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VOLUME 59 NO 2 APRIL 1999

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BRIGHAM YOUNG university GREAT BASIN naturalist httpwwwlibbyueduhttpwwwlibbyuedunmsamsnms FAX 8013783733801 378 3733 editor assistant editor richardwbaumannrighardRICHARD W BAUMANN NATHAN M SMITH 290 MLBM 190 MLBM PO box 20200 PO box 26879 brigham young university brigham young university provo UT 84602020084602 0200 provo UT 84602687984602 6879 8013785492801 378 5492 8013786688801 378 6688 emailE mail richarclbaumannbyuedurichardriehard baumannbyuedu emailE mail nathan smithbyuedu

associate editors JAMES C CALLISON JR JERRY H SCRIVNER department of environmental technology department of biology utah valley state college ricks college orem UT 84058 redburgrexburgRexburg ID 83460110083460 1100 JEFFREY R JOHANSEN STANLEY D SMITH department of biology john carroll university department of biology university heights OH 44118 university of nevada las vegas vegas BORIS C kondratieff las NV 89154400489154 4004 department of entomology colorado state ROBERT C WHITMORE university fort collins CO 80523 division of forestry box 6125 west virginia morgantown PAUL C MARSH university Morgantown WV 26506612526506 6125 center for environmental studies arizona NEIL D WOFFINDEN state university tempe AZ 85287 department of biology of JOSEPH R MENDELSON 111 university pittsburgh ililiiIII johnstown department of biology utah state university PA 15904299015904 2990 logan UT 84322530584322 5305

editorial board richard A heckmann chair zoology john D bell zoology berranjerran T flinders botany and range science duke S rogers zoology bruce A roundy botany and range science richard R tolman zoology larry L st clair botany and range science H duane smith monte L bean life science museum all are at brigham young university ex officio editorial board members include steven L taylor college of biology and agriculture and richard W baumann editor great basin naturalist the great basin naturalist founded in 1939 is published quarterly by brigham young university unpublished manuscripts that further our biological understanding of the great basin and surrounding areas in western north america are accepted for publication subscriptions annual subscriptions to the great basin naturalist for 1999 are 25 for individual sub- scriscribersbers 30 outside the united states and 50 for institutions the price of single issues is 12 allalialtbackbaekback issues are in print and available for sale all matters pertaining to subscriptions back issues or other busi- ness should be directed to the editor great basin naturalist 290 MLBM PO box 20200 brigham young university provo UT 84602020084602 0200 scholarly exchanges libraries or other organizations interested in obtaining the great basin naturalist through a continuing exchange of scholarly publications should contact the exchange librarian 6385 HBLL PO box 26889 brigham young university provo UT 84602688984602 6889 editorial production staff joanne Y abel technical editor emailE mail jyaemailbyuedu copyright 0 1999 by brigham young university ISSN 001736140017 3614 official publication date 30 april 1999 4994 99 700 29403 the great basin naturalist PUBLISHED AT PROVO UTAH BY MLM L BEAN LIFE SCIENCE MUSEUM BRIGHAM YOUNG university

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VOLUME 59 30 APRIL 1999 no 2

gleatgreat basin naturalist 592 019991999 appp 105 111

reproductive ECOLOGY OF BISON ON ANTELOPE ISLAND UTAH

michael L wolfel milan P shipka2 and john FE Kimball 3

abstractABSTRACI autumn musters ofot bison bison bison on antelope island state park utah conducted annually since 1987 provided data on temporal and age specific reproductive patterns and a basis to evaluate the efficacy of management measures implemented to elevate reproductive performance in the heidheldherd pregnancy rates were vanablevariable and low xY 46246 2 in comparison to other free ranging noncommercial bison herds in north america cows inm the 3 and gyigylgyr6 yiyr age classes exhibited lower than expected pregnancy rates P 0050.05oos0 05 annual pregnancy rates showed a significant r 0640 64 P 004700.047047 linear decline of 252.52 5 per annum variance in distribution of fetal ages observed in 6 yr indicates substantial temporal fluctuation longtermlong term reproductive performancepeipel firmanceformanceformaneemance of cohorts born prior to implementation of management measures did not differ from that of cohorts born subsequent to these changes

