural heritage section nongame and endangered wildlife program CAICCO S J CIVILLECIVILLF AND D HENDERSONHFNDERSON 1983 note- idaho and game worthy collections of idaho astragalus leptaleusleptaleus and department of fish boise 12 appp MOSELEY R AND C GROVES COMPILERSCOMPILFRS 1990 rare benPenpenstemonpenstenwnpentstemonstemon procerus var formosusformosus Madromadronofioflo 306430 64 threatened and endangered plants and animals daubenmire R 1954 alpine timbertimberlinestimberlmeslinesiines in the americas of idaho natural section nongame and and their butler university botanical heritage interpretation program studies 11119 136 endangered wildlife idaho department of fish and game boise 33 appp DOVER J H 19819811 geology ofthe boulder pioneer wilder- REVEAL J L 1989 new combinations and novelties ness study area blaineblamebiame and custer counties idaho in eriogonum polygonaceae pages 16 75 in mineral resources of the boulder pio enogoniodeaeeriogoniodeae phy tolopiatologia 6625166 251 265 neer wilderness study area blameblainebiame and custer coun- WARDLE 1974 pages ties idaho USU S geological survey bulletin 1497 P alpine timbertimberlineslines 371 402 in J D ives and R G berry eds and EVENSON E B J F P COITFRCOTTER AND M CLINCH 1982 arctic alpine BJ andaandjJ methuen and co glaciation of the pioneer mountains a proposed environments inc london model for idaho pages 653 665 in B bonmchsenbormichsen and R M breckenndgebreckemidgebreckennedgebreekeBreckemidge eds cenozoicaeolozvcenozoic geology of idaho received 3 march 1992 idaho bureau of mines and geology mosowmoscow accepted 29 september 1992 great basin naturalist 524 appp 34434451344351351

LAKEWARD AND downstream MOVEMENTS OF AGEageo 0 ARCTIC GRAYLING HYMALLUSTHYMALLUST ARCTICUS originating BETWEEN A LAKE AND A WATERFALL

I1 mark A deleray and calvin M kaya12

alisABSIrracrvracrracraoKAC 1 arctic grayling in deer lake montana spawn only min the 350 m segment of outlet stream between the lake and a waterfall the purpose of this study was to examine consequences of and possible adaptations by this population to spawning above the falls by determining the extent of loss over the falls of ageageo 0 young the daily and seasonal patterns of such losses and the seasonal pattern of movement upstream into the lake by the remaining young we measured fish niovementsmovements during 1989 and 1990 with traps placed at the outlet and at the falls from fry swimup in july until october or novemberNovem beibel young went over the falls predominantly as newly swimming fry at night in 1989 about 5000 9000 were lost downstream representing adan estimated 4 7 or less of young produced most young thus appear adapted to maintaining thentheir position above the falls A few started entering the lake in august and september but only 95 in 1989 and 23 in 1990 had done so by the time observations were ended by the onset ofwmteryofsinterywintery conditions most movement into the lake appealedappeared to occur sometime during the six to seven months of annual ice cover this extended period of stream residence contrasts with early lakewardd movements reportedrepol ted for other inlet spawning lacustrine grayling populations and may be an adaptation for avoiding piedapledapredationtaontuon by large conspecificsconspecifics in deer lake

key mordiwordsmondi migration fish grayling thymallus arcticusarcticus salmomcksalmosalmonicissalmonsaimongaimo mchmckmohicislols waterfall stream lake

limited information is available on move- based positive rheotaxis by young arctic gray- ments of young fish from populations inhabiting ling thymallus arcticusarcticus in streams kaya 1989 or spawning in small headwater streams above 1991 there have been no previous studies on waterfalls an innate tendency of young fish their possible loss over waterfalls young gray- from such populations to hold position or move ling may be more susceptible to such loss than upstream in water current positive rheotaxis young trout since young grayling are much would be highly advantageous in preventing smaller and appear to be weaker swimmers at their irretrievable loss over the falls such loss swimup initiation of swimming young gray- should be limited to enable the population to ling are about 9 11 mm in length kaya 1991 maintain itself and appropriate behavioral compared to 20 mm or more for rainbow trout adaptation would be promoted through removal northcote 1962 the present observations from the gene pool of young fish that did go were conducted on a population of grayling that downstream evidence for such adaptation is lives in a lake near the head of a mountain valley provided by studies reporting little or no loss and spawns only in a short stream section over waterfalls of young fish from long estabbestab between the lake outlet and a waterfall object- lisheddished native populations of rainbow trout ives of the study were to determine whether oncorhynchus mykiss and cutthroat trout 0 ageageo 0 first year young are lost downstream clarki in north america northcote 1969 over the falls the daily and seasonal patterns of northcote and hartman 1988 and brown trout such losses and the seasonal patterns of their salawsalnwsalnwtruttatnttta in europe jonsson 1982 exper- upstream movement into the lake perpetuation imental studies have provided evidence for a of such a population would depend on limited genetic basis of such rheotactic adaptation in downstream loss of their progeny and residence rainbowtrout and brown trout northcote 19819811 in the lake would require upstream migration by northcote and kelso 1981 jonsson 1982 the young the study was designed to include although there is evidence for genetically movements of the earliest mobile larvae an

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344 199211992 STREAM MOVEMENTS OF AGEAGEOO LAKE GRAYLING 345 aspect that appears lacking from most past stud- ignateeignated wilderness area and is reached via a trail ies involving downstream movements from sal- that extends about 10 km from and climbs about monid populations above waterfalls 1000 m above the nearest motor vehicle access loss of ice cover from the lake and stream and STUDY SITE AND population spawning activities by grayling were monitored through weekly hikes to the site starting in late observations were conducted in 1989 and may observations ofoffishfishhishbish behavior started as the 1990 on the 350 m long section of deer creek ice thawed and adults began entering the that flows from deer lake to a 3 m vertical stream mid june in 1989 and late june in 1990 waterfall the lake is located at 2780 m altitude and ended as ice started forming on the lake and near the head of a mountain valley in the mad- stream margins 10 november 1989 or as snow ison range of southwest montana dimensions accumulations on the trail made access difficult of the stream on 19 august 1990 measured 11 october 1990 stream temperature was bank to bank at five locations along each of 34 continuously recorded throughout both obser- transects between the lake outlet and the water- vation seasons with a peabody ryan model D fall deleray 1991 were mean width of 5885.88 m thermograph placed about 30 m downstream range ios1081.08 21021.0910gio mean depth of 0100.10olo m range from the lake daily mean temperature was 000.0oo 0410.41 and mean water velocity measured calculated as the average ofdaily maximum and at 060.6og X depth at each location of 0050.05 msecmseeasee minimum range 000.0oo 0480.48 estimated discharge volume trapsarithTraptraps withsArith 1 mm mesh netting were placed to ranged from about 0020.02 to 0050.05oos isecmsecc between determine the dates fry became free swimming 2 july and 9 september 1990 and to monitor their movement downstream previous observations had indicated that and upstream out of the outlet stream in 1989 arctic grayling the only fish in the lake spawn three fry emergence traps 45 X 45 cm of the only in the outlet stream kaya 1989 the outlet type described by fraley et al 1986 were stream is inhabited by grayling fry ageoage 0 fish placed over the substrate after most spawning smaller than about 252.5 cm in length piper et al had ceased over areas where fish had been seen 1982 and other young up to about 14 cm in spawning and where concentrations of eggs length larger fish are rare except when spawn- were visible because arctic grayling spawn ing adults are present during early summer over the substrate without excavating reddsnedds numbers of adults spawning in the stream were eggs were readily visible among the substrate estimated by electrofishingelectro fishing mark and recap particles one emergence trap was placed in a ture methods at 803 95 CL 104 in 1989 and spawning area about 30 m below the outlet and 1109 95 CLcl124124 in 1990 with similar num- the other two were placed in the principal bers of males and females deleray 1991 the spawning area about midway through the 350 m segment between the lake and the water- stream length swimup fry in the traps were fall is the only part of deer creek inhabited by removed and tabulated daily or on alternate grayling near the base of the waterfall the days until emergence ceased stream disappears beneath the surface ofa steep one way traps were placed across the lake talus slope before reemerging about 200 m outlet and at the top of the waterfall to monitor downslope grayling are not present in the 10 movement of young out of the stream the km of stream between the lake and the gallatin upstream trap had V shaped screened barriers river perhaps because of the stream s steep extending completely across the outlet and lead- gradient about 1000 miomlom10 km and numerous ing upstream into a holding box this trap cascades fish habitat is absent upstream from retained fish as they entered the lake the trap the lake and the population is thus physically was installed after most adult spannersspawnersspawners had left isolated within the lake and the short section of the stream but before the young became free stream above the waterfall swimming after installation the trap was in continuous operation through both observation METHODS seasons it was inspected at intervals varying from several days to about one week young methods and observation schedules were were removed measured and released influenced by the relatively remote location of upstream into the lake the study site the lake is located within a des the downstream trap was a drift net with its 346 GREAT BASIN naturalist volume 52

260 about 11 july peaked in midmonthmid month and con- ergeemerge tinued until about 25 july fig 1 spawning in 220 n 0 falls 1990 occurred during the first week in july and 180 swimup of fry began in midmonthmid month and contin- ctcc ued to the end of the month UJ 14- n 92 in 1989 fry started appearing in the falls trap 03 z 100 f as they became free swimming fig 1 highest daily totals of fry in the falls generally over n trap 600 i 200 per day occurred 15 22 july as numbers of fry becoming free swimming in the emergence 20 L n h in traps peaked and then declined the swimup 6 8 10 12 14 16 18 20 22 24 26 28 30 1 3 period ended about 25 july thereafter within a JULY AUGUST week numbers of young in the falls trap declined to 0 6 per day no young entered the fig 1 total number ofyoungofowyoungyoung arctic grayling thymallus falls trap after 20 september arcarcharchcusarcheusorcharcticusticusficuscus in three emergence traps placed over the sub- movement of fry over the falls was concen- strate and in the waterfall trap deer creek montana 1989 trated within a 19igdayday period 13 31 july the falls trap was operated for 13 of these days and opening positioned at the lip of the waterfall it the mean number of fry per 24 h sample was collected young that were going over the falls 1273127.3 extrapolation from the estimated 30 in 1989 this trap sampled about 030.3 050.5 of the 50 of total stream volume that passed through stream volume as estimated by comparing flow the net and application of the 13 day mean to rate into a plastic sack attached to the trap versus 19 days yielded a crude estimate of 4837 8062 estimated stream discharge volume in 1990 young grayling lost over the falls 13 31 july V shaped barriers were added to direct all flow numbers in the falls trap averaged only 272.7 per through the net in 1989 the trap was installed day during the 11 days sampled from I1 august on sampling days and left in place for about 24 h to 20 september the last day young entered the before the young within were tabulated and trap similar extrapolation to this 51sidayday period measured the trap was deployed on 6 july yielded a crude estimate of an additional 275 before fry became free swimming and oper- 459 young lost thus the number of young lost ated at intervals of one to two days until num- downstream over the falls in 1989 during the bers in the trap declined sharply thereafter the period from swimup of fry to onset of ice cover trap was operated at intervals of several days to over the stream was roughly estimated at 5000 two weeks until 19 october in 1990 this trap 9000 was operated less at intervals frequently rang- fry were already becoming free swimming from five ing days during the swimup period to when the falls trap was installed on 23 july 1990 about four weeks in and to september october numbers of young per day in the falls trap determine diel of patterns movement over the peaked at 561 on 28 july diminished to 49 ten falls of voung at different s i young ages postpostsswimupwmupwhup days later on 6 august and to 5 by 8 september sampling began on 23 july as fry started to no young entered the trap on 12 october the swim on sampling dates the trap was deployed last day sampled in 1990 during the swimupsa4mup at 1000 or 1100 h mountain standard time period fry went over the falls predominantly at the trap was emptied of young at 1400 h and night 23 and 28 july fig 2 however there thereafter every 4 h until 1000 h the next day was no consistent pattern of diurnal vs noctur- nal movement among the fewer young fish that RESULTS went over the falls on later dates 6 and 17 august fig 2 too few days were sampled at spawning occurred through much of the the falls in 1990 to estimate total numbers lost 350 m length of the stream from about 10 in in contrast to early losses over the falls below the lake outlet to within 15 20 in of the upstream movement of young grayling into the falls the most heavily used area was a 10lom m lake did not begin until late summer when the reach about 130 140 in above the falls in 1989 fish were larger and water temperatures were spawning occurred during the last week ofjune cooling fig 3 small numbers of young were and swimup offry in the emergence traps began trapped at the lake outlet starting in mid august 199219921 STREAM MOVEMENTS OF AGEAGEOO LAKE GRAYLING 347

100lov 16 90- july 23 1989 14 1990 80 70 12 i LU T 60 to UJ co S 505 8 ZS 40 Q Wx 30 4 20 2 to10

A lake 89 july 28 70 alake 260 lake 90 AA a 60 0afallshfallsfallsfalis 90 220- 50 0 180 UJLU 40 03 t 140 luZ Zi j 30 100 20 00 60 10

20 aB 1989 26 1990 40- august 6 0cr 22 aujn 303- i cca 18 1 20 UJ 10 14 Z 10 40 august 17 cr 6 lu 30- xM 89 90 D 20 t 2 t Z1 f

10 1 10 20 30 10 20 30 10 20 30 10 20 0n- july august september october 12 16 20 0

HOUR OF DAYOAY fig 2 diel pattern of young arctic grayling thymallusThy maliusmaltus fig 3 mean daily temperature OCC of deer creek ardicus accumulated in waterfall trap during 3 or 4 h montana mean total lengths of ageageo 0 grayling thymallus sampling periods 1000 or 1100 h to 1400 h then at 4 h ararcticusarcticusdicus inm waterfall and lake traps and numbers of ageageo 0 intervals thereafter deer creek montana 1990 note grayling in the lake trap 1989 and 1990 arrows indicate change of yaksyansy adsaksans on july 28 mean sizes of young on these datesates when numbers of young becoming free swimming sampling dates are in figure 3 arrows indicate sunset and collected in emergence traps peaked during 1989 and sunrise 1990

1989 and early september 1990 total numbers although numbers of resident young in the ofyoung trapped per 3 day to I1 week periods in stream were not estimated visual observations september and october were 0 26 in 1989 and indicated that ageageo 0 fish were abundant in 0 14 in 1990 fig 3 only 95 ageageo 0 young had november 1989 as ice was starting to form on moved up into the lake in 1989 and 23 in 1990 the stream but were present in much fewer before observations were terminated by the numbers as agelage I1 fish when ice cover melted onset of winterlike conditions in november the following june the agelage I1 fish still in the 1989 and october 1990 while ageageo 0 grayling in stream in june 1989 and 1990 had upstream the falls trap were mostly newly swimming fry patterns of movement similar to those of the that averaged 12 14 mm in length the smallest affeageageoaggeaggeo 0 fish very few entered the lake during the moving upstream into the lake trap averaged june november study period and these lim- 52 54 mm in length fig 3 ited upstream movements occurred mostly 348 GREAT BASIN naturalist volume 52 between early september and the end ofobser- stream movement of the young was similar to vations in october or november in 1989 only those of young from inlet spawning populations seven agelage I1 fish were in the lake trap from june ofgrayling kruse 1959 lund 1974 wells 1976 to the end ofaugust and 38 more from septem- and other salmonsalmonidsids mccart 1967 northcote ber to the end of observations in november in 1969 brannon 1972 these observations were 1990 only two agelage I1 fish were trapped both in also consistent with results of experiments in an september age I1 fish were nearly absent from artificial stream kaya 1989 which indicated the falls trap three were trapped in 1989 and that although young deer lake grayling had an two in 1990 fish older than agelage I1 were rare in innately greater tendency to swim upstream the stream when ice cover thawed in june of than those of an inlet spawning population many both years moved downstream especially in darkness during the summer of 1990 six adults if loss over the falls results from deliberate remained in the outlet stream these fish were downstream migration by the young then this seen in shallow water 5 10 cm deep chasing may indicate that the deer lake population has groups of young in late july one was captured not yet completely adapted to outlet spawning with a dip net and had 12 ageageo 0 grayling in its if so then the waterfall is continuing to act as a stomach selective factor removing those young with inappropriate responses incomplete adapta- discussion tion has also been suggested as an explanation for downstream movement by many swimupswim up fry ofrainbow cutthroat hybrid trout that spawn in we did not estimate the number of since the outlet of a colorado lake lentsch 1985 produced in the stream we do not know young lake had first been planted with trout about the percentage of total young lost over the fallsbailsilslis the 100 years earlier little or no downstream loss between swimupswimup and the end ofobservations in has from populations of brown october and november two considerations been reported and rainbow trout native to waters above falls suggest that the losses represented a relatively northcote 1969 1981 jonsson 1982 small percentage of young produced first it northcote and hartman 1988 in contrast to visually apparent that ageageo 0 young was cascades ofan esti- remained abundant and widely distributed downstream movement over marked rainbow trout in a stream throughout the stream until the end of each mated 22 of stocked repeatedly in preceding observation season second we estimated that that had been rainbow trout chapman the number of eggs that could have been years with nonnativenormative may population spawned by this population during 1989 was and 1986 the deer lake almost certainly originated through a transplant about 131.3 million this was based on the esti- of 2988 eggs in each of seven of young from an inlet spawning population mated average mon- females sampled range 2459 3674 and the sometime during the present century in estimated number of 426 adult females in 1989 tana grayling were not present above natural and only deleray 1991 if we assume as an example barriers to upstream movement the natu- that sa4mupswim up fry resulted from 10 ofthis poten- lakes within the original range that were have tial egg deposition then the estimated loss of rally accessible to fish and known to con- and young over the falls 5000 9000 would be tained native grayling were upper lower about 4 7 of fry produced in 1989 we do not red rock lakes and perhaps elk lake of the know of any estimates of the relationship red rock river drainage nelson 1954 vincent between potential egg deposition and actual fry 1962 another lacustrine population originated production by grayling however a figure of with the creation of ennis reservoir on the 10 seems conservative compared withwi&recentrecent madison river which contained native grayling estimates of lis11511.5 22222.2 for chum salmon the red rock elk and ennis populations are oncorhynchus keta and 16416.4 29129.1991 for coho inlet spawning populations in other lakes orig- salmon 60 kisutch in a canadian stream with inated through stockings that began after artifi- the lower percentages associated with poor sub- cial culture of the species was initiated in 1898 strate quality scrivener and brownlee 1989 henshall 1906 unpublished records of the grayling lost downstream were predom- regional state and federal hatcheries involved inantly small newly swimming fry that went in these stocking programs indicate that fertil- over the falls at night the nocturnal down ized eggs were obtained from upper red rock 199219921 STREAM MOVEMENTS OF AGEAGEOO LAKE GRAYLING 349 lake or ennis reservoir or other inlet spawning from the immediate spawning areas some of populations established through transplants them apparently downstream for those from these two sources kaya 1989 1990 becoming free swimming near the falls even outlet spawning populations are known to have localized downstream dispersal could result in evolved elsewhere from transplants of inlet some being carried over especially under con- spawning grayling kruse 1959 and rainbow ditions of poor visibility at night trout northcote 1969 the results indicate that deer lake grayling it is possible that downstream loss of many spend at least the first and possibly also their young fish occurs even from populations well second summer and early to midautumnmid autumn in the adapted to spawning above a waterfall with outlet stream however the results did not native above falls populations that have been permit us to determine the exact timing of most studied the young sampled were brown trout movement by young into the lake or whether from about 10 cm to over 20 cm in length they move upstream predominantly as ageageo 0 or jonsson 1982 or rainbow and cutthroat trout as agelage I1 fish the very few young that moved whose sizes were not stated northcote 1969 into the lake during both observation seasons 1981 northcote and hartman 1988 given the could not account for the numbers of spawning rapid post swimup decline of downstream adults produced in the population since there movement observed in the present study con- is no other source of young and since the 1989 clusionsclusions on magnitude ofsuch losses would have observation season extended over the entire ice been very different if the sampling had begun free period on the stream maintenance of the one or two weeks after the end of the swimup deer lake population must depend on period or if the only fish sampled were larger upstream movement ofyoung sometime during than 151.5 202.0 cm the six to seven months of annual ice cover factors other than deliberate downstream although ageageo 0 young greatly diminished in movement could have produced losses over the numbers and agelage I1 fish virtually disappeared falls including passive drift or local dispersal from the stream between the onset of ice cover those young that were lost could have origi- in november 1989 and its thawing in june 1990 nated from eggs either spawned within or we do not know the proportions of these reduc- drifted to locations close to the falls adults tions in numbers attributable to movement into spawned within 15 20 m above the falls and we the lake death or loss over the falls the greatly confirmed visually that many eggs drift down- diminished numbers of young in the falls trap stream from spawning areas after being broad- during late summer and their absence in the cast over the substrate fry originating from trap by october ofboth years suggest that down- eggs near the falls could be lost through passive stream losses duringvinterdurinduringdudinggvinterwinter may be small the drift if they became free swimming at night and chronology of major movement by young gray- were consequently displaced downstream in the ling into the lake and the numbers and ages of darkness as has been described of european fish involved would need to be resolved by grayling T thymallus bardonnetbardonnerBardonnet and gaudin observations during winter 1990 downstream losses could also represent little is known about duration of stream passive drift of dead or unhealthy fish as sug- residence for outlet spawning populations of gested by a report that 81 of young brown arctic grayling young from inlet spawning pop- trout produced in a section of stream did not ulatulationsions of the species typically have an early survive and drifted downstream mostly at night descent to the lake ranging from immediately elliott 1986 we did not attempt to determine after swimup kruse 1959 lund 1974 wells the health of young grayling in the falls trap 1976 to within several weeks nelson 1954 loss over the falls could be an indirect con- we are not aware of other studies on stream sequence of local dispersal of young within the residence times of young grayling from outlet stream as they became free swimming young spawning populations and so do not know sockeye salmon oncorhynchus nerka of whether extended period of stream residence is outlet spawning populations have been typical for such populations young rainbow reported to temporarily disperse downstream trout of outlet spawning populations tend to before holding position or swimming upstream remain for extended periods of at least a month into lakes mccart 1967 brannon 1972 young to a year or more before migrating upstream to grayling in deer dreek also disperse locally lakes while those ofinlet spawning populations 350 GREAT BASIN naturalist volume 52

migrate when newly swimming in some popula- residual adult and agelage 1 grayling remaining tions and after extended periods of stream resi- through the summer and an occasional belted dence in others northcote 1969 the kingfisher aves ceryle alcyon thus the extended stream residence ofyoung deer lake movements of ageageo 0 deer lake grayling that grayling is also consistent with their lesser ten- remain in the outlet stream appear adapted both dency to swim upstream in an artificial stream to beginning their existence a short distance as early fry from swimup to three weeks com- above a waterfall and to avoidance of predation pared with their responses when older within a by larger conconspecificsspecifics in the lake study period of up to 10 weeks post swimup kaya 198919911989 199 1 acknowledgments it may be that young of an outlet spawning population need to attain larger sizes and these observations were part of a study sup- thereby become stronger swimmers before they ported by a grant from the montana depart- can swim upstream into the lake however this ment of fish wildlife and parks possibility appears contradicted by our casual ageo 0 of all sizes in observations that age grayling CITED deer creek starting from those newly swim- literature of ming were capable swimming upstream BARDONNFT aandpgaudin 1990 dieldielpatternofpattern of first when they were disturbed by our presence downstream post emergenceernergencegenee displacement in grayling those young originating from spawning areas thymallus thyrthymallusmiliusmillus L 7581758 journal of fish biology within a few meters ofthe lake outlet could have 3762337 623627623 627 BRANBRANNONNON E L 1972 mechanisms controlling migration of entered the lake by moving only a short distance sockeye salmon fry international pacific salmon fish- upstream eries commission bulletin 21 86 appp another possible factor quality of rearing CHAPMAN D WWANDAND B MAY 1986 downstream move- habitat also does not to favor extended ment of rainbow trout past kootenai falls montana appear north american journal of fisheries management 6 residence in deer creek deer lake grayling 475147 51 grow slower during their first two years than DELEDFLFRAYRAY M A 1991 movements and utilization of fluvial those of other lacustrine populations studied habitat by ageageo 0 arctic grayling and characteristics of gallatin thus far they grow at spawning adults in the outlet of deer lake in montana but thereafter county montana unpublished mastermasterss thesis mon- similar or faster rates deleray 1991 unlike tana state university bozeman 75 appp young deer lake grayling those from inlet elliotELLIOIELLIOTT 1 J M 1986 spatial distribution and behavioural spawning populations in montana spend their movements of migratory trout salmo trutta in a likelakehikehuke ofanimal 55 907 922 season in district stream journal ofanimal ecology first summer and autumn growing in ERIKSEN C H 1975 physiological ecology and manage- lakes the slower early growth of deer lake ment of the rare southern grayling ThymathymallushusUus aratiarcti grayling thus appears related to their spending cus tricolor cope Internationinternationaleinternationalsale vereinigungvereimgung fur their first growing seasons in the stream rather theoretischeTheore tische und angewandteangewandto limnologielimnologicLimnologie 19 2448 lake 2455 than in the fraleyFHALPYFRALFY J J M A GAUB AND J R CAVIGLICAVIGLL 1986 emer- we speculate that young deer lake grayling gence trap and holding bottle for the capture ofsalmoof salmo may remain in the outlet stream to avoid intra- nid fry in streams north american journal of fisheriesoffisheries specific predation in the lake eriksen 1975 management 6 119 121 HENSHALLHFNSIIALL J A 1906 culture of the montana grayling observed that ageageo 0 grayling in several montana report of the commissioner offisheries for the fiscal lakes occupied shallow near shore areas among year 1906 and special papers USU S bureau of fisher- rooted aquatic vegetation and suggested that ies document 628 washington DCD C 7 appp salmosainto their distribution provided protection against JONSSON B 1982 diadromous and resident trout trutta is their difference due to genetics oikosbikos 38 predation by the adults behavior of the few 297 300 post spawning adults that remained in deer KAYA C M 1989 rheotaxis of young arctic grayling from creek during the summer of 1990 confirmed populations that spawn min inlet or outlet stistreamsearnsearms of a that adults will prey on the young young gray- lake transactions of the american fisheries society ils118474118 474 481 ling would likely be susceptible to predation by 1990 status report on fluvial arctic grayling thy- larger conconspecificsspecifics in deer lake because of its mallus arcticusarcticus in montana prepared for montana high water clarity throughout the summer and department of fish wildlife and parks helena 9797ppappp outlet 1991 rheotactic differentiation between fluvial the lack of rooted macrophytes in the and lacustrine populations of arctic grayling thy- stream the only potential predators of young mallus arcticusarcticus and implications for the only remain- grayling that weesawwsawsaw were the relatively few ing indigenous population of fluvial montana 199219921 STREAM MOVEMENTS OF AGEAGEOO LAKE GRAYLING 351

grayling canadian journal of fisheries and aquatic NORTHCOTEORTHCCHE T G AND G F HARTMAN 1988 the biology sciences 48 53 59 and significance of stream trout populations salmo KRUSE T E 1959 grayling of grebe lake yellowstone sppapp living above and below waterfalls polslaepolskie national park wyoming USU S fish and wildlife ser- archiwumArchiwum hydrobiolognhydrobiologyhydrobiologii 35 409142409 442 vleevice fishery bulletin 149 307 351 northcote T G AND B W KELSOKFLSO 1981 differential LENTSCH L D 1985 evaluation of young of the year response to water current by two homozygous LDH production in a unique colorado wild trout population phenotypes of young rainbow trout canadianCanaalandlandian journal unpublished mastermasterss thesis colorado state univer- of fisheries and aquatic sciences 38 348 352 sity fort collins PIPERiperIPFR R G I1 B MCELWAIN L E ORMEORMF J P MCCRARFNme CRARFN LUND J A 1974 reproduction of salmonsalmonidsids in the inlets L G FOWLERfowlfrandjAND J R LEONARD 1982 fish hatchery of elk lake montana unpublished master s thesis management USU S department of the interior fish montana state university bozeman and wildlife service washington DCD 517 appp mccart P 1967 behaviour and ecology of sockeye salmon SCRIVENER3rivfner J C AND M J BROWNLEE 1989 effects of fry in the babine river journal of the fisheries forest harvesting on spawning gravel and incubation researchReseaichsealch board of canada 24 375428375 428 survival ofchum oncorhynchusoncorhynchusketaketa and coho salmon NELSON P H 1954 life history and management of the 0 kisutch in carnation creek british columbia american grayling thymallus signifer tricolor in canadian journal of fisheries and aquatic sciences 46 montana journal of wildlife management 18 325 681 696 342 VINCENTINCFNT R E 1962 biogeographical and ecological factors NORTHCOTE T G 1962 migratory behaviour of juvenile contributing to the decline of arctic grayling thy- rainbow trout salmo gairdnergairdnengairdgairdnerinerinefinen in outlet and inlet mallus arcticusarcticus pallas in michigan and montana streams of loon lake british columbia journal of the unpublished doctoral dissertation university of mich-Mich fisheries research board of canada 19 201 270 igan ann arbor 1969 patterns and mechanisms in the lakeward WELLSELLS J D 1976 the fishery of hyalite reservoir during migratory behaviour of juvenile trout pages 183 203 1974 and 1975 unpublished mastermasterss thesis montana in T G northcote ed symposium on salmon and state university bozeman 47 appp trout in streams university of british columbia van- couver BC canada received 7 may 1991 1981 juvenile current response growth and matu- accepted 15 august 1992 rity of above and below stocks of rainbow trout salmo gairdnergairdnengairdgairdnerinerinerlnefinen journal of fish biology 18 741 751 gigreateat basin naturalist 524 appp 352356352 356556

EFFECTS OF BROWSING BY MULE DEER ON TREE GROWTH AND FRUIT production IN JUVENILE ORCHARDS

I1 dennis D austinlandand philip J urness

aiisrracABSIRACIrthearthethe effects of big game depredation on juvenile fruit trees were studied in northern utah utilization of trees was determined by counts of nipped and intact buds in spring height width basal diameter number of buds and initial flintfruit production of peach and apple trees were determined from trees protected from or browsed by mule deer in wmteiwinter results from the 10 orchards studied indicated that removal of buds at the observed browsing levels had no effect on tree growthglowth 01or initial fruit production

ketikeifkeykeg worchwordsmorchmonchmonds depredation mule deer orchards truitfruittrutt trees deer dandanagedanvigedanwgevige evaluation apple trees peach trees winter browsing

whenever depredation occurs in commer- than protected limbs suggesting possible cial orchards potential crop losses due to big growth stimulation as a result of deer browsing game browsing become a major concern to in our project only bud removal browsing growers browsing of juvenile fruit trees has was studied and since browsing during summer important economic consequences because the was negligible we considered only overwinter effects may limit future crop production and depredation the purpose of this study con- increase tree mortality research has clearly ducted in northern utah was to measure the shown that browsing by big game on mature degree of browsing in young fruit trees and to apple trees causes significant crop loss within assess the browsing effects on tree growth and initial the browsing zone katsma and rusch 1979 crop production 1980 austin and urness 1989 however lim- ited information on the effects of browsing on METHODS juvenile fruit trees is extant westwood 1978 suggested deer browsing the percentage of buds browsed by mule may be especially damaging to young trees but deer was determined in march during late dor- rarely would browsing be expected to cause mancy after deer switched diets from winter mortality harder 1970 reported no differ- browse to herbaceous spring growth kufeld et al 1973 umessurness percent ences in trunk diameter growth between pro- austin and 1983 bud removal tected and unprotected apple trees with one was determined by counting all intact and nipped buds and then dividing nipped buds winter of bud removal browsing by mule deerr dee by the total nipped plus intact buds nipped this colorado study of 160 trees no mortality in buds are easily identified by the exposed and was attributed to bud removal browsing broken woody twigs katsma and rusch 1979 although 8 trees died as a result ofbark damage counted intact buds were restricted to terminal caused by antler rubbing similarly mcaninch buds of the previous summersummerss annual growth et al 1985 in a new york study reported 9 of and all protruded buds along second year and 10 growth parameters measured between pro- older stems 1 cmem in length austin and umessurness tected and browsed trees showed no significant 1987 protruded was defined by visualizing a differences one parameter basal diameter was perpendicular line from the twig to the tip ofthe smaller on browsed trees however this study bud and an observable space was required with white tailed deer also showed that average between the line and the bud twig intersection diameters of browsed limbs appeared greater tree growth measurements were taken after

dantdaut lurturlux litmt ofrangeorrangeof ringehinge science utah stitestate university logan utah 84322523084329323084329393084322 5230

352 199211992 DEER BROWSING IN JUVENILE ORCHARDS 353 the end ofthe growing season but before winter deleted from the sample trees were placed into browsing occurred tree hei4htheight was measured three equal groups of 67 by the percentage of to the nearest 101.0iolo cm from ground level tree bud removal browsing damage heavy 61 width to the nearest 101.0loio cm at the height where 100 moderate 34 60 and light 0 33 maximum width occurred width was measured tree measurements were made following the in north south and east west directions and the 1987 summer growing period no differences in mean recorded basal trunk diameter was mea- tree measurements were found among the three sured to the nearest 010.1oi cm using dial calipers at intensities of browsing by mule deer table 1 10 cm above the graft scion diameter was sim- 3 ilarly measured on north south and east west orchard directions and the mean recorded the number twelve pairs of equal age and size yellow of intact buds using the same definition as that delicious apple trees were carefully selected by for bud removal determinations was counted ocular observation within a commercial orchard using hand tally registers where harvestableharvestable planted during spring 1984 one tree of each crops were produced all fruits were hand- pair determined by coin toss was protected picked and counted specific methods are from browsing by fencing during five winters reported in the results for each orchard 1984 89 during the sixth winter 1989 90 for data were analyzed between protected and the same reason as described for orchard 1 all browsed trees and between trees with various trees were fenced intensities of browsing using the standard t test percent bud removal from browsing was ofthe means confidence level was set at P 05os05.05 76460541723676460541723.676476.4 60560.5 41741.7 23623.6 and 63263.2 for years 1985 89 respectively no differences between pro- RESULTS tected and browsed trees were found for any tree measurements or fruit production table 1 orchard I1 orchard 4 A 4 x 6 block of24 equal age aiandauldid size elberta peach trees planted in spring 1986 was twelve pairs of equal age and size red deli- selected for study alternating trees deter- cious apple trees were carefully selected by mined by coin toss were fenced during three ocular observation within a commercial orchard winters 1986 89 during the fourth winter planted in spring 1983 one tree of each pair 1989 90 all trees were fenced becausebecausea4thinwithin determined by coin toss was protected from year browsing effects decrease fruit production browsing by fencing during three winters katsmakatsina and rusch 1980 austin and umessurness 1984 87 during winter 1986 87 a deer proof 1989 trees were protected from browsing to fence was constructed around the orchard and compare production between previously consequently deer use was close to zero 04040.4 browsed and protected trees tree measure- during the two previous winters 1984 86 per- ments were taken and peaches were hand- cent bud removal was 71071.0 and 170ito17.0 respec- picked and counted in late summer 1990 the tively no differences between protected and first year of commercial harvest browsed trees were found for either tree mea- percent bud removal as measured in spring suresurementsments or number of fruits table 1 also which collected 198719881987.19881987 1988 and 1989 was 35676635676.635635.6 76676.6 and 73573.5 flower cluster counts were in respectively even with this high degree of spring 1987 as part of an ongoing parallel study by trees fully recovered during austin and urness 1987 showed no difference browsing deer x the summer growing seasons no differences between protected x 166 and browsed x between protected and browsed trees were 169 trees fruit found for any tree measurements or pro- orchard 5 duction table 1 twelve pairs ofequal age and size red deli- orchard 2 cious apple trees were selected within a com- A small commercial orchard comprising 210 mercial orchard planted in spring 1985 one elberta peach trees was planted in spring 1986 tree of each pair determined by coin toss was percent overwinter bud removal was deter- protected from browsing during four winters mined in early spring 1987 since 9 trees 1985 1989 during the fifth winter 1989 90 all showed bark scraping damage they were trees were fenced 354 GREAT BASIN naturalist volume 52

from TABLETABLT 1 mean growth measurements and initial fruit production from juvenile peach and apple trees protected or browsed by mule deer in winter

meanmf an tree irmeasurementsleasuremenleasuremen ts basal no of no of orchaorchardrd buds height width diameter no fruit tree treatment N years removed cm cm mm buds fruits 104 I1 elberta peach browsed 12 1986 90 62 225 257 56 protected 12 230 247 57 103

2 elberta peach heavily browsed 67 198619868787 61 100 120 88 26 61 moderately browsed 67 34 60 124 92 27 67 lightly browsed 67 003333 122 91 27 65 3 yellow delicious browsed 12 1984 90 53 192 136 51 250 72 70 apple protected 12 193 149 52 238 75 4 red delicious browsed 12 1984 87 44 569 248 44 349 apple protected 12 588 262 44 375 59 577 3 5 red delicious browsed 12 1985 90 24 259 163 54 apple protected 12 250 158 54 570 3

6 golden Dedehcousdelicousdeliciouslicous heavily apple browsed 20 1987 65 92 198a198 093 35 96 moderately a browsed 20 28 64 1921921 88 35 93 lightly browsed 20 0 27 17511 8pap80 35 92

7 red delicious heavily apple browsed 8 1985 86 49 88 22 17 11 moderately browsed 8 21 98 30 18 10 protected 8 92 21 16 7

8 McIntomcintoshapplemcintoshmclntoshmeintoshshappleshappieapple heavily browsed 8 1985 86 50 132 62 24 31 moderately browsed 8 35 126 47 21 22 protected 8 129 44 26 17

9 jonathanappleJonathajonathan appleappienapplenappie heavily browsed 8 1985 86 28 147 69 24 26 moderately browsed 8 22 123 48 20 22 protected 8 131 69 20 45

10 red delicious browsed 12 1985 87 394 167 67 51 90 apple protected 12 159 63 50 107

figures witlivvithavith differentditdifdlf fuent superseriptedsuperscriptedsiupfcrunpted numbers within columns were significantly different P 50505

percent bud removal from browsing was orchard 6 16716.7 000.0oo 16716.7 and 61061.0gio for years 1985 89 two old golden respectively no differences between protected A 2 X 30 block of 60 year and browsed trees were found for any tree mea- delicious apple trees was measured for over- suresurementsments or fruit production which was winter bud removal browsing use in spring greatly reduced in 1990 due to cold tempera- 1987 utilization during the previous winter was tures in spring table 1 unknown but was probably similar to the use 1992 DEER BROWSING IN JUVENILE ORCHARDS 355 measured in 1987 percent bud removal ranged balance with the root system this was the from 0 to 92 with a mean of 46746.7 table 1 observed case trees were placed into three groups of 20 by in this study trees were not browsed bud removal classes 0 27 28 64 and 64 92 severely As a suggested definition severely surprisingly heavily and moderately browsed browsed trees would include browsing of90of 90 trees had significantly greater height at the end of the available protruded buds removal of of the growing season than lightly browsed 70 of the current annual growth scraped trees and heavily browsed trees also had greater bark on the central leader andor scraped bark width than lightly browsed trees table 1 on two or more primary branches or limb although other factors such as pruning could breakage certainly as the level of browsing have accounted for these increases height and increases toward severe levels the potential for width may have been increased by browsing no permanent damage and reduced growth also differences were found in basal diameters or increases the level of browsing intensity number of buds needed to damage juvenile fruit trees is unknown but it is apparently higher than that orchards 7 8 9 which occurs in most depredation situations in twenty four equal age and size trees of red northern utah and elsewhere harder 1970 delicious meintoshmclntoshmcintosh and jonathan apples were mcaninch et al 1985 planted in spring 1985 for this study in winter the intensity of browsing needed to cause 1985 86 one third 8 of each species of the measurable damage would also be expected to trees randomly selected were protected one vary with the quality of the horticultural prac- third received moderate browsing by tame mule tices involved in managing the orchard in this deer as modified by temporary fencing and study all orchards received high intensity care one third received heavy browsing mean bud including adequate irrigation periodic spray- removal varied from 21 to 35 under moderate ing weed control etc orchard trees receiving browsing and 28 to 50 under heavy browsing lower intensities of care and increased environ- table 1 following the summer growing mental stress from pests or competition from season in 1986 no significant growth differ- weeds may respond negatively to similar levels ences in tree measurements were found of deer browsing between protected moderately browsed or in conclusion the results from this study of heavily browsed trees table 1 juvenile apple and peach fruit trees were con- sistent with previous research 1970 orchard 10 harder mcaninch et al 1985 browsing by mule deer twelve pairs of equal age and size red deli- at the intensities observed had no negative cious apple trees were selected within a com- effects on tree height width basal diameter mercial orchard planted in spring 1983 one number of buds or initial fruit production tree of each pair determined by coin toss was protected from browsing during winters 1985 87 percent bud removal from browsing was acknowledgments 76676.6 37437.4 and 414.1 respectively no differ- ences between protected and browsed trees this report is a contribution of the utah were found table 1 state division of wildlife resources federal aid project W losios105 11 discussionDiscussiondiscussion CITED percentages of bud removal measured from literature these 10 orchards10orchards were mostly less than 65 browsing by mule deer duringa4nterduringduning winter dormancy austinauslinAUSIIN D DANDPJD AND P J URNFSS 1983 Ovenoverwintervinterminter forage selection by mule deer on seeded big sagebrush grass at this level of use was not sufficient to cause a range journal ofwildlife management 47120347 1203 1207 decrease in tree growth parameters measured 1987 guidelines for evaluating crop losses due to from the view of carbohydrate reserves depredating big game utah division of wildlife decreased pproductivity would not be expected resources publication 87 5 42 appp the total buds 1989 evaluating production losses from mule deer if number of intact available for depredation in apple orchards wildlife society bulle- spring growth were sufficient to maintain tin 1716117 161 165 356 GREAT BASIN naturalist volume 52 lianderHARDER J D 1970 evaluatingwinterevaluating winter deerusedekrusedeer use oforchards department of agriculture forest service research in westelwesternn colorado transactions of the north amer- paper RM 111 31 appp ican wildlife conference 35 351735 47 mcaninch J B M R ellingwoodELHNGWOOD M J FARGIONEFARGIONF karsKAISMAMA D E AND D H ruscilruscruse II11 1979 evaluation of deer anupplcqnfAND P PICONE 1985 assessing deer damageinyoungdamage in young damage in mature apple orchards pages 123 142 in fruit orchards proceedings of the wildlife damage J R beck ed vertebrate pest control and manage- control conference 2 215 223 ment materials ASTM STP 680 american society for WFSIWOODWE STWOO D M N 1978 temperate zone pomology W H testing material freeman and company san franscisco californiaCahforma 1980 effects of simulated deer browsing on 428 appp branches of apple trees journal of wildlife manage- ment 44 603 612 KUFELDkunkum LD R caC 0 C WALLMO AND C FEDDEMAFFDDFMA 1973 received 20 april 1992 foods ofc0the rocky mountain mule deer united states accepted 18 september 1992 great basin naturalist 524 appp 357 363