key words bison bison bison nutrition reproduction utah antelope island

A population of bison has existed on ante- reproductive anomalies may also exist alterna- lope island utah for over a century this herd tive hypotheses to explain the long calving per- originated from a very small n 12 founder iod are the absence of predation and a low qual population popov and low 1950 and its sub- ity and temporally unpredictable forage resource sequent population growth was punctuated by prior to 1981 antelope island state park reduction to very low levels ie 30 animals AISP encompassed only a small area at the on at least 2 occasions increasing the potential north end of the island in that year the utah of low levels of genetic variability and raising division of parks and recreation udp&r questions concerning possible demographic acquired the remainder of the island includ- effects of the small founder population and sub- ing the bison herd which was thought to sequent bottlenecks preliminary field observa- number 250 280 animals initially the populations of reproductive patterns in the herd by tion was subject to minimal management with wolfe and kimball 1989 revealed an unusually the removal of only 3 male bison reported dur- protracted calving period for this herd other ing the following 6gyryr period

department of fisheriesfisberiesandand wildlife utah state university logan UT 84322521084322 5210 department2department otof animal dairy and veterinary science utah stolestate university logan UT 84322481584322 4815 sutah division of wildlife resources 1594 west north temple salt lake city UT 84114

105 106 GREAT BASIN naturalist volume 59

in 1987 the udp&r initiated an active 327c while minimum winter dec feb bison management program featuring annual temperatures were 62c lower elevations musters selective culling inoculation of anion the island experience about 200 frost free mals artificial winter feeding of calves and days annually higher elevations about 150 limited hunting of adult males in 1987 grazing by approximately 1500 domestic cattle on METHODS the island was also terminated the annual roundups provided an opportunity to collect the udp&r has conducted an annual information on reproductive patterns in the roundup of the antelope island bison herd in population and test hypotheses regarding the the last week of october or first week of impact of these management changes on november since 1987 these gatherings employ reproductive patterns within the population helicoptershelicopters and ground support on horseback specifically we hypothesized that a presum- and in wheeled vehicles to haze the bison into ably improved nutritional plane resulting from large holding paddocks at the north end of the the management program would increase island all animals in the herd are captured in reproductive rates and synchrony of parturi- the roundup or are at least accounted for dur- tion shipka et al 1995 provided a prelimi- ing followupfollow up flights bison typically remain in nary analysis of the data obtained through the holding paddocks for approximately a 1992 this paper includes data collected sub- week prior to being processed through a set of sequently and more comprehensive analysis working chutes during the period 1987 1997 this operation typically commenced on julian STUDY AREA day 309 and lasted 3 4 d upon entering the squeeze chute adult antelope island has been described previ- animals are identified and weighed every ously by wolfe and kimball 1989 briefly it bison in the antelope island herd is perma- is the largest 104 km2 of several islands in nently identified by means of metal ear tags the great salt lake lying approximately 6 kmkni and calves are branded according to their birth from the mainland topographically the island year beginning in 1991 animals were also is characterized by a north south ridge maxi- identified by means of micromicrochipchip implants in mum elevation above lake level 600 in with their ears adult females were examined for steep west facing escarpments and generally pregnancy by rectal palpation in 1987 1988 1994 fetal more gentle east facing slopes 1990 1992 1993 and age was esti- annual grasses constitute the principal veg- mated during pregnancy palpation etation on the island with cheatcheatgrassgrass bromus during the 1987 roundup provisional ages tectoriumtectorumtectorum and threethreeawnawn aristida sp compris- were assigned to younger animals based on ing the dominant taxa these conditions are the schedule of replacement of lacteal incisi- the result of range deterioration due to over- form teeth incisors and canines as described grazing erosion and recurrent wildfireswildfires jones by hogben in larson and taber 1980 the 1985 isolated pockets of juniper juniperus series of known age cohorts established by osteospermdosteospenna and big tooth maple acer gran marking calves of the current year provided didentatumdidentatum occur primarily on steeper slopes the opportunity to verify this sequence and and canyons on the western side where they subsequently correct initial assignments of have been protected from fire there are also animals in the 1987 roundup through 4 yr old limited stands of sagebrush artemisia triden age classes kimball and wolfe 1989 the lin tata on some of these higher slopes some gual labial width of the central incisors 111 of portions of the eastern slopes have been re most older bison was also measured during seeded with perennial grasses and grass legume the 1987 and 1988 musters novakowski 1965 mixtures further estimates of age were obtained by weather records for antelope island exist counts of cementum annuannulationslations in 111 teeth only for the period 1952 1972 during that collected from 32 animals mortalitiesmortalities and interval annual precipitation averaged 39339.3 hunter killed animals 11I width data collected cm with snowfall of 25125.1 cm average maxi- during the 1987 roundup were regressed against mum summer jun aug temperatures were age estimates from cementum annulation counts 1999 BISON reproduction 107 and animals derived from known age cohorts 70 for the purposes of this analysis we established 60 the following age categories 2 3 and 4 yr 50 olds 5 6 and 6 ab9b yr yr 2 40 statistical analysis of reproductive data P uti- 30 lized procedures outlined by cochran and cox tU b 20 1957 for categorical data two discrete cate- P gories ie pregnant not pregnant were ana- 10 0 lyzed by chi square procedures for a 2 X n 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 contingency table using the following formula year