CHANGES IN RIPARIAN vegetation ALONG THE COLORADO RIVER AND RIO GRANDE COLORADO

1 warren D snyder and gary C miller2

ABSIRACABSTRACT 1 changes in vegetation including area occupied canopy cover and maturity class of cottonwoodscottonwoods PopopulisPopupopuhispopithispithiskishis sppapp within lower elevation zones of the colorado river and clandecrandeglandegrande in rio in colorado weiewelewere monitored over 25 and37and 37 yeaiyealyear intervals respectively using photo methods interpretative estimated loss of cottonwoodscottonwoods along the colorado rivelriver was 2 hairnhakmhakrn 17517 5 and remaining stands had become more open and older cottonwoodsCottonwoods along the rio grande mclmcimelincreasedeased 161 6 baimhaimbaemhakmbakm 93939.39 3 with canopy and minor cover maturity class changes area occupiedoccupie d by shrubssh rubs and nverriver channel changed little along the colorado rivelriver but declined along the grande rio loss of hay meadow occurred along both aveisnveisrivers whereas developed land inciinclincreasedeased along the colorado river and farmland increased along the rio grande wildlife habitats the coloradodo deteriorated along Coloia much more rapidly than those along the rio clandeglandegrande during monitored intervals

tuomi keykeijwoids riparian colorado inventonjinventory cottonwood populusspppopulus sppapp wildlife habitat riverine systems in the great basin and melton et al 1984 awareness of these values southwestern united states are important hab has increased in recent years alongwithalong with concern itaas for itats resident and migratory wildlife ander- for increasing activities in and degradation of son and ohmart 1980 hunter et al 1985 two these critical wildlife zones windell 1980 major river systems colorado and rio grande these habitats are of special concern in moun- in the southwestern united states originate tainous areas because valleys are frequently within colorado while substantial work has narrow and centers of human activity been conducted to identify wildlife use and to before attempting to manage riparian vegeta- manage riparian habitats in lower reaches of tion for wildlife it is necessary to learn whether these river systems stevens et al 1977 ander- these habitats are declining in ability to sustain son et al 1978 anderson and ohmart 1980 species richness and abundance this paper swenson assesses 1985 and mullins 1985 little infor- recent changesehanchanbes and status of riparian mation has been published from studies con- vegetation along the rio grande and colorado ducted near the headwaters of these rivers river in southern and western colorado the cottonwood willow populus salix riparian ecosystem along colorados s colorado major STUDY AREA rivers has the highest wildlife species richness and density in the state beidleman 1978 fitz- lower elevation gerald 1978 hoover and wills 1984 and is used zones of the rio grande and colorado by 283 species of vertebrate wildlife however river in colorado were selected for study 1 1 most studies have centered on the south platte fig table I the colorado river and its tributariestributa ries drain about km2ki112 river in northeastern colorado graul and 46196 of western colorado ugland et al 1984 vol 2 bissell 1978 wildlife values ofriparian habitats the colorado river is confined to along streams and rivers in the relatively mountainous narrow valleys until it is joined by the gunnison western two thirds of colorado have received river near grand junction where the valley little study among ecosystems in mountainous broadens with reduced stream gradient areas cottonwood willow it riverriverbottomsbottoms usually leaves the state with flows approximately 75 possess high values for resident and migratory greater than at the upstream end wildlife of the study schrupp 1978 thomas et al 1979 area table 1

cocolliadocoloiadodo division of wildlifefwildlifi 306 2 Coloia cottonwood lane steilingsteistelling Coloiacoloradodo 80751 C ado pro Coloiacolliadocoloiado division olof wildlifeolwildlife 317w317 W prospectPio spectspeet roidroldroad fortfoit collins coloradoColoia do 80526

357 volume 52 358 GREAT BASIN naturalist

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GUNNISON RIVER GUNNISON MONTROSE

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inventoriesinventoriedoried and segments 1 I in western and south central fig 1 colorado river and rio grande with invent portions colorado

grande drains approximately of variables measured along the the rio tableTABLF 1 characteristics closed colorado river and rio grande colorado 19194 km2 ofwhich 7612 km2 is within a basin in south central colorado ugland et al rio grande variable colorado river 1984 vol 1 river flow originates primarily in from a the san juan range with lesser amounts Faf3f sampling interval arsyrs 250 368 11 enters the distance sampled km 1673 1174 the sangre de cristo range the river sample units 21 20 western part of the san luis valley a high 1632 I1 hasampleha sample unit 870 2286 2438 m park and travels 20 27 elevation sampling intensity approximately 100 km elevation m through farmed areas for upper 1829 2438 where most stream flows are used for irrigation 1372 2286 lower table 111 before entering a canyon that extends i daily stream flow m3sass upper loos1005loo100 5 25325.325 3 into new mexico lower 1755its175 5 70 harrington 1954 noted that narrownarrowleafleaf angustifolia dominate along grindegrande mdand from 1951571951 57 cottoncottonwoodswoods P aerial photos weicwelcwelewere froin 1941 to 1973831973 83 rio the 1 of 111to 1980olondorivli98 colorado rier the grande and upper portions col- of the rivernver channelch innelone rio 11 distancedist wasw is inasedinineincasedtsui cd atit thetiietile dentercenter linelinearlind me orado river whereas lanceleaflanceleaf cottoncottonwoodswoods 1992 COLORADO RIPARIAN vegetation 359 P acuminataacuminateacuminata occur sparsely over a slightly types k 1 oftotal area were omitted the area broader elevation range grande rio cotton per vegetation type was recorded to 00.1oi01 1 ha woods FP wislizeni 01 using dominate at lower eleva- an electronic planimeterplanometerplanimeter on site tions along the colorado inspections river willows are the were conducted within several plots along both primary shrubs along the grande rio and upper rivers to verify that photo interpretation was portion of the colorado river giving way to accurately assessing cottonwood stand tamarisk Ta maturity tamarixmatix fallicagalgallicalica at lower elevations canopy cover and vegetation types photo along the latter inter- plant names follow harrington pretationpretation accuracy approximated 95 195419541 maturity classes trunk diameter were esti- mated from tree crown size using photo inter- METHODS pretationpretation the relationship between trunk diameter and tree crown size was based on approximately 167 linkinkm ofthe colorado previous sampling of cottoncottonwoodswoods along the river south platte and 117 linkinkm of the rio grande were selected for river in morgan county colorado getter 1977 study and resrespectivelypactivelyectively stratified into four A close relationship rar2 8181.81 and between three segments strata based on empirical tree crown size and trunk diameter at breast height dm assessments ofvegetation area occupied by cot dahdbh was indicated however data relating dahdbh to tontonwoodswoods plot width etc fig 1 segments tree age were lacking as from increment boring to estimate age of cotton numbered upstream to downstream fifig9 1 woods did not yield were used to distribute random sample units notyield satisfactory age data matu- I1 rity classes included linear 161igi linkinkm river tracts more uniforuniformuniformlymy stands dominated by trees 1515151.5 1.515 4.040 4.141 76 along the rivers twenty sample units were dis- 1515151.515 40 41 767.6 and 76767.6 dindm dahdbh stands of trees tributedtributed along the rio grande whereas the were classified by canopy cover as open 10 35 colorado river study area contained 21 an intermediate 36 55 and closed 55 electronic planimeterplaniplanometermeter positioned at mid chan changes nel on US geological survey topographic in stands ofcottoncottonwoodswoods from early to maps was used to delineate the randomly recent photos were analyzed using paired selected km t tests appropriate for stratified segment sam- 161 river mile sample units I1 ples based width of sample units varied and was based on on the hypothesis that mean change was zero initial flood plain width primarily encompassing nat- tests included analyses of indi- vidual ural riparian vegetation readily discerned on maturitycanopy cover classes however sample aerial photos some adjacent cropland and sizes were inadequate to yield meaning- grassland were included ful results therefore maturity class data for pooled the earliest scale 120000 and most canopy cover classes and canopy cover recent data for scale 140000 aerial photos available pooled maturity classes are presented US in addition department of agriculture were acquired for early to recent changes were pre- sented where each sample unit to yield changes over time canopy cover and maturity classes were the same area was inventoriesinventinventoriedoried within each partitioned changes for other cover types were sample unit during both early and recent inter- analyzed using paired t tests ANOVA used vals to assess changes earliest aerial photos was to detect differences among segments mean were from 1941 and the most recent photos comparisons were considered significant 05os.05 were from 1973 through 1983 for the rio at P 05 grande those for the colorado river were from 1951571951 57 early and 1980 recent RESULTS interpretative analyses of aerial photos were contracted to the colorado state forest service colorado river vegetation types including trees primarily cot estimated loss of cottonwood stands along tonwoodstonwoods shrubs tamarisk colorado river the colorado river was 191.9ig hakmhaam sample and willow hay unit meadows grasslands agricul- 175 table 2 losses in the upper ture farmland developed segment roads towns etc fig 1 where cottonwoodscottonwoods initially averaged river standing water and vegetatedunvegetatedun sand- only 222.2 hakmhaam were 90 table 3 area bars were delineated on acetate overlays using occupied by cottonwoodscottonwoods was highest in seg- a stereoscope river and vegetatedunvegetatedun were ment 2 where they declined 444.4 hakmhaam within combined as river channel minor vegetation downstream segments cottonwoodscottonwoods averaged volume 52 360 GREAT BASIN naturalist

duringearlyanddunngduang early and recentintervalsrecent intervals alongthealong the colorado river 1 haam byvegetationland usense type tabietable I 2 area occupied 7 hakm7hakm by vegetationland and rio crandegrandeci ancle Coloiacoloradodo colorado river rio grande

eailyeallyearly recent early recent

SE X7 SE P type 7 SE 7 SE P 7 29 190 33 NS Cottoncottonwoodswoods 112 21 92 17 NS 174 18 101 21 NS 65 09 49 07 05 shrubs 95 03 147 29 112 31 NS 686 70 545 63 hayilayliay meadow 14 05 grassland 31 08 41 10 NS 09 06 31 51 26 NS 01 01 135 53 03 agriculture 55 16 NS 07 03 32 09 01 07 02 10 03 developed 03 01 channel 93 07 88 08 NS 62 04 39 rivelriver 12 03 NS standingstandmgwateiwater 01 005 23 08 03 10 03

cottonwoodswoods from early to lecentrecent sampling intervals along the colorado taniTABI 1 3 area occupiedsegmentoccupiedsegrnentoccupied segment 7 ham by cotton riverandreverandRivriveireveierandand rio grande colorado colorado river rio grande

eailyeallyearly recent early recent P 7 SE Y SE P segment 7X SE 7 SE 06 02 01 02 147 12 184 23 NS upper 23 35 NS 240 42 196 23 NS 299 30 322 middle 29 43 29 NS lowerlowel 78 31 74 22 NS 49 lowest 93 13 82 18 NS

10 of the total overall river about 757.5 939.3 hakmhaam and declined at more increased to among seg- modest rates channel changed little but variance channel widened in the eight percent of the cottoncottonwoodswoods along ments was evident the fifty and narrowed down- the colorado river were in the two younger two upstream segments stream maturity classes fig 2 the percentage of shrubs primarily tamarisk occupied 17 young trees dm dahdbh declined almost 50 18 of the sampled riverriverbottombottom and increased P oi0101.01 during the 25 year interval numbers slightly primarily in the second segment of large trees 76 767.6 dm also declined dramati- shrubs occupied only 191.9ig 252.5 hakmhaam within the 0202.02 cally P segment 12412.4 hakmhaam within the second of cottoncottonwoodswoods were similar upper hectares and third segments and 939.3 hakmhaam within the cover classes during the early among all canopy lower segment sampling interval however by the recent sample interval open stands increased 11 rio grande whereas intermediate and closed stands declined 42 FP oloi01.01 and 27 P 0505.05 Cottoncottonwoodswoods were moderately abundant respectively fig 2 within the upper segment of the rio grande hay meadow the most abundant vegetation increasing 373.7 hakmhaam 24924924.9 and were most middle segment where type along the colorado river declined 23723.7 abundant within the 3 1 during the sample interval table 2 with the they increased 232.3 hakmhaam tt77777.7 table fig absent within several downstream primary decrease occurring in the lower seg- they were estimated loss was 070.7ot hakmhaam ment grassland occupied 575.7 of the area sample units and cottoncottonwoodswoods occupied iti17117.1 during early year sampling but increased 31 13813813.8 initially area increasing to 188iss18.8 by the about 10 of the sampled area was in agricul- of the sampled second table 2 ture during both surveys developed land and survey 10410.4 of standing water were initially minor but small trees isls 151.5 dm represented 199211992 COLORADO RIPARIAN vegetation 361

5 TRUNK DIAMETER TRUNK DIAMETER 8 dm dahdbh dm dahe dbhe 76 4 1 7 417641 76 15 4 15 4 154040 15

6

3 ilelictid

W LU 4

W W W

2

0 EARLY RECENT EARLY RECENT EARLY RECENT 0 CANOPY COVER EARLY RECENT EARLY RECENT EARLY RECENT CANOPY COVER fig 2 early to recent changessamplechanges sample in maturity class and canopy cover cottonwoodsofofcottonwoodscottonwoods along the lower colorado fig 3 early to recent changessamplechanges sample in maturity class river western colorado and canopy cover of cottoncottonwoodswoods along thetiietile lower rio grande southern colorado the composition during both samples and increased 939.3 in occupied area fig 3 trees surveys river channel decreased 36736736.7 of intermediate size 15151.5 404.0 dm declined P throughout the study area area occupied by shrubs was 13 overtheovertoeover the 367 year interval livingwaygivingwaygivingivingwaytogivinggwaywaytoto the minor and estimated loss was 25 next larger 41414.1 767.6 dm maturity class that table 2 increased 27227.2 P iglg16.16 fig 3 this latter group dominated among maturity classes discussion during both surveys large trees 76tg 767.6 dm rep- resented only 3 of the total during both sur veys and showed little evidence of increasing in comparison of changes along the two rivers occupied area leads to greatest concern for habitats along the open stands initially occupied 31 of the colorado river the much larger of the two 1 timbered area and declined P 2525.25 to 25 table the 25 year interval along the colo- rado fig 3 in contrast stands of intermediate clo- river was considerably less than that for the grande sure increased P 0202.02 from 33 to 40 closed rio but a 17517.5 decline occurred in area occupied by stands increased modestly P 4949.49 9 repre- trees development along the river increased dramatically senting 35 of the total during both surveys and replaced fig 3 many stands of trees lack of natural hay meadows dominated among vegetation reproduction andor high mortality ofyoung trees types table 2 decreasing from 68 to 54 of was indicated by a 50 reduction in stands of young the sampled area declines occurred primarily trees along the colorado river reduction of stands dominated within the two upper segments initially grass- by old trees which provide habitat for land was minor but it increased primarily primary cavity nesting wildlife was also evident within the upper segment only 2 of 20 how- samples ever rapid shifts toward more originally contained cropland but the open stands propor- which indicated excessive mortality within tion increased to 9 of 20 samples oi01010.1 to 13413.4 stands were more discouraging than changes in developed land and standing water were maturity structure thus there were fewer and minor components in both early and recent smaller stands and those remaining were more volume 52 362 GREAT BASIN naturalist of the san luis valley open and occupied by intermediate maturity in the west central portion vol 1 however upstream classes ugland et al 1984 peak flows and of cottoncottonwoodswoods were especially dra- impoundments have reduced losses with stabilized increased vol- matic 90 in the upper segment where altered patterns late summer for irrigation flows occurrence was initially low expansion of urban umes into at alamosaalamona fig 1 averaged about areas highway construction and other develop- downstream of those at norte and average flows ments were responsible for much ofthe riparian 30 del have been about one half of those habitat loss in a relatively narrow valley that since 1930 in channel width possessed limited riparian habitat and from 1913 to 1930 reduction initially and stabilized stream rapid stream flows loss of trees to was indicative of reduced relatively dominated by peren- canadensis was noted and may flows Streamstreamsidessides were beaver castor which provides lim- especially in the upper segments nial herbaceous vegetation be important for establishment of many stands of cottoncottonwoodswoods were con- ited opportunity since such as cottoncottonwoodswoods and is fined to streamstreamsidessides by valley relief pioneering species of moderately stable and slow stream of tamarisk was evident along indicative expansion flows through the relatively flat san luis valley of the colorado river within a lower reaches farmland was the most pronounced and slower stream flows increased broadened floodplainfloodplain along the rio grande whereas of tamarisk severely limits land use change increasing expansion little development occurred for natural regeneration of cot opportunities shrubs primarily willows have not been tontonwoodswoods and willows russian olive elaeagnus major components along the rio grande in also is pioneering along the colo- angustifolia recent decades severe cold winters due to high rado species possesses a growth form river this 1 may prevent invasions of height and like tamarisk may elevations table of intermediate tamarisk which has developed as a streamside monoculturescultures Knopknopfandknopffandand olson 1984 form mono monoculture at lower elevations elsewhere the colorado river have stream flows along along riparian systems in the southwest rus- declines in recent decades not shown major sian olive was not yet invading the inventinventoriesinventoriedoried high elevation diver- large impoundments and grande riverriverbottombottom during the last 50 rio sions primarily occurring similar inventories of riparian vegetation and reduced peak flow years have altered changes and status were conducted along the colorado and gunnison sequences on the south platte and arkansas rivers in the high riversnversavers plains of eastern colorado snyder and miller the colo- extensive flooding occurred along 1991 deterioration of habitat along the arkan- in considerable rado river in 1983841983 84 resulting sas river was much greater than along western natural reproduction of seedlings however rivers in colorado however conditions along offset tthee infrequent flooding is not likely to the colorado river seemed to be deteriorating streamside impacts of stream flow regulation more rapidly than along the south platte river developments and invasions of exotic species was also much less riparian habitat along the there vegetation conditions and changes along western rivers making that which remained of the pat- colorado river appear to be following greater importance sampling of changes tern of disrupted recruitment of native riparian between two points in time may not give an phreatophytes occurring along many western accurate assessment of longtermlong term trends A rivers howe and knopf 1991 third inventory of these same sample units is in contrast to changes documented along the recommended in the near future colorado river riparian habitats along the rio grande were relatively stable during the sample acknowledgments interval with an increase in area occupied by cottonwoodswoods however several of the sample cotton of the colorado within the lower segment contained few or T E owens and D teska units performed aerial photo cottoncottonwoodswoods little evidence of seedling state forest service no sampling design establishment was noted subsequent to interpretation assistance in provided by D C increased stream flows during 1983841983 84 which and statistical analysis was graul and G R assisted concern for future trends stream flows bowden W D craig raises implementation A E averaged over 10 year intervals since 1890 in study designdesi9n and R W hoffman and showed little evidence of decline at del norte anderson C E braun 199219921 COLORADO RIPARIAN vegetation 363

R C kufeld provided constructive review of tamarix habitats in three southwestern desert riparian the manuscript this project was supported by systems pages 128134128 134 in R R johnson C D ziebell patton federal aid to wildlife D R P F ffolliott and R H hamre techni- restoration project cal coordinators riparian ecosystems and 152 their man- W R and the nongame checkoff program agement first north american riparian conference of the colorado division of wildlife USU S dedepartmentartmentartmont of agriculture gen- anicalenical forest service eral technicalteetec report RM 120 KNOPF F L AND T E OLSON 1984 naturalization of literature CITED russian olive implications to rocky mountain wildlife wildlife society bulletin 1228912 28929828999828912289298298998ggs ANDERSON B W AND R D OHMART 1980 designing MELIONMELTON B L R L HOOVFR R L mooneMOORFMOORE AND D J and developing a predictive model and testing a devegreveg PFANKUCH 1984 aquatic and riparian wildlife pages statedetated riparian community for southwestern birds 261301261 301 in R L hoover and D L wills eds manag- pages 434449434 449 in R M degraff technical coordina- ing forested lands for wildlife colorado division of tor management of western forests and grasslands for wildlife and USU S department of agncultureagricultureAgnculture forest nongame birds USU S department of agriculture service rocky mountain region denver colorado forest service general technical report INT 86 SCHRUPP D L 1978 the wildlife values of lowland river 1985 managing riparian vegetation and wildlife and stream habitat as related to other habitats in col- along the colorado river synthesis of data predictive orado pages 425142 51 in W D graul and S J bissell models and management pages 123127123 127 in R R technical coordinators lowland rivernver and stream hab- johnson patton C D ziebell D R P F ffolliott and itat in colorado a symposium colorado chapter the R H hamre technical coordinators riparian ecosys- wildlife society and colorado audubon council tems and their management first north american greeley conference U S riparian US department ofAgnagricultureculture SNYDERSNYDFH W DANDGD AND G C MILLER 1991 changes in plains forest service general technical report RM 120 cottonwoodscottonwoods along the arkansas and south platte ANDERSON B W R D OHMART AND J DISANO 1978 rivers colorado prairie naturalist 231651762316523 165176165 176 vegetatingrevegetatingRe the floodplamfloodplainfloodplamplainpiam for wildlife pages riparian STEVENS L RTR T BROWN J M simpsonandSIMPSIMPSONSONANDAND R RJOHNR JOHN 318331318 331 R R in johnson and J F mccormick tech- SON 1977 the importance of riparian habitat to nical coordinators for strategies protection and man- migrating birds pages 156164156 164 in R R johnson and agement of floodplainflood plain wetlands and other riparian D A jones technical coordinators importance pres- ecosystems USU S department of agricultureAgnculture forest erervation and management of habitat general riparian a sym- service technical report W 0 12 posium USU S department of agriculture forest BEIDLEMANBFIDLEMAN R G 1978 the cottonwood willow ripariannpanan service general technical report RM 43 ecosystem as a vertebrate habitat with particular ref- SWENSON E A AND C L MULLINS 1985 revegetatingRevegetating erence to birds pages 192 195 S 192195 in W D graul and J riparian trees in southwestern floodplainsfloodplains pages 135- bissell technical coordinators lowland rivernver and 138 inw R R johnson C D ziebell D R patton P F stream habitat in colorado a symposium colorado ffolliott and R H hamre technical coordinators chapter the wildlife society and colorado audubon riparian ecosystems and their management council greeley first north amerleanamerican riparian conference USU S depart- fitzgerald J P 1978 vertebrate associations in plant ment ofAgnagricultureculture forest service general technical communities along the south platte river in northeast- report RM 120 ern colorado pages 738873 88 min D graul S W and J THOMAS J WCW C MASERMASFR AND J E RODIFKRODIEK 1979 bissell technical coordinators andaandj ripar- lowland rivernver and ianlan zones pages 404740 47 inm J W thomas ed wildlife stream habitat in colorado in a symposium colorado habitats in managed forests the blue mountains of wildlife chapter the society and colorado audubon oregon and washington USU S of council department agricul- greeley ture forest service agricultureAgn culture handbook 553 GETTER J R 1977 procedures for inventoryinginventor plains ying UGLAND R C J T steinheimerSTEINHEIMFR J L BLAITNFRBLATTNER AND cottoncottonwoodswoods morgan county colorado CJ colorado R G KRETCHMANKRFTCHMAN 1984 water resources data state forest service report 44 col- appp orado water year 1983 vol 2 colorado river basin GRAUL W DDANDSAND S BISSELL TECHNICAL COORDINATORS J technicalcoordinators1echnicalcoordinators above deloresdolores river USU S geological survey 1978 water lowland rivernver and stream habitat in colorado a data report CO 83 2 symposium colorado chapter the wildlife society UGLAND R C J T steinheimer J L BLAIFNFRBLATTNER AND and colorado audubon council CJ greeley R D STEGERSTFGER 1984 water resources data colo- harrington H D 1954 manual of the plants of colo- rado water year 1983 vol 1 missouri river basin rado sage books denver colorado 666 appp arkansas river basin and rio grandegi cinde basin USU S geol- HOOVER R L AND D L WILISWILLS EDS 1984 managing wllis ogical survey water data report CO 83 1 forested lands for wildlife colorado division of wild- WINDELL J T 1980 colorado s water resources uses life and USU S department of Agnagricultureculture ser- forest abuses and current status proceedings annual meet- vice rocky mountain region denver colorado 459 appp ing ofthe colorado wyoming chapter americanamerlean HOWE W H AND F L KNOPF 1991 on the fish- imminent eries society 154757154715 475747 57 decline of rio grande cottonwoodscottonwoods in central new mexico southwestern naturalist 3621836 21822421836218224224 HUNTER W C B W ANDERSON AND R D OHMART received 28 january 1992 1985 summer avian community composition of accepted 20 september 1992 creatgigreateat basin natmalist524naturalist 524 appp 364372364 372

RESIDENT UTAH DEER hunterspreferencesHUNTERS preferences FOR management OPTIONS

1 1 2 dennis D austin philip J urness and wes shields

distributed at checking stations in fall 1989 ABSAHSITRACiAraciHACI A total 013291of 3291 lesiesresidentident deeldeer hunters returned questionnaires crowding and and 1990 providing opinions and management data concerning the utah rifle hunt hunters reported hunter and too fewfiew big bucks as critical reasons foifolfor possibly choosing to quit deer hunting in utah indeed hunter age structure of several ineaineasuredmeasuimeaskisureded satisfaction suggested a negative future trend in hunter participation results suggested the adoption huntcrprcferrcdhuntel piefened management options would inciincreaseease satisfaction motivation and success

methods keokelikeifkey words nude leericerdeerdeen questionnairesquestionliresmires checkingstationschecking stations leerlehrdeerdehr management hunter opinions wildlife wildlife techniques

competition for wildlife recreation in the tive communication between state wildlife offi rocky mountain region will increase inm the elais and utah hunters future while projected populations of major one means of communicating information is which wildlife species will show little change in the through hunter opinion questionnaires have become an important data source for game next 30 yeaiyealyears s the number of big game hunters management decisions in utah during the is expected to slowly increase from about 151.5isls1 5 to 1980s six questionnaire surveys were con- 171.71 7 million compared with the rapid increase in ducted and that number will likely double in nonconsumptivenon users of 39 to 71 million consumptive the 1990s bunnell and austin 1990 the use and hoekstra 1989 certainly the per- flather of surveys distributed to the total population will postcard questionnaire centage of hunters in homeward bound hunters at deer checking sta- decline while the percentage of nonconsump tions is one method this simple technique will consequently to bal tive users increase developed in utah during the late 1980s is wildlife managers must ance resource use inexpensive demographically unbiased and obtain a clear understanding of user prefer- accurately representative of hunters opinions ences particularly among those users who his- concerning deer management austin and toritoricallycally and currently have paid most jordan 1989 austin et al 1990 management costs via license permit fees and excise taxes on sporting equipment METHODS in utah mule deer are preeminent among hunted wildlife species min terms of income Questionnaires3 were printed on 4 14 X received for wildlife management and hunter 6ginchinch postage paid cards during opening days afield however compared with the 1970s weekend of the 1989 utah rifle deer hunt 7040 and in contrast to past regional trends flather questionnaires were distributed to hunters at 11 and hoekstraHoekstia 1989 total big game license sales checking stations and in 1990 8750 question- decreaseddecieased slightly 08080.80 8 in the 1980s while naires were distributed at 16 locations one total rifle hunters afield declined 31 jense questionnaire was given to each licensed hunter and shields 1990 these figures warn ofpossi checked until the supply was exhausted ble negative trends for deer hunter participation data were analyzed within years using the and along with uncertain hunter satisfaction pearson chi square statistic the cross tabula- strongly suggest a need for constant and effee tion method from the SPSS program on a VAX

dpartnuntD paitinriil of nalurange selensciensekSLK nerncrnee utah statstalestatestalp uniumveisitrity logan utah 84322 5230 butah2ulall2utah division ofwildlifol01 wildliliWildliliilicilie rsourkcsoniccsraour s 159615 wistwest north tn11cteinpliTeinpli salt laklake city utah ml8411616 33opks oftheodthe questionnaircquestionnaire codscmds are avajlabk froiii the iiior mithormishor

364 199211992 HUNTERhuWTER OPINIONSOPINIC NS ON dlDEERLER managementMANAGISMENT 365

tableTABLF 1 questionnairereturdque stionnanestionnauenane return ratesrates

1989 survey 1990 survey no no no no region location distributed returned retuifetuireturnedned distributed returned returnedi etui ned

northern Snowsnowvilleville 500 107 214 500 109 218 blacksmith 500 95 190 500 83 166 ogden 1000 151 151 500 126 252 kamas 500 91 182 wellsville 72 20 278

salt lake canyons 250 48 192 nor northeasternthetheastelasteiastel n veinalvernal 666 44 66 500 114 228 bookcliffsBook cliffs 650 131 202 500 113 226 current ck 500 120 240

central thistle 1000 206 206 125012 50 291 233 tucker 300 37 123 sheepcreekSheepcreek 300 48 160 250 41 164 vernon 452 151 334 375 95 253 stansburyStansbmy 300 57 190 southeastern areawide sio850830810 95 114 south central fishlikefishlakeFishlake 300 14 47 oak creek 195 63 323 southern bloomington 1600 423 264 1500 418 279

total 7040 1413 201 8750 1878 215

computer was used for question number 18 25825825.8 and 1988 2012012020.11 expected statewide 1990 survey the values given were objectively return rates usinusingetleti this method are thus about placed into 11I1 I1 monetary classes for analysis the 20 25 significance level for all data interactions was set at P 05os05.05 for question number 20199020 1990 survey hunter demographics success because the question was written with innumer- and satisfaction able potential responses and many question- most resident hunters male 90 naires contained more than one response data are age 25 444 52 and have than 10 of were not statistically analyzed but are reported 254 more years utah deer hunting numerically the responses were subjectively experience 60 during 1989 and 1990 hunters had less than 50 grouped into 71 categories data from the 1987 party success for bucks on and 1988 surveys are added and opening weekend and rel- compared low where applicable atively hunter satisfaction table 2 hunter party success noticeably declined between 1987 and 1988 and again between 1988 and 1989 but RESULTS remained about the same between 1989 and 1990 return rates the percentage of hunters 20 in the checking stations used for distribution by youngest age class 14 24 years is lower than utah division of wildlife resources regions expected participation by hunters in this age the number of questionnaires distributed and class should be highest because few people return rates are shown in table 1 although begin hunting after about age 25 these figures rates varied considerably by region and location consistent over four years alone suggest possi- total return rates were 20901golgoi20120.11 in 1989 and 21521.5 ble future declines in the number of utah deer in 1990 and consistent austin and jordan 1989 hunters however the sharp drop in hunter austin et al 1990 with those reported for 1987 participation between the third and fourth age 366 GREAT BASIN naturalist volume 52

1987 1990 tolfTAILF 2 demographics party success and hunter satisfaction of utah resident deer hunters sampled sample sizes in parentheses

a1 sex age class year male femafemaleie N 1 2 3 4 5 6 N

1987 904 96 863 1901igoe 318 230 142 81 40 869 1988 896 104 444 197 330 234 138 83 20 458 1989 928 72 925 186 281 259 139 90 46 936 1990 927 73 1429 220 266 262 129 82 41 1429 means 914 86 198 299 246 137 84 37

experience class b

0 d year 1 2 3 4 N success satisfaction

1987 187 213 272 328 867 69693f3 411 53 871 1988 217 184 284 315 461 553 459 43 456 1989 227 152 252 368 932 480 904 41 934 1990 251 142 260 346 1418 479 1406 46 1413 means 221 173 267 339 551 46

age classes 1 144 24 2 25 34 3 35 44 4 45 54 5 55 64 6 65 years fxpencnce classes 1 1 5522 6 10 3 11 20 4 21 years 11 weekend ilhunting party success foi one 01or moiemole bucks on opening A score 5 0 would be expected for the average hunt iiuntei1 lo10 sllslismisatisfactionfaction of currentcorrent yeaiyealyears s hunt inin companioncomparisoncompansoneomcompanson to all previouspieviouspi evious deer huntsbunts of50of agengo class lh16 24 I1 lanteiluntei11iinters s aged 14 and 15 years were ineligible for big gainegame licenses alo harvested rootinghunting party success toiloifolfor buckshocksbocks and antleantieantlerlessilessliess deerdeel on thistins hunt for the 1987 season 6251662 516 bucks and 1168 antlerless deer were classes 35 44 and 45 54 is also of concern another concern for hunter participation is because in these age groups many hunters chil- noted by comparing the trend of hunter partic- dren are beginning to hunt and parent partici- ipation by experience classes between survey pation is a key factor in longtermlong term sustained years table 2 no trends in hunter participa- interest of new hunters decker and connelly tion were evident for hunters with 11 or more 1989 mean age of all hunters was 36336.3 35435.4 years of experience however hunters with 6 37037.0 and 36036.0 years for 1987 90 respectively 10 years of experience decreased ti717.17 1 between in a completely randomized survey of utah 1987 and 1990 while hunters with 1 5 years of hunters krannich et al 1991 reported a mean experience increased 646.4 of 37 and similar hunter age and sex age years between hunt types characteristics comparison one probable explanation for the sharp drop utah has had four basic types of hunts since in hunters in the 45 54 age class is the signifi- 1951 with each hunt type having a variable cant interaction between age and hunter expe- number ofantlerless control permits either sex rience with hunter satisfaction P 0404.04 hunts dominated from 1951 to 1973 with buck hunters with 20 years of experience who gen- only hunts dominating from 1974 to 1990 as erally hunted deer before the 1970s when the well as before 1951 from 1985 to 1990 hunter number of hunters was lower fig 1 and number restrictive limited entry and high hunter success rate was higher jense and country hunts and from 1984 to 1989 shields 1990 show lower satisfaction scores antler restrictive three point and better hunts than younger less experienced hunters mean were established on some units satisfaction scores of experience classes 1 3 BUCKONLYBUCK ONLY HUNTS total buck harvest versus 4 table 2 for both years combined werewere averaged 63250 per year with 8633 antlerless 454.5 and 393.9 respectively similarly mean satis- harvest and 181235 hunters afield fig 1 the faction scores of age classes 1 3 versus 4 were number of unretrieved deer reported per 100 454.5 and 373.7 respectively consequently hunting buck only hunters in these surveys for 1987 90 motivation for hunters with 20 years of experi- was 19919.9 21721.7 15915.9 and 16016.0igo respectively ence has likely decreased because of perceived using the weighted mean of 17917.9 total lower quality hunting unretrieved deer for this period was 32441 per 199211992 HUNTER OPINIONS ON DEER management 367

250000 T BUCK

0 DCE

HUNTERSHUMTERS 200000

150000 X

100000

50000 0olaotala

0 c0ca LO r 0 M LO r 0 0

fig 1 total harvest of buck and antlerless deer and combined hunters afield from all buck hunts in utah 1951 90

year and mean total annual hunting mortality decrease in the small buck two point and less was 104324 mean hunter satisfaction 1987 to doe ratio between preseason and postseasonpost season 90 with 0 representing the worst hunt and 10 classification counts jense 1990 the best hunt was 444.4 hunting party success our analysis confirmed the adverse impacts was 45845.8 of three point and better hunts reported by EITHERSEXeltherEITHER SEX HUNTS during 23 years of jense 1990 with the highest number of either sex hunting the statewide total buck har- unretrieved deer at 39639.6 per 100 hunters vest averaged 66992 and the antlerless harvest including 21721.7 bucks this number of bucks was 39228 using the estimated mean for mostly two point and less is compared to 464.6 unretrieved deer robinette et al 1977 stapley bucks per 100 hunters on buck only areas how- 1970 of 808.0so deer per 100 hunters and the mean ever hunters from antler restrictive areas were number of rifle hunters afield 153666 a cal- moderately satisfied with a mean index of 48 culated yearly loss of 12293 unretrieved deer is and mean hunting party success was 55655.6 obtained bringing the mean total annual hunt- during 1989 the last year of three poinpoint taand ing mortality to 118513 hunter preference for better hunts 40040.0 n 931 of utah resident buck only versus either sex hunting has not hunters had hunted at least once on three point been addressed and better areas but only 26726.7 n 906 pre- ANTLER restrictive HUNTS three ferred to continue this type of hunt indeed less point and better antler restrictive hunts were than half 47747747.7 of hunters who chose to hunt available on some units during 1984 89 and these units in 1989 preferred to continue them then discontinued in comparison with buck even though antler restrictive hunts were only hunts three point and better hunts not successful over entire deer management showed a reduction in hunters afield buck har- units selection of conscientious hunters to vest and hunter success jense 1990 however avoid high unretrieved deer losses may lead to these hunts also showed a small increase in the successful antler restrictive management for postseasonpost season total buck to doe ratios but a large example at the east canyon resort 10000 decrease in the number of postseasonpost season mature acres in northern utah protecting only 22x2X 2 bucks counted these areas also showed a large point bucks 1988 90 increased the mean 368 GREAT BASIN naturalist volume 52 number of total antler tines of harvested bucks restricted areas in terms of size locations and from 454.54 5 1985 87 to 616.16gi 1 1988 90 the per- number of areas of harvested bucks 2 X 2 or smaller cent license fees decreaseddecidecl eased from 60 to 35 while the number of trophy bucks larger than 4 X 4 increased from with the current cost of a big game hunting 0 to 8 unpublished data 44east canyon resort license set at 1500150015.00 hunters were asked what HUNTER NUMBER restriction HUNTS they believed to be the fair value although limited entry hunts have been used on some schreyer et al 1989 reported increased units since 1985 in comparison with buck only license fees were opposed by most hunters a hunts they provide higher hunter success mean value of 159015.90 was determined n FP 01 and satisfaction FP 001 with an 1391 in our study most hunters 58858858.8 indi- index of 636.36 3 but no difference in the total cated isoo150015.00 was the fair value sixty eight hunt- number of unretrieved deer 177 total deer per ers 49494.9 indicated the fair value was 3000300030.00 or 100 hunters with 919.19gi 1 bucks and 868.68 6 antlerless more while 58 hunters 41414.1 indicated the hunting party success 1987 90 was high at value was less than 1000100010.00 it was interesting to 68868.868 8 in 1989 2282222.88 of resident hunters n note that costs were not related to hunter suc- 935 had hunted deer on limited entry areas cess satisfaction hunter choice of hunt type or and most 65665665.665 6 indicated the fee of 9900220022.0022 00 whether private or public lands were hunted was fair while most hunters n 908 favored although license fees are strongly and the samesarne 37837837.837 8 or increased 38938938.938 9 number broadly approved by utah hunters few of limited entry units hunter preferences for improvements in the quality of the deer hunt various permit drawing and landowner hunting can be made without the economic tradeofftrade off of options were unclear increased hunter fees hunter preferences for A second type of hunterhunteante r number restrictive balancing potential increased fees with hunt is the high country hunt this uncrowded increased hunt quality need to be defined high quality hunthulithullthurit but one that harvests bucks hunter concerns not then available during the october rifle hunt received positive support from most twentyfivetwenty five categorical responses were 59659659.659 6 utah hunters given by 1 or more hunters as reason to quit deer hunting table 3 although the list con- vehicle access to public lands tains several areas of low management influ- ence such as old age high associated costs of A strong majority of hunters 76276276.276 2 indi hunting and personal attitude most areas of batedcated that at least some lands should be closed responses are influenced by management deci- to vehicle access during the deer hunt to sions the most common reasons directly influ- increase the quality of the hunting experience enced by management decisions included too however the percentage of hunters indicating many hunters too few deer bucks and big at least half cfallofallof allaliail public lands should be open to bucks private land problems and poor game vehicles was 74574.574 5 overall hunters indicated management that a mean of 37537.537 5 of public lands should be closed to vehicle access varying by location discussion from 2892828.99 to 45445.445 4 the percentage of hunters who hunted on areas with vehicle restrictions reasons to quit deer huntinghuntin9 was 33833.833 8 while the percentage of hunters who indicated preference to hunt on areaswithareaareas withswith cehivehi the proportion of mature bucks in the har- cle restrictions was 45245.245 2 using the logical vest is an area of management control it is clear assumption that the percentage of areas most hunters prefer harvesting large bucks restrictedrestncted to vehicles should be closely proporpoopor infrequently as opposed to harvesting smaller tionaldional to the percentage of hunters preferring bucks frequently austin et al 1990 as well as themthern our data suggest the current amount of reducing some hunting opportunity to increase area with restricted vehicle access is close to the proportion of mature bucks in the harvest hunter preference but that an additional 373.73 7 austin and jordan 1989 toweill and alienallenailen 37537537.537 5 33833.833 8 to 11411.411 4 45245245.245 2 33833.833 8 ofpublic lands 1990 furthermore with the hunting media should be restricted more information is emphasis on trophy bucks the potential harvest needed on hunter preferences for vehicle of mature bucks adds considerably to hunter 199211992 HUNTER OPINIONS ON DEER management 369