alpi pyap 1 1 average annual pregnancy rates for antelope 2 pj peaai fig X island bison cows 2 yr of age 1987 1997 aqpqP 4 where aia number in category I1 within the ith age class p overall proportion of the total we further tested the hypothesis that cohorts population in category 1 and subject to the effects of management changes iai1q I1 p overall proportion of should have a higher reproductive rate than the total population in category 2 animals born before the changes were implemented the longtermlong term 5 yr of consecutive this procedure allows for the examination of data histories of females 2 yr of age and within population differences when a known born during the period 1987 1991 were com- or expected value is not available in addition pared with similar performances of those one way ANOVA was used to evaluate the females which had already achieved reproduc- observed difference in pregnancy rates among tive age in 1987 table 1 the mean longtermlong term various age classes with years being pooled pregnancy rates of cows born during the 2 differences between individual age classes time intervals 0530.53 and 0490.49 respectively did were analyzed by means of bonferroniBonferroni t tests not differ significantly z 0460.46 P 0320.32 bartieBartlebartletttf s test for homogeneity of variance was likewise maximum numbers of consecutive used to evaluate observed differences in the pregnancies between the 2 groups were virtu- distribution of fetal ages ally identical 262.6 and 272.7 respectively t 0350.35 FP 0720.72 RESULTS one way ANOVA revealed significant dif- pregnancy rates ferenferencesces F 4054.05 P 00070.007 among mean pregnancy rates for various age classes for the pregnancy determination for the period 11 yr period 1987 1997 fig 1 bonferroniBonferroni t 1987 1997 indicated a mean annual preg- tests indicated that pregnancy rates in the 3 nancy rate of 46246.2 range 32532.5 66666.6 fig and 6 yr old groups were significantly lower 1 the rates observed in 1993 and 1997 were P 0.05005 than the other age groups 2 significantly higher xax22 40940.9 df 9 P 005 fig 000050.0005 and lower xax22 25025.0 df 9 P fetal ages 00050.005 respectively than expected regression of annual pregnancy rates against time indi- fetal ages were available for 6 yr of the study 2 cated an average annual decline of 252.5 r table and indicated that 1988 had 0640.64 P 00470.047 the value for 1987 was the lowest measures of central tendency with excluded from this analysis because we rea- a suggestion of bimodal distribution it should soned that pregnancy rates in that year were be noted however that the sample size in that not affected by management changes described year was less than half of the next lowest year above regressions using pregnancy rates the results of bartlett s test for homogeneity weighted by sample size and ludinghidingexexludingexhidingexcluding the of variance suggested a tendency P 00980.098 1993 value an unusual observation but not for this population to experience random fluc- technically an outlier yielded qualitatively tuationstuations in the variance of fetal ages over time similar results fig 3 108 GREAT BASIN naturalist volume 59