TABU 3 utah residentlesiesident deer hunters responses to the question if you were to quit deerde ei hunting in utah what leasonreason would you list

number of questionnaires i eturnedreturned 1430 number of questionnaires with no response 88 number of questionnairesa4thquestionnaires with would not quit none 46 number of questionnaires with responses 1296 number of total responses 2087

number of response categories responses hunters too many hunters 479 370 too few deeldeer 199 154 private land problems 164 127 too few big bucks 122 94 old age or physical impairment 108 83 high associated costs of hunting 83 64 no areas to hunt or access to public lands 81 63 too few bucks 79 61 poor game management 75 58 unethical hunters 72 56 low success or no limit on statewide license sales 63 49 children aged 14 and 15 years can hunt 48 37 deer are too small 44 34 too much ATV use or too many road hunters 41 32 safety 39 30 high costs of licenses 35 272 7 personal attitude 33 25 too few vehicle access roads 31 24 too many nonresident hunters 30 23 poor hunt quality 29 22 proclamation too long or complicated 27 21 no either sex or antielantler restriction hunts 19 15 too many limited entry areas 17 13 too few limited entry areas 16 12 too many does 14 111 1 46 otherothel categories 139 107

motivation and krannich et al 1991 reported the authors believe a reasonably high per- that about two thirds of hunters 66366366.3 were centage 20 40 of mature bucks in the har- dissatisfied with the size of bucks vest is critical to successful deer management compared with either sex hunting age and hunter motivation it is clear to us that structure ofthe male population declines under decreased hunting pressure on the buck popu- buck only hunting mccullough 1979 in utah lation is necessary the data strongly suggest a austin 1991 the percentage of mature bucks need to establish statewide minimum standards age 3 12 years and older harvested decreased for 1 age structure of the buck harvest 2 from about 44 during the pre 1951 buck only post season buckdoebuck doe ratios and 3 hunter suc- hunts to about 30 during the period of either cess for bucks sex huntingbunting 1951 73 the percentage of problems associated with private lands are mature bucks harvested sharply decreased and important to hunters these problems include has remained at about 10 during the period of poorly marked lands trespass private lands cur- reestablished buck only hunting 1974 90 on tailing access to public lands and depredation limited entry hunts the percentage of mature private lands provide deer hunting for 14814.8 bucks in the harvest has exceeded 30 on most 1990 survey of utah resident hunters and units not only has size of harvested bucks 14714.7 of hunters reported owningowninor 10 or more decreased due to decreasing mean age but age acres used by wildlife 1989 survey one possi- specific size has also declined austin et al ble partial solution may be to give landowners 1989 more flexibility in management by allowing 370 GREAT BASIN naturalist volume 52 either sex hunting on private lands advantages the lack oleitherofeitherof eitherelther sex hunts and too many does include increased landowner control over deer especially since changes to buck only manage- numbers on their lands decreased unretrieved ment were made beginning in 1974 have merit deer kill austin et al 1990 reduced depreda- hunter crowding before about 1969 when tion complaints and improved opportunity for license sales were less than 180000 fig 1 was harvest furthermore liberal hunts on private probably a much smaller problem bureau of lands may increase incentives for landowners to government and opinion research 1971 mark their boundaries and allow additional however the crowding problem of increased hunting opportunity human population and finite resources leo- the categories of unethical hunters safety pold 1930 has been exacerbated because of the and minimum age for hunters are closely related longtermlong term leopold 1919 and more recent to hunter education courses since the begin- increasing urbanization closures of private ning of the hunter education program 1958 lands to public hunting and increased vehicle and the required wearing of hunter orange access on both private and public lands mann clothing 1973 the mean number of total utah 1977 reed 1981 hunting accidents and fatalities per year has our findings indicate the majority of hunters averaged 111illiiiliilil11.111 1 and 343.4 respectively with about prefer reduced hunting opportunity for higher three accidents and one fatality occurring quality when hunters were asked to indicate during the rifle hunt before about 1958 when the effect of crowding on their hunt quality neither hunter education nor hunter orange was using an I111 I point scale where 0 means crowding required over 100 accidents and about 20 fatal greatly decreased the quality and 10 means itiesaties occurred yearly from all hunts combined crowding had no negative effect only 27827.8 of hunter preference to allow persons aged 14 and hunters scale 8910 indicated crowding had has 15 years to hunt big game not been little effect compared to 60260.2gog of hunters addressed scale 0 5 who indicated a large effect x the length and complexity of the proclama- 4924.92 krannich et al 1991 reported 71 of 1979 the tion is a concern of hunters before hunters believed there were too many hunters deer measured one page utah proclamation in their areas crowding effects were not signif- 175 X 22.5225 inches and on high 17517.5 225 was printed icantly related to hunter age sex years of expe- with the rules and regulations on quality paper rience unretrieved deer reported or whether side and a multicolored map of utah s deer one hunters were on private or public lands surpris- on the 1990 the units reverse in newsprint ingly the means for hunters from successful proclamation sheets were close to the same size 5045045.04 and unsuccessful parties 4964964.96 were not 14514.5 X 230 inches but contained six pages 145 23023.0 different these data indicate the effects of quality of the hunt in terms of the ratio the crowding are felt by almost all groups equally of deer or bucks harvested per hunter is con however hunters from limited entry areas trolledstrolled by management although management P 002002.002 where hunter numbers are limited can alter the buckdoebuck doe ratio the total number of rated the effect of crowding less negatively x deer is limited by habitat and conversely hunt- 6.16gig while hunters preferring to hunt in areas ers have not been numerically limited the utah 616 from vehicles P ooi001001.001 buck harvest has remained rather constant restricted were more negatively affected x 4614.61 than hunters pre- mostly 50000 8080000000ooo since 1951 fig 1 while the antlerless harvest has sharply decreased ferring no restrictions x 5555505.50 since 1974 with the resumption of buck only management options to reduce hunting total buck hunters afield from all com- hunter crowding bined hunts increased steadily between 1951 and 1964 decreased for three years 1964 67 several options are available to reduce slowly increased during 1967 69 but abruptly hunter crowding split deer hunting seasons increased between 1969 and 1973 after a were opposed by utah hunters in recent studies second three year period of decreasing hunters krannich and cundy 1989 austin et al 1990 afield 1973 76 hunter numbers have fluctu- krannich et al 1991 this option would likely ated but remained high throughout the 1970s increase hunting pressure on bucks by and 1980s consequently the hunter responses increased hunter days longer seasons and of poor game management poor hunt quality hunting during the more vulnerable rutting 199219921 HUNTER OPINIONS ON DEER management 371 period it would thereby further decrease mean weigh the values of increased wildlife manage- age and size of harvested bucks ment income and hunting recreation opportu- A second option is to require hunters to nity indeed hunter responses from these choose either a buck or doe tag our survey surveys confirm our view that hunting pressure indicated 78478.4 of resident hunters would on bucks should be reduced to the pre 1970 choose a buck tag which would reduce buck level hunting pressure by about 21621.6 A third option is to require hunters to choose and hunt only one season since mean hunters literature CITED afield for 1988 89 combined were archery 26613 rifle 180298 and muzzleloadermuzzleloader AUSTIN D D 1991 age specific antler tine counts and 8832 this option would reduce crowding during carcass weights of hunter harvested mule deer from the rifle hunt up to approximately 20 assum- utah checking stations 1932 1988 division of wild- ing hunter proportions remained about the life resources salt lake city utah 35 appp same hunters favor this option in our 1989 and AUSTIN D D S D BUNNFLLBUNNELL AND P J URNFSS 1990 responses of deer hunters to ques 1990 surveys 63.8638 and 64064.0 respectively in a a checking station ques- 638 tionnairetionnaire in utah western associatioassociation of game and 1990 completely randomized telephone survey fish commissioners proceedings 6920869 208 229 of 14305 deer hunters 58058.0 of utah hunters AUSTIN D D AND L JORDAN 1989 responses of utah indicated preference for this option krannich deer hunters to a checking station questionnaire great et al 1991 reported a similar level of support basin naturalist 4915949 159 166 AUSTIN D D R A RIGGS P J urnessUHNFSS D L TURNER 6.19619gig a scale of 0 10 DR PJ mean score 619 using ANDJFAND J F kimKIMBALLBALL 1989 changes in mule deer size in probably the most effective option to perma- utah great basin naturalist 493149 31 35 nently reduce hunter crowding while at the BUNNELL S DANDD ANDDD D AUSTIN 1990 Hunterhunteropimonshunteropinionsopinions and questionnaires pages 10 47 87 in deer manage- establishing a standard for in same time minimum ment workshop june august 1990 utah division of quality in terms of hunter pressure on bucks is wildlife resources salt lake city 94 appp to limit license sales of buck tags hunters con- BUREAUBUREAUOF OF GOVERNMENTgovernmentandAND OPINION researchRFSFARC II11 1971 sistently favor this option in our 1990 survey opinion survey of the people of utah for wildlife resources and outdoor utah state univer- 60.6606gog of resident hunters preferred to limit recreation 606 sity logan 73 appp buck license sales to 150000 with up to 35000 DECKERDFCKER D J AND N A CONNOLLY 1989 motivations antlerless tags available to unsuccessful buck tag for deer hunting implications for antlerless deer har- applicants 39439.4 favored unlimited license vest as a management tool wildlife society bulletin sales since hunters who favored limiting license 1745517 455 463 FLATHFRFLATHER C H AND T W HOEKSTRA 1989 an analysis sales also favored to choose sex of having tag of the wildlife and fish situation in the united states P 004004.004 most hunters would favor having to 1989 2040 USDA rocky mountain forest and range choose sex of tag krannich et al 1991 deter- experiment station general technical report RM mined most hunters 61761761.7 supported choos- 178 147 appp JENSF G 1990 three point strategy in utah pages 78 88 sex and yearly harvest in ing the of tag havincrhaving in deer managermanagementnent workshop june august 1990 restricted to one deer per hunter utah division ofwildlife resources salt lake city 94 appp in the 1989 survey only 36636.6 of hunters jenseJINSFJENSF G K ANDWAND W SHSHIELDSIELDS 1990 utah big game annual publica- indicated preference to hunt every year regard- repoidepoireportt utah division of wildlife resources tion no 90790 7 less of future growth in hunter numbers while KRANNICHKRANNICH R S AND D T CUNDY 1989 perceptions of the majority 63463463.4 selected some level of crowding and attitudes about a split deer hunting hunter number limitation austin et al 1990 season among resident and nonresident utah deer hunters preferring the limitation 38238.2 hunters institute for social science research on nat- of ural resources utah state university logan 29 appp selected the limit at 160000 and 25225.2 selected KRANNICH R SSJJ S KEITH ANDVAND V A RIIFARHEA 1991 utah the 200000 limit previously in 1987 55855.8 of deer hunters opinions about deer hunting and alterna- hunters showed preference to limit hunters to tive season formats project summary report institute less than 200000 austin and jordan 1989 for social science research on natural resources utah state university logan 75 appp it is apparent to the authors that some LEOPOLD A 1919 Wildwddliferswildliferslifers vs game farmers a plea for restrictions are needed we believe the democracy in sport AGPA bulletin pages 54 60 in increased buck hunting pressure beginning in D E brown and N B carmony 1990 aldo leopoldleopoldss wilderness stackpole books harrisburgHarnsburg 1970 1 has had effects on hunter fig negative pennsylvania 249 appp success satisfaction motivation and harvested 1930 game management in the national forests buck size these negative effects appear to out american forester july 412 414 372 grestGREAT BASIN naturalist volume 52

CUNDY 1989 MANN 1 K 1977 land use and acquisition study on SCIIKFYFR RRRR S KRANNICIIKHANNICIL AND D T wildlife langesranges utah division of wildlife resources public support for wildlife lesourcesresources and programsprogiamslams in joijotob completion reportrepoitwW 65 11 D 25 jobjobaA 8 270pp8270pp utah wildlife society bulletin 1753217 532 538 loss 1 1 STAPLEY kill and wounding mctjllotjiiM cm ou ll11 D R 1979 the george reserve deer SIAPI FY H D 1970 deer illegal herdhelhei d universityuniveiuniver ritysity of michigan piessplesspress ann arborarbon 271 appp final report utah division of wildlife resources 1 publication 1 7 1 1 65 R A 8 rlrr i D D F 19819s11 conflicts with civilization pages 509 536 W appp S fromhormhomm inMI 0 C wallmo ed mule and black tailed deer of toweill D E AND T ALLENALLFN 1990 results horn the northnoi th ainencaamerica university of nebraska presspi essegg lincoln 15 year policy plan questionnaire idaho department 605 appp of fish and game boise 14 appp roninROBINEITEROBIN 1 rn W L N V IIANWCKHANCOCK AND D A lonesJONFSJONES 1977 the oak creekcreckci eek mule deer heldherd in utah utah division of wildlife resources publication no 771577 15 148 appp received 17 december 1991 accepted 3 november 1992 great basin naturalist 524 appp 373378

LIST OF OREGON scolytidae coleoptera AND NOTES ON NEW RECORDS

1 2 malcolm M furniss james B johnson richard L westcott and torolf R torgersen 3

ABSTRACT listed are 121 species of scolytidae from oregon ten species are reported from oregon for the first time hylastesHylastes tennis eicuioffphloeosinusscopulorumscopulorumeichhoff phloeosinus scopulorum scopulorurn swaine phloeosinus hoferhofenhopen Blackmblackmandn trypodendron betulaehetthetibetthetiilaeflaeilaefiah swaine xyleborus xylographxylographusus say trypophloeus striastnatulusstriatulustulus mannerheimManmannerheimonerheim T thatcherthatchenshatchenthatchen wood procnjphalusprocryphalus inucronatusmucronatusmucronatus leconte pityophthorus11ityophthorus sculptorscalptorscalp tor blackman and MonaTmonarthnimmonatthrumthrum dentigerurndentigerumdentigerum leconte the second oregon specimen of an exotic species xyleborus califorcalifornicuscahfomicusnicus wood is reported also

kegkeykhykeywordswords scolytidaescolytidaefaunalhstfaunal list oregon

oregon is a large state with diverse vegeta- found by further collecting they likely will tion that occurs there due largely to the wide include more pityophthorus a genus that is range of physical and climatic environments relatively rich in species in western forests and the climate results in part from the interplay elsewhere and species of other genera from the between maritime and continental air masses diverse california fauna blightbright and stark 1973 and the intervening cascade mountain range that infest trees endemic to both states that divides the state into distinct western mar- other new scolytids are likely to be intro- itime and eastern continental regions frank- duced accidentally by commerce for example lin and dyrness 1973 for example average the exotic xyleborus alfinisaffinis eichhoffeichhoffwaswas inter- annual precipitation varies from approximately cepted in 1961 at portland in dracaena 60 300 cm west of the cascades to 20 100 cm massangeanamassangeana from puerto rico of the 121 spe- eastward cies listed here 8 are clearly exoticsexotica that have the exceptionally diverse forests of south- become established at unknown times western oregon have an affinityaffinitya4thwith california huiahulastinusHulahylastinusHylastinus obscuresobscurus marsham scolytus whereas those of northeastern oregon are rugulosusrugulosus mimMifmullermulierfler S multistriatusmultistiiatus marsham related to rocky mountain forest types xyleborus disbardispar fabfabriciusrictus X xylographxylographusus because scolytidae are host specific to some say X californicuscalifornicus wood xylebolinusxyleborinus degree their distribution in oregon is linked saxesenisaxeseni ratzeburg and MonaimonatmonaithrummonarthrumMonthrumarthrum denti closely to the distribution of species of trees and gerum leconte of these X californicuscalifornicus was shrubs known heretofore in oregon from only one oregon scolytids were listed by chamberlin specimen wood 1982 a second specimen was 1917 but that list is greatly out of date we caught by JBJ in flight after sunset 6 VIII herein update the list to include records and 1990 Champochampoegeg state park marion co it synonymiessynonymicssynonymies published by wood 1982 similar probably was introduced by commerce at port- lists have been published recently for idaho land although its native range is still unknown furniss and johnson 1987 and montana gast we speculate that it may infest distressed deciddecad et al 1989 bousuous trees along the willamette river six species not previously reported from by their habits oregon scolytidae are char- oregon were collected by us on field trips in acterized as true bark beetles living in cambium 1990 and four species were found among 105 species ambrosia beetles living in xylem museum specimens more species will surely be although they may feed entirely or partly on

I1 divisiondivision of entomology universityumveiumpei litysity of idaho moscow idaho 83843419683843 4196 2 20regonoregon departmentdepirtmentotagncultuieofagriculture salem oregon 97310011097310 0110 3 olloolio USDAusdafoiestforestFoiest service GrandelagrandeLi oregon 97890

373 374 GREAT BASIN naturalist volume 52 fungi that they transmit 14 species living in phloeosinus hoferihofetihoferd blackman pine cones conophthorus pondeponderosaerosae hop- BIOLOGY monogynous infests branches kins or living in roots of clover H obscuresobscurus of juniperus deppeanadeppeana JJ osteosperma and J conifers are hosts of 98 species while the other scopulorum 23 occur in angiospermsangiosperms species AND NOTES CANADA of listed for distribution abbreviations repositories spec- BC USA all western states except wash to oregon are ODAC oregon imens new OREGON about 9 km W enterprise wallowagallowa of collection salem department agriculture co 9 XI 1990 juniperus scopulorum M M pacific northwest and range PNW forest furniss and A equihua 12 9 5 6 WFBM station collection forest service experiment infesting branches 050.5os 30 cm diameter of a oregon W F USDA corvallis WFBM barr small standing tree larvae parasitized by an museum of idaho entomological university abundant braconid wasp phylusecphylusEc sp proba- moscow idaho and SLW stephen L wood bly nicus rohwer host is restricted in young provo californicuscalifor collection brigham university oregon to the vicinity of the wallowagallowa river for utah a distance of approximately 30 km downstream from enterprise SPECIES NEW TO OREGON subfamily scolytinaescolytidae

subfamily hylesininae trypodendron betulae swaine BIOLOGY monogynous infests betula hylastesHylastes tenuis eichhoff sppapp rare mainusin alnus sppapp tunnels are constructed BIOLOGY monogynous infests pinus sppapp by females radially into sapwood other females presumably the roots construct branches from the radially aligned distribution AND NOTES MEXICO tunnel at close intervals left or lightrightdight in the hildago and mexico state USA mass to fla horizontal plane eggs are laid in niches ori- all southern states westward to calif and ida ented above and below the gallery larvae exca- OREGON eugene lane co 22 IX 1971 black vate short cradles in which they develop and light trap K J goeden 1 I1 ODAC Priprindvilleprinevilleneville feed on ambrosia fungus males are active in crook co 25 VII 1934 1 I1 PNW VIII 1935 1 I1 keeping the tunnels clean and aerated PNW pinus ponderosa hopk 189608318960 83 W J distribution AND NOTES CANADA buckhorn paraparatypestypes of the synonym H alta BC man NB NS NWT ont minutusminutes blackman que USA ida me mass minn mont NH NJ NY SD wise OREGON mill phloeosinus scopulorum scopulorum swaine creek umatilla co 8 XI 1990 betula papyrifera M M furniss and A equihua 1 Y BIOLOGY monogynous infests stems of papyrifera 3 dS WFBM I1 Y I1 d& ODAC infesting lower juniperus scopulorum galleries parallel to stem of a 23 cm diameter wind felled tree also graintraingrain with a nuptial chamber just above the 9 present were xyleborus disbardispar fabricius and entrance appearing as though the chamber xyleborinus saxesenisaxeseni ratzeburg were halved and one side shifted forward half its diameter bright 1976 fig 182 xyleborus xylographxylographusus say distribution AND NOTES CANADA alta BC USA wash OREGON sisters biobloBIOLOGYLO GY unstudiedunstudied in species of this deschutesDeschutes co 8 V 1978 juniperus sp R L genus that are studied haploid males are pro- penrose 4 Y 3 S& ODAC canby klackamasKlackamas duced parthenogenetically they are dwarfed co april 15 1965 K J goeden 1 JS ODA and flightless diploid females are produced by north plains washington co 20 IV 1969 mating between siblings or between a female thuja alicataplicataplicata K J goeden 2 6 1 Y ODA 2 parent and a male offspring infests quercus S& SLW portland multnomahco22multnomahMultnomah co 22 X 1971 sppapp rare in other hardwoods the galleries are chamaecyparis lawsonianalawsoniana R L westcott 1 made obliquely into sapwood in a horizontal Y 161 S ODA I1 Y SLW northbendNorthbend coos co plane to a depth of an inch or more after which 9 VI 1974 on cypress J mclaughlin 3 Y they branch the arms following the annual rings ODA beal and massey 1945 199211992 LIST OF OREGON scolytidae 375

distribution AND NOTES CANADA nissassmss and J B johnson 9 WFBM infesting ont que USA twenty two states and DC stem of a 26 cm diameter tree jackman park east of looth meridian calif 1 I1 specimen tex steens mtnman harney co 14 VIII 1990 pop- OREGON 5 km NW newberg yamhill co 20 ulus tremultremuloidesoides M M furniss and J B john- VI 1970 black light trap K J goeden 1 I1 son 14 WFBM 10 ODAC attacking lower ODAC stem of a 25 cm diameter dead tree foliage shed bark moist trypophtrypophloeusloeusaloeus striastriatulustulus mannerheimManmannerheimonerheim BIOLOGY monogynous infests outer bark pityophthorus scalptorsculptorscalptor blackman of salix sppapp most commonly S scoulerscoulerianaiana BIOLOGY presumably polygynous infests also recorded from alnus sppapp may reinfest stem small branches of living pines progressively downward for several genera- distribution AND NOTES CANADA tions cave type egg gallery larvae mine shal- BC USA calif ida OREGON 15 km N lowly under bark palmer junction union co 16 VIII 1990 distribution AND NOTES CANADA pinus ponderosa M M fumisslumiss and J B john- neafnewf NS que yukon USA alas colo son 2 Y 2 6 WFBM infesting 1 cm diameter ida minn ut OREGON hot springs camp- freshly faded lower branch on a live merchant- ground hart mtnman nati antelope refuge lake able tree each gallery contained only one co 14 VIII 1990 salix scoulerscoulerianaiana M M female and one male no eggs or larvae they furniss and J B johnson 34 WFBM 5 appeared destined to overwinter before repro- ODAC infesting necrotic bark lesions in a live ducing stem having a deep frost crack diameter of infested part 5105 10 cm mature larvae present MonaTmonatthrammonarthrumMonthramthrumarthrum dentigerumdentigerum leconte trypophloeus thatthatcherithatcherscheri wood BIOLOGY not studied infests quercus sppapp most species of monarthrumMonarthrum are polygy- BIOLOGY monogynous infests outer bark nous and their galleries branch from a radially of standing unhealthy or dying populus oriented entrance tunnel inm the xylem larvae tremultremuloidesoides cave type egg gallery larval mines ofthis genus develop inm niches apparently feed- confined bark to outer ing on a mixture of ambrosial fungus that grows distribution AND NOTES CANADA on gallery walls and xylem of the host tree BC USA ida OREGON calif hot springs distribution AND NOTES MEXICO baja mtnman campground hart nati antelope california USA arizanz calif tex OREGON co 14 refuge lake VIII 1990 populus medford jackson co 18 villVIII 1968 black light tremultremuloidesoides M M B furniss and J johnson trap 1 I1 ODAC 27 WFBM 5 ODAC adults attacking and walking on bark of a dying 15 cm diameter tree OREGON scolytidae

procryphalus mucronatusmucro natus leconte hylesininafhylesininae BIOLOGY monogynous infests populus hylastiniHylastini tremultremuloidesoides prefers soft fermenting dead sciurusscierus annectensleconteannectensannectentannectenstengleconte bark usually follows invasion by hylurgops borawsporomsporaws leconte primary hylurgops reticularetimlatusreticulatesreticulatustus wood trypophloeus papulipopuli hopkins petty 1977 the hylugopshylurgops rugipennis rugipennis MannermannerheimmannerbeimManmannerheimonerheimhelmbeimheim gallery is narrower and the bark overlying the hylurgopshylurgyps subcostulatus subcostulatus mannerheimManmannerheimonerheim gallery is thicker than that of T populipapuli hopkins hylastesHylastes gracgraciosgracihsilslis leconte andandpresumablytpresumably T thatcherithatthatcherscheri one and one half HylasteshylastesHylastes tongicollistongicoffis swaine hylastesHylastes macer leconte to two annual generations utah overwinteringoverwintering hylastesHylastes grinusnignnusni mannerheimmannerheimoMannerheim as larvae and adults eggs appear first in late hiasteshylastesHylastes ruber swaine may hylastesHylastes tenuis eichhoff distribution AND NOTES CANADA hylesinini alta BC USA alas colo ida mont nev hylastinusHylastinus obscurusobscumsobscures marsham NM ut OREGON hot springs campground hylesinusHyle sinus califomicus swaine mtnman hyieHylehylesinushylesznussinus oregonuscoregonusoregonus blackman hart catlnatl antelope refuge lake co alniphagusalmphagus asperiaspenaspencollisaspericolliscollis leconte 14 VIII 1990 populus tremultremuloidesoides M M fur alniphagusalmphagus hirhirsutussutus schedl 376 GREAT BASIN naturalist volume 52

tomicinitomicinieTomicini ipiniigini1pini xylechinus montanus blackman Pityopitwgenespityogenesgenes cannulacannulatustus leconte pseudohylesinus disbardispar chspar blackman pityogenesPityopityogenesfissifronsfenesgenes fossifossifronspossifrons leconte pseudohylesinus disbardispar pulpullatuslatus blackman pityogenesPityogenes knechtknechtelieli swaine pseudohylesinus granulagranulatusgranulatestus leconte pityokteinespityoktemes elegansdelegans swaine pseudohylesinus nebulosus nebulosus leconte pityokteinespityoktemes lasiocarpi swaine pseudohylesinus nobilis swaine pityokteinespityokteincs minutesminutus swaine pwudohylesinuspseudohylesinus pini wood pityokteinespityoktemes ornatus swaine elatus eichhoff pseudohylesinus serisericeussericeoussenceusceus Manmannerheimmannerheimonerheim Orthotomicorthotornicusorthotomicusus ca Eichhoffl pseudohylesinus sitchensis swaine ipsaps concinnusconcinnousconcinnus MannerManmannerheirnmannerheimmannerheimonerheimheirn pseudohylesinus tsugietsugae swaine ipsaps ernarginatusemarginatus leconte dendroctonus breulbrevibrevicornishrevicormscornis leconte ipsaps integer Eichhoeichhofffo dendroctonusdendroctonusjeffreyijeffrejeffreyjjeffreyiyi hopkins eps latidenslatidens leconte dendroctonus pondeponderosaerosae hopkins ipsaps mexicanusmexic anus hopkins dendroctonus pseudotsugae hopkins ipg montanus Eichhofeichhofffl dendroctonus rufiniftpennisrufipennisrupipennis kirby ipsaps paraconfusus lanierlamer dendroctonus valens leconte ipsaps pini say ipsaps plastographus marmantimusmaritimusmanmaytimusitimus lanierlamer phloeotribini ips1pseps plastographus plastographus leconte phloeotribusphlocotnbus leconteilecontei schedl ipsaps tedenstndenstridens engelengelmanniengelmanmengelmanniamanni swaine mannerheimonerheimhelmbeimheim phloeosinini ipsaps tedenstndenstridens tndenstedenstridens MannerManmannerheim Phloephloeosmusphloeosinusosmus antennatusantennatus swaine xyloterini phloeosinuphloeosinus cupressipressl hopkins trypodendron betulae swaine phloeosinusfulgensphloeosinu fulgensfulgensi swaine trypodendron lineatumlincatumlinealincalineatumtum olivier phloeosinus hofen blackman trypodendron retusumretusuntretusrutusuniunt leconte phloeosinus punctatuspunct atus leconte trypodendron rufitarsisrufitarsis kirby phloeosinus scopulorumscopulorurn scopulorumscopulorurn swaine phloeosinus hopkins xyleboiinixyleborini sequoiae californicuscaliformcuscalifornicus wood phloeosinusphloeosinuf secratusserratusserratus xijleborusxyleborits leconte dispar fabricius phloeosinus vandyvandykeivandykeskei swaine xyleborus disbar xyleborus intrususintrusus blandford hypoboriniHypoborini xyleborus xylographylographusxylographusus say chaetophloeus heterodoxheterodoxusus casey xyleborinusxylebonnus saxesenisaxesaxesemseni ratzeburg Polygraphpolygraphiniini cryphalini carphoborus intermediusintermedium wood trypophloeus salicissalichs hopkins carphoborus picenepiceae wood trypophloeus triatulusstnatulustriatulus mannerheimManmannerheimonerheim carphoborus pinipimpinicolenspimcolenscolens wood trypophloeus thathatchertchentohen wood carphoborus ponderosaeponderosae swaine procryphalus mucromucronatusnatus leconte carphoborus sansoni swaine procryphalus utahensis hopkins carphoborus vandykeivandyvandykeskei bruck cryphalus pubescentpubescenspubes cens hopkins Polygraphpolygraphusus rufirufipennisruf1pennisrupipennis kirby cryphalus rufiruficolliscollis rufimficollisruficollisrupiruppcollis hopkins scominarScoscolyrinaemiNAr corthylini alnar pseudopityophthorus pubipennispubipenms leconte scolytiniscolyfini conophthorus ponderosaeponderosae hopkins scolytus lancis blackman pityophthorus boychiboycei swaine scolytus monticmonticolaeolae swaine pityophthorus confconfertusertus swaine scolytus inultistriatusmultistnatus marsham pityophthorus confilsconfimsconfinisconfinis leconte scolytus opacusopacous blackman pityophthorus digestusdigestus leconte scolytus aregonioregoni blackman pityophthorus electus blackman scolytus picenepiceae swaine pityophthorusjeffreyipityophthorus jeffrey i blackman scolytus praeceps leconte pityophthorus murraymurrayanaeanae blackman scolytus rugulosusrugulosus Mtmullermulieriller pityophthorus nitinitidulusdulus Manmannerheimmannerheimonerheim scolytus subscabersubscaber leconte pityophthorus nitidusnitidous swaine scolytus tsugietsugae swaine pityophthorus pseudotsugaepseuclotsugae swaine scolytus unispinosus leconte pityophthorus scalptorsculptorscalptor blackman scolytus venventralistralis leconte pityophthorus toraliscoralis wood pityophthorus tuberculatuberculatustus eichhoff micracini gnathotrichus returns leconte hirhirtellushirtelloustellus leconte cnathotnchus hylocurus gnathotnchusgnathotrichus sulcatussulcatus leconte crypturgini monarthrummonarthrurnMonarthrum dentigerurndentigerumbentidentigerum leconte dolurgus humilispumilispurnilispurnilis MannermannerheimmannerheirnManmannerheimonerheimheirn monarthrumMonarthrum scutscutellarescutellateellare leconte cnfpturgusct71pturgus borealishorealis swaine dryocoetini acknowledgments dnjocoetesdryocoetes affaber Manmannerheimmannerheimonerheim dnjocoetesdryocoetes autographautographusalitographusus ratzeburg gary L peters provided collection data for dnjocoetesdryocoetes confususconfusesconfusus swaine H schmidt dryocoetesdnjocoetes sechettisecheltisechelti swaine specimens in the ODAC fred 1992 LIST OF OREGON scolytidae 377

USDA forest service lagrande oregon seg- BRIGHTRIGHT D E JRJK AND R W STARK 1973 the bark and regated undetermined scolytidae in the PNW ambrosia beetles of californiaCahgahfomiaromiabomia coleoptera scolytidae and platypodidaePlatypodidae bulletin of the california insect collection locations ofofbetulabetula papyrpapyriferaifera host survey vol 16 169 appp oftrypodendronbetulaeof trypodendron betulae swaine andjunipemsandjuniperusandjuniperus chamberlinHAMBERLIN WWJJ 1917 an annotated list of the scolytid scopulorum host of phloeosinus hofethodethoferihoferd black- beetles of oregon canadian entomologist 49 321 man were provided by charles johnson USDA 328353 356 service oregon FRANKLINRANKLIN J FFANDCAND C T DYRNESSDYRNFSS 1973 natural vegeta- forest baker the manuscript tion of oregon and washington USDA forest service was reviewed by frank W merickel university general technical report PNW 8 portland oregon of idaho and dr stephen L wood brigham 417 appp young university provo utah who also identi- FURNISSURNISS M M AND J B JOHNSONJOIINSON 1987 list of idaho and notes on new records fied X ornicus X xylographusus and P s scolytidae coleoptera califorcalifornicuscalif xylograph great basin naturalist 47 375 382 those collected us scopulorum other than by casGASASTT S jaj5J M M FURNISS J B JOHNSON AND M A lvieivleIVIFIVIE this is university of idaho agriculture experi- 1989 list of montana scolytidae coleoptera and ment station research paper no 92714 notes on new records great basin naturalist 49 381 386 pettyPFTTYFTTY J L 1977 bionomics of two aspen bark beetles literature CITED trypophloeus populipopuhpapuli and procryphalus mucronatusmucronatus coleoptera scolytidae great basin naturalist 37 BEAL J A AND C L MASSEY 1945 bark beetles and 105 127 ambrosia beetles coleoptera scolytidae with special jooojhooWOOD S L 1982 the bark and ambrosia beetles of north reference to species occurring in north carolina duke and central america coleoptera scolytidae a taxo- university school of forestry bulletinbnlletin 10 178 appp nomicnomie monograph great basin naturalist memoirs brightBRIG hi D EEJRJR 1976 the bark beetles of canada and no 6 1359 appp alaska the insects and arachnids of canada part 2 biosystematics research institute research board canada department of agriculture publication 1576 received 3 april 1992 1 241 accepted I1 october 1992 great basin naturalist 524 appp 378381378 381

RIFFLE BEETLES coleoptera ELMIDAE OF DEATH VALLEY NATIONAL MONUMENT california

william D shepard

abst11actABSI racKAC 1 three species of elmidaeelmidge occur in death valley national monument Stenstenelmiselmis calida is in three springs in the ash meadows area microcylloepusformicoideusmicrocylloepusformicoicleus is only in travertine springs and microcylloepus similis isis in several springs throughout death valley and ash meadows only permanent springs support elmiaselmids considerable morphological variation occurs in the disjunct populations of M similis the evolution of elmiaselmids in death valley national monument is equivalent to that of the local pupfish Cyprincypnnodoncyprinodoncyprinodontodon sppapp

key words death valley insecta coleoptera eimelmidaeelinidaeElmElinelmidgeidae distributions decertificationdesertification evolution

death valley national monument DVNM of the aquatic organisms occurring in is located mostly in southeastern california DVNM only the fishes have been studied with two small extensions into southwestern extensively soltz and naiman 1978 reviewed nevada DVNMDVN M includes death valley proper the past work and presented an excellent syn- its adjacent mountain ranges and the ash thesis particularly so for Cyprincyprinodoncyprinodontodon sppapp pup meadows area of nevada which surrounds fish for aquatic insects studies have been devil s hole biogeographically this is a transi- primarily descriptions of new species and their tion area between the mojave desert and the type localities eg chandler 1949 usinger basin and range desert desert conditions 195611956 only one study colburn 1980 directly here are the result of the drier and warmer addressed the ecology of any species however post pleistocene climate and a rain shadow deacon 196719681967 1968 discussed insects as part of effect from the panamint mountains the sierra the community ecology of saratoga spring nevada and the coast range mountains to the during a vacation I1 found a single specimen west of a riffle beetle in saratoga spring at the south water sources in DVNM are unexpectedly end of DVNM that chance discovery led me to common palmer 1980 cites over 100 springs embark on a survey of the water sources in alone hunt 1975 has classified these springs DVNM to determine if other elmiaselmids riffle bee- into four types based upon volume of discharge tles occurred there and if so in which sources and geomorphic origin the amargosaamargoso river flows when it does into the southern end of METHODS death valley two permanent streams salt creek and furnace creek are located inthein the water sources central portion of DVNM numerous wells the water sources examined were chosen shallow subsurface water sources and seeps primarily because of their accessibility those are to be found scattered throughout DVNM that required more than a daydayss travel by auto these are not reliable water sources being andor foot were not examined in all 27 water more or less intermittent wherever a water sources were examined in death valley and its source does occur however it may not be very environs and in ash meadows death valley amenable to aquatic organisms because of lethal water sources include grapevine spring temperatures andor salinitiessalinities discussions of scotty s castle spring mesquite spring day- the local hydrology can be found in hunt et al light spring hole in the wall spring midway 1966 and soltz and naiman 1978 well stovepipe wells salt creek nevarenevaress

idedeputml t1lenttalenttment of ententomologyri olgy california academyAcadelayelnyerny of sciences golden gate paikpark san francisco california 94118 mailing address 6824 linda sue way fanfairfaig oaksXs california 95628

378 199211992 DEATH VALLEY RIFFLE BEETLES 379

springs texas spring travertine springs emi- TABLE 1 the occurrence of riffle beetles coleoptera elmidge grant spring navel spring tulethletuie spring Elmelmidaeidae inm water sources ofdeath valleymalleymailey national monument Bapadwater s dwaterbadwater shorty well eagle borax spring average warm spring ibex spring and saratoga spring temp elevation ash meadows water sources include indian water source C m species 1 spring school spring devil s hole point of rocks spring Jackjackrabbitrabbit spring big spring death valley and 1 grapevine spring 25 29 820 2 an unnamed spring 2 nevare s springs 300 2 the only permanent water sources are large 3 travertine springs 32323636 122 23 volume springs on the east side of death valley 4 saratoga spring 26 29 46 2 and in ash meadows and devils hole no ash meadows water flows from devildevilss hole but here the 5 indian spring 243024 30 705 12 surface of the ground intersects the hydrologic 6 devildevilss hole 735 1 head of the groundwater so water is always 7 point of rocks spring 705 12 8 big spring 681 2 present in the bottom of a large crevice these permanent sources are all connected with the 1I stalstaist ai calida 2 milrocyllolptiimicnwylloepus sindsimililisils 3 microcylloqmsmilroullicii ash meadows groundwater basin fomicoideusfonnicoicleus collections spring run M similis completely replaces M all of the above water sources were exam- formicoideusformico ideus microcylloepus similis also occurs ined for the presence of riffle beetles where in several other springs grapevine spring possible collecting was accomplished with a nevare s springs saratoga spring indian standard kick net however many of the seeps spring point of rocks spring and big spring and wells had such low discharge andor narrow all water sources inhabited by elmiaselmids were width that collection could be done only by located either on the east side of death valley manual removal of rocks and sticks for visual or in ash meadows with the exception of examination voucher specimens for all species devildevilss hole these springs all exhibit perma- collected were deposited in the author s collec- nent flow of a relatively large volume most of tion at callcailCalicaliforniafomia state university sacramento the water sources not inhabited by elmidselmias are low volume seeps eg daylight spring sub- RESULTS surface sources eg shorty s well or pooled water eg Badwater note added author of the 27 water sources examined 8 were after review richard found to contain populations of elmiaselmids table zack washington state university has found S scruggs 1 Stenstenelmiselmis calida chandler was still resident calida in skruggs spring and mexican spring in meadows WDS in devildevilss hole its type locality however ash during this survey two additional populations were located in nearby indian spring and point discussion of rocks spring la rivers reports unsuccess- fully searching springs near devil s hole in an the major factor linking those springs inhab- attempt to locate additional populations chan- ited by elmidselmias is their association with the ash dler 1949 it is not known whether these addi- meadows groundwater basin this large water- tional populations were missed or ifthey are the shed undoubtedly maintains the constant flow result of colonization or transplantation the required by elmidselmias the only large volume spring run coming from indian spring is very spring not inhabited by elmiaselmids Jackjackrabbitrabbit narrow and deeply incised into the desert floor spring was pumped dry during a local battle making it extremely inconspicuous over water rights microcylloepus fonnicoideusformicoideusformico ideus shepard although it may at first seem incongruous to occurred only at travertine springs shepard find riffle beetles in a desert area one must 1990 near the spring heads a complex of remember that the regional desertificationdecertification is a several upwellings and for many meters below rather recent event geologically and ecologi- M formicoideusformico ideus was the only elcidelmid to be found cally speaking during the several pleistocene further downstream though it co occurs with glacial periods and perhaps even before the M similis hobhomhorn in the lower third of the basin and range desert was far cooler and 380 GREAT BASIN naturalist volume 52 wetter the death valley area is thought then to evolution of pupfish of DVNM see soltz and have had a climate much like the present day naiman 1978 microcylloepusformicoideusmicrocylloepus formicoideusformico ideus is lake mono area 240 kmkin 150 mi to the north similar to Cyprincyprinodoncyprinodontodon diadiabolisdiabolicbolis in being located hildreth 1976 evidence from the distribu- in only one water source and in being very tions of fishes in the desert of california and distinct from and smaller than its convenerscongenerscongeners nevada and along the east front of the sierra Stenstenelmiselmis calida is similar to C salinusgalinus and C nevada suggests that many of the pleistocene billerimilleri in that there are two taxa subspecies lakes overflowed their basins and were con- that inhabit two separate locations along a once nected by extensive river systems miller 1946 free flowing water course la rivers 1949 hubbs and miller 1948 soltz and naiman microcylloepus similis is similar to C nevadannevaden 1978 thus pre pleistocene distributions of sis in being widely distributed but having iso- aquatic organisms would have been subject to lated somewhat morphologically distinct changes during the pleistocene ultimately populations throughout DVNM and surround- these distributions were then subjected to the ing areas influences of the warmer and drier current elmidselmias like most aquatic insects accom- interglacial period present distributions are plish dispersal primarily by flying adults As the therefore the sum of pre pleistocene distribu- post pleistocene desertificationdecertification proceeded tions pleistocene disperdispersalssals and post water sources in the DVNM area became pleistocene vicariantvicariant andingsstrandingsstr smaller fewer and farther apart A point even- small isolated populations that were tually had to be reached at which aerial dispersal stranded in reliable water sources presented became hazardous mutations reducing the ideal situations for rapid evolution given the ability to fly would then be favored indeed small gene pools and lack of gene flow from relatively rapid fixation of these mutations in the other populations these factors have been population would be expected it is not surpris- responsible for the quick proliferation of pup ing then that adults of all three elmidselmias occur- fish taxa in the death valley area soltz and ring in DVNM are either apterous wingless or with incompletely developed naiman 1978 this may also account for the brachypterous and subsequently incapable of flying speciation mfonnicoideusofmformicoideusof the development wings of subspecies in S calida and the inter populational variation in M similis acknowledgments Stenstenelmiselmis calida had been previously reported from ash meadows in the form of its I1 thank the staff of death valley national nominate subspecies A second subspecies S c monument for granting permission to collect humpamoapanumpa la rivers occurs southeast of DVNM for access to their library and for a myriad of along the muddy river in southern nevada helpful comments about this remarkable area each of the various populations of M similis exhibits minor morphologic variations some CITED even in the aedeagus I1 have vacillated for a long literature time concerning the taxonomic status of these disjunct populations however since the genus CHANDLER H P 1949 A new species of Stenstenelmiselmis from nevada pan pacific entomologist 253 133 136 1 needs revision and because I suspect that the COLBURN E A 1980 factors influencing the distribution variation is ecologically induced I1 have chosen and abundance of the caddisflycaddishlycaddis fly limnephilis assimilis to be conservative and not assign separate taxo- in death valley unpublished doctoral dissertation nomic status to any ofthe populations perhaps university of wisconsin deaodeagonDEACONDFAC ON J E 1967 the ecology of saratoga springs some enterprising future student will examine death valley national monument pages 1 26 in stud- how constant warm temperatures influence ies on the ecology of saratoga springs death valley morphologic expression in riffle beetles if so national monument final report of research accom- no 14 10 1989 DVNM and the basin and range plished under NPSNFS contract 1410043419890434 the springs of nevada southern university las vegas university of desert would offer an excellent natural experi- nevada at las vegas ment and the numerous populations of M 1968 ecological studies of aquatic habitats in similis in those springs and spring runs would be death valley national monument with special refer- material ence to saratoga springs final report of research choice study accomplished under NPSNFS contract no 1410043414 10 0434 the elmidselmias of DVNM represent an inverte- 1989 nevada southern university las vegas univer- brate analog of the already well documented sity of nevada at las vegas 199219921 DEATH VALLEY RIFFLE BEETLES 381