TAlabiIABIiiii1 1 reproductive performance ofantelopeofantelope island bison cows in relation to management changes maximum number sample years pregnancy of consecutive size observed rate pregnanciespi egnanciesegnanclescieseles cohortsts n Yx jx sys x STs bornboinbefoiel987before 1987 79 64 0530 53 0060 06 272 7 021 bomborn 1987 1991 61 65 0490 49 0060 06 262 6 oisols0180 18

70 table 2 summary statististics foiroirolfor estimated fetal ages of antelope island bison 60 age days ab9b 50 samplesampie year size median YX sy 40 1987 53 135 1135 399 30 1988 31 105 1031 708 Q 91 20 1990 82 105 921 349 1992 61 135 1212 326 10 1993 179 135 1322 216 1994 106 135 0 1207 290 2 3 4 56 6 age years

fig 2 age specific pi egnancyregnancypregnancy rates of antelope island bison cows 1987 1997 known reproductive histories spanning intervals of 5 8 yr produced 2 calves during those intervals while 23 and 11 produced a sin- discussion gle calf or 3 calves respectively only 13 gave birth to 5 calves pregnancy rates in the AISP herd are low nutritional stress combined with metabolic compared to some other free ranging bison requirements of lactation may limit the ability herds table 3 the only herd known to have of first calving bison cows to raise a calf and a lowerlowet pregnancy laterate than that on antelope replenish body reserves sufficient to return to island is the santa catalina herd lott and estrous cyclicitycyclicity forage quality on antelope galland 1987 suggested that the low repro- island is thought to be poor due to decades of ductive rate in that herd was likely due to overgrazing combined with erosion despite nutritional deficiency because forage quality increased digestive efficiency by bison com- appears to be low at the AISP location that pared to cattle deliberto 1993 poor forage hypothesis may also be true for antelope quality may limit energy availability to young island calves received supplemental winter bison cows bronson 1989 showed that estrous feed during the latter portion of this study cyclicitycyclicity is related to whole body energy which may have affected pregnancy rates balance furthermore suckling has a negative among 2 yr old cows puberty will be achieved effect on the return to estrous cyclicitycyclicity in at an earlier age within a species group among domestic cattle short et al 1990 and kirk- those individuals enjoying a higher plane of patrick et al 1993 have recently demon- nutrition bronson 1989 an explanation for stratedst through physiological data that lacta- the lower than expected pregnancy rate among tional suppression of ovarian activity reduces 3 and 6 yr old animals is more elusive fecundity in bison however kirkpatrick et al 1993 suggested wolfe and kimball 1989 presented data that calving rates of 35 55 imply an every on the apparently asynchronous calving pat- other year or every third year calving pattern tern of the antelope island bison population whether this applies to antelope island bison in comparison to other herds meagher 1973 is less certain but some variant of less than haugen 1974 rutberg 1986 shaw and carter every consecutive year pregnancy does oper- 1989 they reported that calves were born ate approximately 13 of the females with from march through october with 40 of the 199911999 BISON reproduction 109