HILDRETH W 1976 death valley geology death valley southwestern nevada with a maprnapgnap of the pleistocene natural history association 72 appp waters journal of geology 54 43 53 HUBBS C L AND R R MILLER 1948 the zoological PALMER TST S 1980 place names ofthe death valley region evidence correlation between fish distribution and in california and nevada sagebrush press morongo hydrologic history in the in desert basins of western valley california 80 appp reissue ofthe 1948 printing united states great in the basin with emphasis on SHEPARD W D 1990 microcylloepusformicoideusmicrocylloepus formicoideusformico ideus col- glacial and postglacial times bulletin of the university eoptera elmidaeElmelmidgeidae a new riffle beetle from death of utah 3820 biological series 107 17 166 valley national monument california entomological HUNHUNT r C B 1975 death valley geology ecology archellarcheol news 10131471013 147 153 agyogy press university of california berkeley 234 appp soldzsomSOLTZ D L AND R J NAIMAN 1978 natural history HUNTHUNF sol the C B T W ROBINSON W A BOWLES AND A L of native fishes in the death valley system science WASHBURN general 1966 geology of death valley series no 30 natural history museum oflos angeles california hydrologic basin united states geological county los angeles 76 appp survey professional paper B 494 138 appp USINGERUSINGEH R L 1956 aquatic hemiptera pages 182 228 LA RIVERS 1 I 1949 A new subspecies of stenelmisStenelmis from in R L usinger ed aquatic insects ofcaliforniaafthcalifornia with nevada proceedings of the entomological society of keys to north american genera and california species washington 515 218 224 university ofcalifornia press berkeley 508 appp MILLER R R 1946 correlation between fish distribution and pleistocene hydrology eastern california in and received 16 july 1991 accepted 10 september 1992 great basin NaturnaturalistdistAist 524 appp 382382384384

siphonaptera FLEAS COLLECTED FROM SMALL MAMMALS IN MONTANE SOUTHERN UTAH

2 james R kuceralandand glenn E haashads

key wordswordsfleasfleasgleas siphonaptera utah mammals

recent collections from various small mam- natural history flea specimens are retained by mals of southern utah have helped to elucidate the authors the distribution of fleas siphonaptera within hystnchopsyllahystrichopsylla dippinidippiei truntruncatetruncatacata the special fleas of mam- state of interest were holland 1957 mals found in forested high mountain areas of the southernmost part ofutah an area ofcom- peromyscus inaniculatusmamculatus rufinusmfinusruflnus san juan co abajo mts dalton springs campground plex topography containing habitat varying from 1991 1 ct low desert to subalpine coniferous forests in 2560 m 8 september id microtus ionlonlongicaudusaicaudus alticola idem I1 Y particular we sampled the small mammal flea fauna of the abajo mountains san juan hystrichopsyllahystnchopsylla occidentoccioccidentalsoccidentaleoccidentalisdentalsalis sylvaticussylvaticus county the la sal mountains GrandgrandsangrandmanSan juan campos & stark 1979 pine counties and the valley mountains peromyscus boyki utahensis washington washington county these ranges have been co pine valley mts north juniper park camp- sparsely surveyed in this respect as evidenced ground 2122 m 10 november 1991 3 Y Y by review of the seminal work of stark 1959 peromyscus maniculatusmamculatus sonoriensissononensis washing- after excluding 22 records 13 65 56 37 Y Y of ton co pine valley mts pines campground the ubiquitous deer mouse fleaaethecaflea aethelaaetheca wagnen 2079 m 12 june 1991 1 Y baker which occurs in all counties of utah fleas of the genus hystnchopsyllahystrichopsylla are found beck 1955 we present and discuss the signif- on a variety of small mammals in mesic to moist icance of 42 new records of 12 species of fleas habitats these are the first records ofH dippieidippini A parallel survey offleas found in mammal nests truntruncatatruncatecata from southeastern utah campos and will be presented elsewhere haas and kucera stark 1979 record H 0 sylvsylvaticussylvatwusaticus from san in preparation juan co but our records of this taxon are the first from southwestern mammal nomenclature is that of hall utah 1981 however designations of long tailed corrodopsyllaCorrodopsylla curvatecurvata curvatecurvata vole subspecies should be considered tentative rothschild 1915 because of the present confused state of their sorex palustnfpalustrispalustris navigator baird grand mammals collected with taxonomy were sher- co lasal mts meadow at iowahoowah lake 2769 man live traps at all localities except pines m 15 june 1991 16 1 Y from each of two campground pine valley mountains and snap shrews used at all three localities the pine traps were in the only previous published records from valley mountains and at iowahoowah lake camp- utah of C c curvatecurvata are both from northern ground la sal mountains an asterisk f utah rich co bear lake collection by stan- denotes that the host specimen or at least one ford published by tipton and allred 1951 host specimen was deposited in the mammal 107 and salt lake co wasatch mts egoscue collection of the university of utah museum of 1988 these records are from unidentified

14655465546534555 south locust lamlane 17 siltsaltgilt lake city utah 84117 2557wcalifoimastieetfficalifornia street 7 boulder Ccitygityity nevada 89005

382 199219921 NOTES 383

sorex sppapp records of C c obtusata ex water peromyscopsylla hesperomys adelpha shrews are given from tooelethoele co by egoscue rothschild 1915 1966196619881988 published records of C c curvatecurvata peromyscus maniculatus ruflnusrufinus grand from southwestern states are sparse et al haas co lasal mts iowahoowah0owah lake campground 1973 record it from new mexico additional 26822682min 15 june 199iggi1991 1 0sa 2 Y99Y collecting may reveal its presence in the abajo s2 in johnson and traub 1954 record this mts since water shrews are known to spe- since occur cies from beaver box elder millard san there schafer 1991 juan and washington counties rhadinopsyllarhadmopsylla secsectihssectilissextilistilis secsectihssectilissextilistilis opisodasys kehnl baker 1896 jordan & rothschild 1923 peromyscus maniculatus sonoriensissonotlensis wash- peromyscus mamculatusnwniculatus sononensissonorlensis wash- ington co pine valley mts N juniper park ington co pine valley mts north juniper park campground 2119 in 10 november 1991 1 Y campground 2119 in 10 november 1991 IS16 from each of three hosts peromyscus boyliiboylis this species is known from only one area of rowleyirowleyi idem 2122 in 3 Y Y P m sonorien southern utah garfield co vievic panguitch sis washington co pine valley mts pines stark 1959 but not previously from southwest- campground 2079 inm 11 june 1991 1 9 P m ern utah this specimen possesses 5 spines in sonoriensis idem 12 june 1991 1 Y P m the genal comb as do other utah specimens sonohensissonoriensis idem 161 0 1 peromyscus crinitusctlnituscrinitus sspasp stark 1959 and unpublished data but charac- washington co below dam at baker dam res- ters of the genitalia are clearly referable to R s ervoir foothills of pine valley mts 1463 in 12 secsectihssectifistifis rather than R s goodi hubbard 1941 may 1991 1 9 stark 1959 noted that the genal spine number no known records ofthis species from south- of utah specimens is not consistent with the ern utah are published other than that of hub- original description of R s secsectilissectihssextilistilis bard 1947111 garfield co stark 1959 noted that this species is collected only in moun- catallagiaCatallagia decipiensdecipiens rothschild 1915 tainous areas or moist habitats we have also found this to be true unpublished data per- peromyscus mamculatusmaniculatus rufinus san juan omyscus the usual host co abajo mts dalton springs campground sppapp are 2560 inm 8 september 1991 1 6c P m rufinus malaraeusMalaraeus telchinustelchinestelchinus rothschild 1905 idem 161 S tamias sp idem 2 S& 0S P m rufinus peromyscus maniculatus grand co lasal mts iowahoowah lake camp- sonoilensissonoriensis wash- ington co pine valley mts N park ground 2682 in 15 june 1991 ibe1 0 2 Y Y juniper 129 campground 2119 10 1991 2 these are apparently the first specific in november 6c dS I1 Y P m idem 1 1 Y P records published for that part of utah south of sonoriensis 0d I m sononensissonoriensis idem 2 Y 9 P m the colorado river beck 1955 table 3 lists it sonoriensissonofiensis idem 1 9 peromyscus 1 as in san juan county boyliiboylis rowleyirowleyi idem I occurring in 9 peromyscopsylla selenis rothschild 1906 Malaramalaraeuseus sinomussinemus jordan 1925 peromyscus maniculatusmamculatus rufinus san juan peromyscus crinituscrinitus sspasp washington co co abajo mts dalton springs campground below baker dam reservoir foothills of pine 2560 in 8 september 1991 1 06 microtus valley mts 1463 in 11 may 1991 1 6 4 9 9 longicaudus alticola idem 1 6 M I1 alticola P crinituscrinitus sspasp idem 12 may 1991 1 6 2 9 9 idem I1 Y M I1 latuslotusiotuslafus washington co pine P crinituscrinitus sspasp idem I1 dS 4 Y Y peromyscus valley mts pines campground 2079 in 12 june trueithuei truel idem 3 Y Y P t truel idem 161 S 2 199121991.2iggi1991 2 6 6 1 Y 9 9 P boyliiboylis rowleyirowleyi idem I1 dS P boyliiboylis this species was not previously known from rowleyirowleyi idem 2992 Y Y utah south of the colorado river johnson and beck 1955 table 3 lists malaraeusMalaraeus traub 1954 give a record from iron county telchinustelchinestelchinus as occurring in washington county bordering washington county to the north hubbard 1947 200 gives the only specific this species is most commonly collected from record from southern utah garfield co microtus sppapp but also from other small mam- however several records of M sinemussinomus from mals sympatricsympatnc with the voles desert areas of southern utah are given by stark 384 GREAT BASIN naturalist volume 52

1959 also many specimens of M sinomussinemus acknowledgments from peromyscus stinituscrinitusctinituscrinitus were taken by the senior author in snow canyon washington co we thank H egoscue and C pritchett for incidental to the search for traubellaTraubella grundtwnnigrundgrundmannimanni their review of the manuscript and E rickart museum of egoscue 1989 M telchinestelchtelchinusinus seems to be like 0 university of utah natural history some peromyscus keenideeni in being found only in nonnondesertdesert habitat for assistance in identifying specimens megabothris abantisabantes rothschild 1905 literature CITED microtus longicaudus alticola grand co BECKRFCK DDEE 1955 distributional studies of parasitic arthro- 2676 m reck lasal mts meadow at iowahoowah lake pods in utah determined as actual and potential vec- 14 june 1991 161 0 1 Y peromyscus maniculatus tors of rocky mountain spotted fever and plague with young rufinus idem I1 Y M 1 alticola idem I1 Y M notes on vector host relationships brigham university science bulletin biological series 1 1 1 ticola 15 1991 1610 1 Y M I1 al idem june lo CAMPOS E A AND H E STARK 1979 A revaluation siesic alticola san Jjuanuan co abajo mts dalton springs of the hystrichopsyllahystnchopsylla occidentoccidentahsoccidentalisoccidentalistalisails group with descrip- campground 2560 m 8 september 1991 1 6 tion of a new subspecies siphonaptera hystrichopsyllidaehystnchopsyllidae journal of medical entomology 1955 table 3 listed this as beck species 1543115 43144444 occurring in beaver iron sevier and wayne ecoEGOEGOSCUESCU E H E 1966 new and additional host flea associ- counties this was apparently overlooked by ations and distributional records of fleas from utah great 26 71 75 196 who stated flea appears basin naturalist stark 19591961959 this 1988 noteworthy flea records from utah nevada confined to the northern half of the state and oregon great basin naturalist 48 530532530 532 egoscue 1988 reported collecting one male 1989 A new species of the genus traubellaTraubella bulletin of the from a pika at johnson reservoir siphonaptera ceratophyllidae specimen southern california academy of sciences 88 131 134 sevier county in south central utah the dis- HAAS G E R PE MARHNMARTIN M SWICKARD AND B E tributiontribution map of haddow et al 1983 map 76 MILLERMILLFR 1973 siphonaptera mammal relationships in new of medical indicates a locality record in that same region of northcentralnorthcentral mexico journal ento- mology 10 281 289 the first for southeastern utah our records are HADDOW J R TRAUB AND M rothschild 1983 dis- utah megabothrismegabothtis abantisabantes is usually found on tributiontribution ceratophyllidofceratophyllidof fleas and notes on their hosts various species of microtus pages 42 163 in R traub M rothschild and J F haddow the rothschild collection of fleas the ceratophyllidae keys to the genera and host relation- eumolpianus eumolpieunw1pi americanosamericanusameric anus ships with notes on their evolution zoogeography and hubbard 1950 medical importance 288 appp privately published I1 HALL E R 1981 the mammals of north america and2nd ed tamias sp san juan co abajo mts dalton 2 volumes john wiley & sons inc new york 1181 appp springs campground 2560 m 8 september HUBBARD C A 1947 fleas of western north americaamenea iowa state college press 533 appp 1991 2 Y Y 1950 A pictorial review of the north american these specimens seem closer to E e amer chipmunk fleas entomological news 60 253 261 icanus than to E e eumolpi recorded by beck JOIINSONJOHNSON P T 1961 A revision of the species of mono icarus kolenaty monopsyllus several psylluspsyllos kolenatiKolenati in north america siphonaptera 1955 then in the genus ceratophyllidae technical bulletin no 1227 agri- of the type specimens were collected in san cultural research service USU S department of agricul- juan county hubbard 1950 johnson 1961 ture 69 appp TRAUB of flea indicates that intergradation between E e ametiameliameriameniamenn jonnsonJOIINSONJOHNSON PPTANDRT AND R 1954 revision the ee genus peromyscopsylla smithsonian miscellaneous banuscanus and E e eumolpieunw1pi occurs in the county collections vol 123 no 4 68 appp SCHAFESCIIAFFRii TTSS 1991 mammals of the abajo mountains an isolated mountain range in san juan county southeast- in summary the significant findings among ern utah occasional papers no 137 the museum 64 collection records of 13 species of fleas are as texas tech university lubbock slarkSTARK H E 1959 the siphonaptera of utah US depart- follows the first records south of the colorado ment of health education and welfare communica- river in southeastern utah for hysttichopsyllahystrichopsylla ble disease center atlanta georgia 239 appp dippieidippini truntruncatatruncatecata corrodopsyllaCorrodopsylla c curvatecurvatacurvata per tipton VVJJ AND D M ALLRFDALLRED 1951 new distribution with the description of a 0myscopsyllaomyscopsylla selenis and megabothtlsmegabothris abantisabantes records of utah siphonaptera new species of megarthroglossus jordan and roth- and the first in washington county southwest- schild 1915 great basin naturalist 11 105 114 ern utah for H occidentoccidentalisoccidentalistalis sylvaticussylvaticus rhad received 22 may 1992 inoinopsyuainopsyllapsylla s secsectilissextilistilis P selenis and opisodasys kehnl accepted 2 november 1992 great basin naturalist 524 appp 385386385 386 NOTES ON SPIDER theridiidae salticidae PREDATION OF THE HARVESTER ANT pogonomyrmex SALINUS OLSEN hymenoptera formicidae myrmicinaemyrmicidae AND A POSSIBLE parasitoid FLY chloropidae

2 william H clarkclarkaclark1 and paul E blomBIOM

key words pogonoinyructexpogonomyrmex salinus harvester ants euryopisEuryopis formosa xysticusxysticusspiderXysticusleus spider predatorspredatory inelneIncInertellacertelLi parasite

spiders are known predators of ants pres- that was carrying a worker of pogonomynnexpogonomyrmex sure exerted by consistent spider predation can salinusgalinus olsen hymenoptera formicidae alter the behavior of ant colonies mackay myrmicinaemyrmicidae across a large area of basalt rock 1982 and may be a selective pressure contrib- the ants were actively foraging in the area the uting to the seed harvesting behavior of air temperature shaded was 31 C and the soil pogonomyrwxpogonomyrmex mackay and mackay 1984 surface in the sun was 39539.5 C no other spiders we observed the spider euryopisEuryopis formosa of this species were encountered prey capture banks araneae theridiidae capture and was not observed transport workers of the harvester ant on 31 august 1991 at 1725 h at the clark pogonomymwxpogonomyrmex salinusgalinus olsen hymenoptera county site we observed a crab spider of the formicidae myrmicinaemyrmicidae in southeastern genus xysticusXysticus preying on P salinusgalinus about 20 cm idaho additional observations revealed a crab from the ant nest entrance the ants were still spider of the genus xysticusXysticus preying on P actively foraging at this time one spider was salinusgalinus and the presence of a chloropiachloropidchloropid fly riding on the ant in the shelter of an isolated Incincertellajncertellaertellaentella that may have been parasitizing the clump of indian ricegrassricegrass oryzopsis hyme moribund prey subdued by the spider noides at the edge of the ant mound the ant was initially very active walking around an old STUDY SITE grass stem while the spider made periodic attacks on the ant As time progressed involun- one collection site is located along road tary spasms in the ant increased the spider was T 20 county s6sa on the butte t4ntan r31e generally oriented toward the posterior of the idaho national environmental research park ant biting it at the base of the petiole some- INERP in the cold desert of southeastern times the spider was perpendicular to the ant second set ofobservations was made idaho the holding on to the ant with only its mandibles on the INERP clark county s34 t7ntan r31e after five minutes the ant fell onto its side and along highway 28 voucher specimens of all movements slowed at 1740 h only its antennae species have been deposited at the orma J were slightly and a minute later the smith museum of natural history albertson moving spider moved the ant under a small stick two college of idaho caldwell idaho 83605 USA small flies approached the ant and one flew onto CIDA its head occasional movements jerks of the ant s legs were observed at 1751 h this time RESULTS AND discussion at we collected the spider the ant and one of the on 3 july 1988198810201020 h at the butte county flies WHC 9170 the fly is a female Incincertellaertella collection site we collected a single individual of diptera chloropidae and may represent an euryopisformosaeuryopisEureunyopisformosa banks araneaeAranaraneaecae theridiidae undescribed species brown and feener 1991

1 ormiorma J smith museum of natural history albertson college of idaho caldwell idaho 83605 usaandUSA and departmentdepar tmentament ofplant soil and entomological Scisnescisneesscienceses univeisityuniversityuniver srtyarty of idaho moscow idaho 83843 USA department ofot plant soil and entomological sciences university of idaho moscow idaho 83843 USA

385 386 GREAT BASIN naturalist volume 52 have found the phorid apocephalus paraponerae small gray spiders capture ants on their mounds selectively parasitizing moribund workers ofofparaokparaearneanapara and drag them away by a web sling attached to ponera alavataclavataclavata it may be that these Incincertellaertella the ant and to the tip of the spider s abdomen flies are seeking a similar host and opportunis- euryopsisEury opsis formosa is found from central cali- tically exploiting the spider prey the flies were fornia north to british columbia and east to not observed to interact with living active ants wyoming levi 1954 E fornosaformosafornwsa may also be at 1740 h we noticed a second spider E an important predator of P salinusgalinus at this site formnormpormformosatsa on the same ant mound this spider and of pogonomynnexpogonomyrmex species in the western oriented uphill on the side of the mound facing united states the relatively greater precision the ant nest entrance at 1742 h an ant walked and speed with which euryopsisEury opsis subdued and over and slightly past the spider apparently transported the P salinusgalinus prey suggests an failing to recognize the predator s presence the established predator prey relationship spider remained motionless as the ant passed then spun around and mounted the ant s gaster acknowledgments moved face the spider released the ant and to this work was conducted under the INEL it the ant began convulsing at this time while radioecologyRadioecology and ecology programs sponsored the spider sat I1 cm away from the ant facing by the office of health and environmental away from the ant by 1745 h no motion was research and the division of waste products 1746 h observed in the ant and at the spider through the fuel Reprocessing and waste man- side climbed onto the ant the ant was on its agement division united states department of with the spider on top facing the gaster A fly energy 0 D markham T D reynolds and similar to those mentioned above moved onto love J B johnson have provided assistance J the head of the ant at 1747 h the spider was mccaffrey and H W levi provided spider dragging the ant across the mound using a web determinations C W sabrosky identified the described porter and sling as previously by Incincertellaertella specimen and B V brown assisted eastmond 1982 formor the spider E cokl south- forthe in this paper is published as idaho agriculture eastern idaho the spider dragged the ant to the experiment station paper no 91767 edge of the mound and into the grass clump mentioned earlier several other worker ants literature CITED were observed strung up in the grass clumps at this point we collected the spider WHC 9171 ALLRED D M 1969 spiders of the national reactor test- ing station great basin naturalist 29 105 108 the spider genus euryopisEuryopis is known to prey BLOM P E W H CLARK AND J B JOHNSON 1991 on ants levi 1954 carico 1978 including har- colony densities of the seed harvesting ant vester ants of the genus pogonomyrmexpogonomyrnwx in pogonomyrmex salonus hymenoptera forimcidaeformicidae in national north 1982 porter and east- seven plant communities on the idaho engi- america mackay neering laboratory journal of the idaho academy of mond 1982 mackay 1982 has reported science 27 28 36 E califomicacalifomica preying on P rugosus in southern BROWN B VANDDV AND D H FEENER JR 1991 behavior and california host location cues ofapocerphalusofapocethalus paraponerae dip- tera Phonphondaepboridaephoridaedae a parasparasitoiparasitoiditoi of the giant tropical ant prey of E formosa has not previously been Paraparaponeraponera alavataclavata hymenoptera formicidae reported levi 1954 nor has the spider been biotropicabiotropica2323 182 187 reported from the INERP levi 1954 allred CARICO J E 1978 predatory behavior in euryopisfunerisEureuryopiseunyopis funensdunens aranea and the evolutionary 1969 1954 distribution hentz theridiidae levi gives the of the significance ofweb reduction symposium of the zoo- species over most of idaho except for the south- logical society of london 42 51 58 western comer so its presence here was ex- LFVI H W 1954 spiders of the genus eunjopsiseuryopsisEuryeurgopsis from pected allred 1969 reported a related species north and central america american museum novitatesnovitiatesNovitates 1666 114848 Euryopis scriptipesscriptipes banks from the southeastern euryopis scilptipes MACKAY W P 1982 the effect of predation of western border of INERP during july pogonomynnexpogonomyrmex widow spiders araneae theridiidae on harvester ants salinusgalinus is the dominant seed harvesting ant on the hymenoptera formicidae Occooccologiaoecologia53logia 53 4064406411406 441111 INERP occurring in almost all ofits plant commu- MACKAY W PPANDAND E E MACKAYMAC KAY 1984 why do harvester ants store seeds in their nests sociobiologySociobiology 9 31 47 nities blom etalet al 1991 in poterPORIFR S DANDDD AND D A eastEAhlEASTMONDMOND 1982 euryopisEurEuryeuryoptseurgyopisopts coklcoet porter and eastmond 1982 found euryoneuryop theridiidae a spider that preys on pogonomynnexpogonomyrmex sis cokl levi to be a common predator of ants journal of arachnology 10 275 277 pogonomyrmex owyheei P salinus in south- received 6 february 1992 eastern idaho during july and august these accepted 10 september 1992 dad6 466

T H E GREAT BASIN naturalist F r j W ff

it IK

I1 N D E X

VOLUME 52 1992

BRIGHAM YOUNG university great basin naturalist 524 1992 appp 388 391 INDEX volume 52 1992

authorindexAUTHOR INDEX

R 382 steven A 284 kucera james acker E 122 agenbroad larry D 59 kucera thomas allredaliredailredallredkellyw41kellykeily W 41 kudo j 29 anderson stanley H 139 253 launchbaugh karen L 321 arnow lois A 95 leidyroberta68leidy robert A 68 arnowted95arnouarnow ted 95 lindzey frederick G 232 D 352 364 austin dennis marks jeffrey shaw 166 beck reldon F 300 mcarthur E durant I1 bernatas susan 335 mcnulty irving R 95 blackburn wilbert H 237 mead jim I1 59 block william M 328 meyer susan E 53 blom paul E 385 miller gary C 357 boe edward 290 morrison michael L 328 bonham charles D 174 moseley robert K 335 burge howard L 216 murray leigh W 300 callahan J R 262 negus norman C 95 clark william H 385 nielsonmw160nielson M W 160igo comstock jonathan P 195 norling bradley S 253 A 288 cookecookelynna288lynn palmquist debra E 313 cottrell thomas R 174 pendleton rosemary L 293 75 cronquist arthur pickering russell 309 E 11 cushinggushing C pieper rex D 300 russell 262 davis quinney dana L 269 deleray mark A 344 rasmussen G allenalienailen 185 R 95 195 ehleringer james roberson jay A 25 elias scott A 59 rocheroehe ben F jr 185 evans R P 29 rocheroehe cindy talbott 185 flinders berranjerran T 25 rumble mark A 139 furniss malcolm M 373 rushforth samuel R 131 gaineswlllgaines W L I1 I1 saab victoria ann 166 gill ayesha E 155 scoppettone G gary 216 glenn E 382 seville robert S 309 haas G 226 S 328 sheeley douglas hall linnea william D 378 B A 160 shepard haws wes 364 hovingh peter 278 shields hubert wayne A 253 shiozawa D K 29 simanton J roger 237 jehl joseph R jr 328 slaugh bartel T 25 johansen jeffrey R 131 smith bruce N 93 johnson james B 373 smith loren M 226 johnston N paul 25 smith S D 11 kay charles E 290 snyder warren D 357 kaya calvin M 344 stantonstantonnancyl309nancynaney L 309 kitchen stanley G 53 swiecki steven R 288 knapp paul A 149

388 199211992 INDEX 389

taylor daniel M 179 wagstaff fred 293 thomas J diane M 309 bansiwansi tchouassiTchouassi 300 torgersen torolf R 373 welch bruce L 293 trost charles H 179 weltzweitz mark A 237 petelpeter weltzmarka237 tuttle L 216 westcott richard L 373 tyser robin W 189 wester david B 226 uresk daniel W 35 williams R N 29 urness philip j321J 321 352 364 wood stephen L 78 89 woodward S R 29 van sickle walter D 232 vickery robert Kajrkjrjr 145 yearsley kurtis H 131 voorhees marguerite E 35 yensen eric 155 269 young james A 245 313

KEYWORDKEY WORD INDEX

age growth 216 bunchgrassbunchgrass 284 alien flora 189 burrow stucstucturestructureture 288 alpine vascular flora 335 analysis california 41 122 factor 174 death valley 378 fecal 300 mono lake 328 dietary 269 caribou 321 microhistologicalmicrohistological 300 caves59caveseaves 59 anas acetaacutaacutal 226 centaurea virgatavergata sspasp squarrosasquarrosesquarrosa 185 annual grass 245 cervid 321 aplanusiella 160 checking stations 364 apple trees 352 chromatography 174 aquatic habitat 278 chukar 25 arid lands 149 rearing25rearing 25 aristida 41 cicadellidae 160 artemisia cold desert 11 195 nova 313 coleoptera 11 78 89 378 tridentata 174 colonization 328 tritrldentata dentatahitritridentatahidentatatridentate 284 colonizing species 245 dentatatrih1dentatathitrn wyomingensis 284 colorado 174 357 arthropods 59 colorado plateau 59 195 avifauna 278 columbian sharp tailed grouse 166 bark beetles 78 competitive release 68 cottonwood beardtongue 53 357 cutthroat trout 29 beaver dam creek 131 utah cynomys leuc behavior 25 leucurusurus 288 foraging 293 death valley california 378 benthos 11 deer 321 big sagebrush 284 damage evaluation 352 biochemical differentiation 155 management 364 biogeography 262 mule 122290352364122 290 352 364 biomass 313 white tailed 290 leaf 237 depredation 352 birds 278 desert 278 migrating 179 soil formation 313 black sagebrush 313 streams 131 body desertificationdecertification 378 condition 226 diatoms 131 size 216 dietary analysis 269 browse293browse 293 diptera 11 bumblebees 145 dispersal 262 distributiondistributionss 160 378 volume 52 390 GREAT BASIN naturalist

53 disturbance 253 induced dormancy 378 DNA 29 insecta sequencing west 95 drought response 237 intermountain interspecific hybridization 290 einwria 309 inventory 357 electivities 68 irrigation reservoirs 179 electrophoresis 155 islands 328 elk 321 idaho 335 Elmelmidaeelmidgeidae 378 kane lake cirque ephemeroptera 11 lake 344 etheostomietheostomaetheostoiwiEtheostomi nigrimnigrummgrum 68 land bridge 328 euryopsisEuryeuryopsisfirtnosaopsis fornosafonnosa 385 leaf area index 237 evolution 378 leafleafbiomassbiomass 237 160 factor analysis 174 leafhoppersleafhoppers 300 faunal list 373 lepus cahfomicuscalifomicus 216 fecal analysis 300 life history degradation 284 fecundity 216 long term site felix concolor 232 mammals 382 fish 344 management 166 fleas 382 deer 364 flora forest 139 alien 189 medusmedusaheadahead 245 alpine vascular 335 meleagrismclemcieMeleagns gallogalhpavogallopavopadopavo 139 rare 335 merriam s wild turkeys 139 floristics 41 micromicrohabitathabitat use 68 flower colors 145 micromicrohistologicalhistological analysis 300 food habits 216 269 microtus 262 foraging behavior 293 montanus 328 forest management 139 migrating birds 179 fruit trees 352 migration 122 344 functional groups 11 mimulus 145 moapa conoonconaceacopaceaacea 216 ghost towns 149 moaba coticotiacea park montana 189 moabamoapa dace 216 glacier 328 grand canyon arizona 59 mono lake california grassland 95 montana glacier park 189 aylinggrayling 344 gi specializations 68 35 245 morphological grazinggraying nevada 216 great basin 195 278 muddy river 149 mudmudflatsflats 179 desert 122 290 352 364 ash 35 mule deer green 321 squirrels 155 269 mushroom ground 321 orusgrmgrus canadensis 253 mycophagy nebraska habitat platte river 253 aquatic 278 288 descriptions 139 nest 216 selection 139 253 nevada muddy river 216 slimmersummer h characteristics 166 new genus 160 use 179179216216 160 ants 385 new species harvester breadth 68 hibernaculum 288 niche 75 78 89 160 nomenclature hosts pintails 226 hummingbirds 145 northern nutrients 293 hunterhuntel opinions 364 nutrition 321 idalio166269335IdahoIdalio 166269335166 269 335 idaho166 maple 95 kane lake cirque 335 oak pioneer mountains 335 odocoileus 122 290 293 idaho ground squirrel 155 hemionus vftginianusmr 290 imprinting 25 simanus odonata 11 Incincertellaertella 385 199211992 INDEX 391

oncorhynchus 29 siphonaptera 382 orchards 352 soil 174 oregon284Oregonoregon284373284373284 373 compaction 149 palmer penstemonpenpentstemonstemon 53 nitrate 313 parasite 385 organic carbon 284 partridge 25 organic matter 284 peach trees 352 recovery 149 Penpenstemonpenstenumpentstemonstemonstenum palmeri 53 speciation 145 percidae 68 spermophilus 155 peromyscus maniculatusnwniculatus 328 brunbrunneusbrunneousneus 155 phenology 195 townsendtownsentownsendiitownsendntownsendiadnii 269 pioneer mountains idaho 335 mdecemlineatustndecemlineatus 309 plant adaptation 95 195 spider predators 385 platte river nebraska 253 spring streams 11 platypodidaePlatypod idae 78 89 squarrose knapweed 185 plecoptera I111 I1 state records 335 pleistocene 262 stream 344 pogonomynnexpogonomy rinexriney sahnussahnassalinus 385 summer habitat characteristics 166 pollinator preferences 145 survival 25 polymerase chain reaction 29 symphoncarpossymphohcarpos occidentoccidentalisoccidentalistalis 35 polysporocysticpolysporocystic coccidia 309 taeniatherumTaeniatherum medusaemedusan populus sppapp 357 caput 245 predictive models taxonomy 75 78 155 226 texas prevalence 309 226 thymallus productivity I111I1 arcticusarcticus 344 track survey 232 propagation 25 trichoptera 11 prunus virginiajavirginianamrgimanavirginiana 35 trophic levels 11 quaternary 59 tympanuchus phasianellus columbianuscolumbianus 166 quercus gamberigambehigambehi 293 questionnaires 364 utah131utah 131 232382232 382 beaver dam creek 131 radio telemetry 122 red butte canyon 95 rangerangelandand weeds 185 utilization 293 rare flora 335 red butte canyon utah 95 vicarivicanancevicariancevieavicaanceaneenancenanee 262 reproduction biology 216 vole 262 research natural area 95 riparian 357 water ecology 95 depth253depth 253 river boostsroosts 253 drawdown 179 rosidaeRos idae 75 stress 195 ruminant 321 waterfall 344 waterfowl 226 sagebrush 174 weather 122 big 284 weed dispersal 185 black 313 wetlands 278 salinity 195 white tailed deer 290 salmonsaimonsalmondssalmomdssalmonidsids 344 wild sandbars wildfiresfires 245 284 253 wildlife sandhill crane 253 habitat 357 scolytidae 78 89 313 seed methods 364 techniques bank53bank 53 364 winter 293 germination 53 browsing 352 29 sequencing wisconsin glacial 59 sex differences 122 wooded draws 35 sheep185sheep 185 wool 185 shorebirdsshorebirds 179 wyoming 290 shrubs 35 succession 31 xysticusXysticus 385 392 GREAT BASIN naturalist volume 52

TABLE OF CONTENTS volume 52

no I1 march 1992 articles in memoriammemonam A perry plummer 191119111991iggi1991 teacher naturalist range scientist 14 I1 I1 E durant mcarthur secondary production estimates ofbenthicofbenthic insects in three cold desert streams gainesgames gushing and S D smith 1114 I1 I1 W L caines C E cushing effect of rearing method on chukar survival bartelbarteltbarteitT Slaughslaugh berranjerran T flinders jay A roberson and N paul johnston 25 DNA extraction from preserved trout tissues D K shiozawa J kudo R P evans S R woodward and R N williams 29 relating soil chemistry and plant relationships in wooded draws of the northern great plains marguerite E voorhees and daniel W uresk 35 the genus AristilaristidaartstidaArtstidatodatodo cramGramgramineaemeae inm california kelly W allred 4144 temperature mediated changes in seed dormancy and light requirement for Penpenstemonpentstemonstemon palmenpolmen scrophulariaceae stanley G kitchen and susan E meyer 53 late quaternary arthropods from the colorado plateau arizona and utah 1 agenbroad I1 I1 I1 scott A elias jim I mead and larry D 59 Micromicrohabitathabitat selection by the johnny darter etheostoma nigrimnigrummgrummagrum rafinesqueRafinesque in a wyoming stream robert A leidy 68 nomenclatural innovations in intermountain rosidae arthur cronquist 75 nomenclatural changes and new species in Platypodplatypodidaeidae and scolytidae coleoptera part II11 wood 78 1 stephen L I1 I1 I nomenclatural changes in scolytidae and Platypodplatypodidaeidae coleoptera stephen L wood 89 book review plant biology ofthe basin and range C B osmond L F pitelkaPtteikatelka and G M hidy N smith 93 I1 I1 bricenbrucenbruce

no 2 june 1992 articles red butte canyon research natural area history flora geology climate and ecology james R Ehlerehleringerehlermgermger lois A arnow ted arnow irving R mcnulty and norman C negus 95 influences of sex and weather on migration ofmule deer in california thomas E kucera 122422

diatom flora of beaver dam creek washington county utah USA I1 kurtis H yearsley samuel R rushforth and jeffrey R johansen 131434 stratification of habitats for identifying habitat selection by merriam s turkeys mark A rumble and stanley H anderson 139439 pollinator preferences for yellow orange and red flowers of mimulus verbenaverbenaceusverbenaceousceus and M cordicardicordtcardinalisnalts robert K vickery jr 145445446 soil loosening process following the abandonment of two and western nevada townsitestownsites paul A 149449 1 knapp I biochemical differentiation in the idaho ground squirrel spermophilus brunbrunneusbrunneousneus rodentia in yensen ScurScunscundaescuridaesciuridaeseundaeidae ayeshaayeshaeayeshahE gill and eric 455155 new genus aplanusiellaapkinustella and two new species of leafhoppersleafhoppers from southwestern united states homoptera cicadellidae deltocephalmaedeltocephalinae M W nielsonandbnielson and B A haws 160460 199211992 INDEX 393 summer habitat use by columbian sharp tailed grouse in western idaho

1 I1 ann 166 I victoria saab and jeffrey shaw marks notes characteristics of sites occupied by subspecies of artemisia tridentata in the Picepiceanceance basin colorado thomas R cottrell and charles D bonham 174474 use of lakes and reservoirs by migrating shorebirdsshore birds in idaho daniel M taylor and charles H trost 179 dispersal of squarrose knapweed centaurea cirgatavirgatavergata sspasp squarrosasquarrosesquarrosa capitula by sheep on rangeland in juab county utah cindytalbottcindy talbott roche ben F roche jr and G alienallenailen rasmussen 185 vegetation associated with two alien plant species in a fescue grassland in glacier national park montana robin W tyser 189

no 3 september 1992 articles plant adaptation in the great basin and colorado plateau R 195 I1 jonathan P comstock and james ehleringer life history abundance and distribution ofofmoapamoabamoapa dace moabamoapa coricoriaceaacea peter I1 G gary scoppettone howard L burge and L tuttle 216246 condition models for wintering northern pintails in the southern high plains B 226 I1 loren M smith douglas G sheeley and david wester evaluation ofroad track surveys for cougars felis concolor

I1 I1 I1 walter D van sickle and frederick G lindzey 232 leaf area ratios for selected rangeland plant species mark A weltz wilbert H blackburn and J roger simanton 237 ecology and management ofmedusaheadofmedusaheadmedusahead taeniatherumTaeniatherum caput medusaemedusan sspasp asperumesperum simk melderis james A young 245 roost sites used by sandhill crane staging along the platte river nebraska bradley S norling stanley H anderson and wayne A hubert 253 post pleistocene dispersal in the mexican vole microtus mexicanusrnexicanusmexicanus an example of an apparent trend in the distribution ofsouthwestern mammals russell davis and J R callahan 262 can townsend s ground squirrels survive on a diet of exotic annuals eric yensen and dana L quinney 269 notes avifauna ofcentral tule valley western bonneville basin peter hovingh 278 wildfire and soil organic carbon in sagebrush bunchgrassbunch grass vegetation steven A acker 284 structure of a white tailed prairie dog burrow

I1 lynn A cooke and steven R swiecki 288 hybrids of white tailed and mule deer in western wyoming

I1 charles E kay and edward boe 290

no 4 december 1992 articles winter nutrient content and deer use of gambel oak twigs in north central utah

I1 rosemary L pendleton fred J wagstaff and bruce L welch 293 botanical content ofblack tailed jackjackrabbitrabbit diets on semidesert rangeland murray I1 tchouassiTchou assi bansiwansi rex D pieper reldon F beckbeek and leigh W 300 species ofeimeriaofeimeria from the thirteen lined ground squirrel spermophilus tridecemlineatus from wyoming robert S seville diane M thomas russell pickering and nancy L stanton 309 394 GREAT BASIN naturalist volume 52 plant agesizeage size distributions in black sagebrush artemisia nova effects on community structure james A young and debra E palmquist 313 mushroom consumption mycophagy by north american cervids

I1 I1 I1 karen L launchbaugh and philip J urness 321324 terrestrial vertebrates of the mono lake islands california michael L morrison william M block joseph R jehl jr and linnea S hall 328 vascular flora ofkaneolkaneof kane lake cirque pioneer mountains idaho robert K moseley and susan bernatas 335 lakeward and downstream movements ofageofagoof ageageo 0 arctic grayling thymallus arcticusarcticus originating between a lake and a waterfall mark A deleray and calvin M kaya 344 effects of browsing by mule deer on tree growth and fruit production in juvenile orchards

I1 dennis D austin and philip J urness 352 changes in riparian vegetation along the colorado river and rio grande colorado I1 warren D snyder and gary C miller 357 resident utah deer hunters preferences for management options dennis D austin philip J urness and wes shields 364 list of oregon scolytidae coleoptera and notes on new records malcolm M furniss james B johnson richard L westcott and torolftorolfrR torgersen 373 riffle beetles coleoptera elmidaeElmelmidgeidae of death valley national monument california

I1 william D shepard 378 notes siphonaptera fleas collected from small mammals in montane southern utah james R kucera and glenn E haas 382 notes on spider theridiidae salticidae predation of the harvester ant pogonomyrmex salinusgalinus olsen hymenoptera formicidae myrmicinaemyrmicidae and a possible parasitoid fly chloropidae

1 1 I1 william H clark and paul E blom 385 index to volume 52 387 information FOR AUTHORS great the basin naturalist welcomes previously VOUCHER SPECIMENS unpublished authors are encouraged manuscripts pertaining to the biologi- to designate properly prepare label and deposit calnaturalcal natural history of western north america pref- high quality voucher specimens and cultures docu- erence will be given to concise manuscripts ofup to menting their research in an established 1200012 000ooo words perma- nent collection and to cite the repository in publi- SUBMIT manuscripts to james R barnes editor cation great basin naturalist 290 MLBM brigham references IN THE TWTEXT are cited by author young university provo and utah 84602 A cover date ege g martin 1989 or martin 1989 multiple letter accompanying the man manuscriptascriptuscript must include citations should be separated by commas and listed phone numbers of the author submitting the in chronological order use et al after name of 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4469544 695 700 in condensed form guidelines printed at the are sousa W P 1985 the back ofthe issue additional copies are available disturbance and patch dynam- ics on rocky pages from the editor upon request also check intertidal shores 101 124 the S pickett most recent issue of the great in T A and P S white eds the basin naturalist ecology for changes and refer to the of natural disturbance and patch dy- CBE style manual press ath5th edition council of biology editors one namics academic new york coulson R N illinois center suite 200 illiiiliilil111 east lackerwacker and J A witter 1984 forest drive chicago IL 60601429860601 4298 USA entomology ecology and management john PHONE wiley and sons 312 6160800616 0800 FAX 312 6160226616 0226 24 inc new york 669 appp TABLES TYPE AND DOUBLE SPACE all materials are double spaced on separate sheets and including designed literature cited table headings and figure legends to fit the width of either a single column or a page use avoid 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begin paragraphs photocopies of illustrations ISSN 17361400173614000017171 3614 GREAT BASIN naturalistnaturalistvol volwol 52 no 4 december 19924992

CONTENTS articles winter nutrient content and deer use of gambel oak twigs in north central utah rosemary L pendleton fred J wagstaff and bruce L welch 293 botanical content of black tailed jackrajackerajackrabbitjackrabbitbbitabbit diets on semidesert rangeland tchouassiTchouassi bansiwansi rex D pieper reldon F beck and leigh W murray 300 species of eimeria from the thirteen lined ground squirrel Spennspennophilusophilustophilmtopholm tri decemlineatwdecemlineatus from wyoming robert S seville diane M thomas russell pickering and nancy L stanton 309 plant agesizeage size distributions in black sagebrush artemisia nova effects on community structure james A young and debra E Palinquistpalmquist 343313 mushroom consumption mycophagy by north american cervids karen L launchbaugh and philip J urness 321 terrestrial vertebrates of the mono lake islands california michael L morrison william M block joseph R jehl jr and linnea S hall 328 vascular flora of kane lake cirque pioneer mountains idaho robert K moseley and susan bernatas 335 lakeward and downstream movements of ageageo 0 arctic grayling thymallus arc ticus originating between a lake and a waterfall

I1 mark A deleray and calvin M kaya 344 effects of browsing by mule deer on tree growth and fruit production in juve- nile orchards dennis D austin and philip J urness 352362 changes in riparian vegetation along the colorado river and rio crandGrandgrandecrangde colorado warren D snyder and gary C miller 357 resident utah deer hunters preferences for management options dennis D austin philip 1 urness and wes shields 364 list of oregon scolytidae coleoptera and notes on new records malcolm M furniss james B johnson richard L westcott and torolf R torgersen 373 riffle beetles coleoptera elmidaeElmelmidgeidae of death valley national monument cali- fornia william D shepard 378 notes siphonaptera fleas collected from small mammals in montane southern utah

I1 james R kucera and glenn E haas 382 notes on spider theridiidae salticidae predation of the harvester ant pogonoinyrmexpogonomyrmex salinusgalinus olsen hymenoptera formicidae myrmicinaemyrmicidae and a possible parasitoid fly chloropidaeflychloropidae william H clark and paul E blom 385

index to volume 52 387