60 1987 60 r 1988 n53 n31 40 40

P 4 20 h 20

0 0 45 75 105 135 165 45 75 105 135 165 days pregnant 60 r 1990 60 r 1992 n82n 82 ngin6161 40 40

u 20 20

LEELM 0 0 B B 30 45 75 105 135 165 45 75 105 135 165 days pregnant

60 1993 60 1994 n179 niogn106

40 40

20 20

0 0 45 75 105 135 165 30 45 75 105 135 165 240 days pregnant

fig 3 distribution of fetal ages in antelope island bison cows days piepregnantgnant lefersrefersrehersrebers to midpoints of monthly intervals as determineddetel mined by rectal palpation births occurring from august through octo- those 6 yr this interval was greater than 90 d ber green and rothstein 1993 used the in length this extrapolates to an asynchro- length of time from the first birth until 80 of nous period of parturition in comparison to the year s calves were born as a measure of the bison herds cited above especially consid- birbirthingberthingthing synchrony among bison in wind cave ering that the loss of any of those fetuses to national park WCNP berger and cunning- abortion or resorption would only serve to ham 1994 cited those data combined with lengthen this period during the calving season data of rutberg 1987 from the national anecdotal reports by AISP personnel confirm bison range NBR and data collected from the occurrence of very young 3 months of the badlands national park BNPBNF bison herd age calves as evidenced by their reddish tan indicating that 80 of bison calves were born coloration meagher 1979 during every month during 23 49 and 55 days at NBR WCNPWCNE of the year and BNPBNE respectively within the AISP bison synchrony of parturition has been examined herd estimates of fetal ages by rectal palpa- among populations of wild ungulaungulatesungulatedtes estes tion fig 3 indicated more than 60 d spread 1976 rachlow and bowyer 1991 and related in ages of the oldest fetuses in all 6 yr these to various causal mechanisms namely preda- data were collected additionally during 3 of tion effects and availability of forage bison 110 GREAT BASIN naturalist volume 59

table 3 pregnancy rates of some free ranging north american bison herds pregnancy rate location source national bison range 882 rutberg 1986 niobraraNiobrara national wildlife refuge 784 haugen 1974 witchitawitchataWitchata mountains wildlife refuge 718 shaw and carter 1989 badlands national park 643 berger and cunningham 1994 henry mountains 630 van vuren and bray 1986 yellowstone national park 520 meagher1973meagher 1973 antelope island 462 this study santa catalina island 350 lott and galland 1987

herds at NBR WCWWCNP BNP and AISP are cur- to determine whether the number of repro- rently not subject to natural predators preda- ductductivelyively mature females could continue to tion other than hunting by humans has not increase as it has over the last 10 yr while been a factor of consideration in any of these experiencing a decline in annual pregnancy herds for at least 100 yr in the absence offreofpreof pre rate we constructed a model to generate ani- dation environmental factors may become a mal number estimates using assumptions of more important determinant affecting syn- 90 survival rate and 50 female calf crop chrchronousonous parturition berger 1992 suggested along with the actual annual pregnancy rates that natural selection would still favor animals we arrived at estimates of reproductively that calve at some optimal time that is linked mature female herd size that were very close to the annual peak bloom of spring forage to to actual numbers of animals in this group support lactation and calf growth spring for- through 1994 estimates generated for 1995 age blooms on antelope island are predictable through 1997 were higher than actual num- but short lived furthermore minor blooms may bers of mature females observed neverthe- occur at widely disparate times of the year less growth in numerical size of this herd whenever sufficient moisture is available to observed over the period 1988 through 1997 is produce germination of annual grasses these not inconsistent with the observed rate of conditions may produce some interinterannualannual vari- decline in mean annual pregnancy rate ation in the timing of forage availability it is noteworthy that each year during the green and rothstein 1993 postulated life- roundup some 15 20 red calves are encoun- long negative effects to late born bison calves tered these animals are generally reunited citing reduced growth and reproductive suc- with their dams following processing at a cess and negative effects on early dominance minimum these cows would not have been relationships as examples ofbirthof birth order effects pregnant at that time and the return of their in addition berger 1989 further indicated calves could result in a delayed onset of estrus that earlier calving cows were in better body during the following breeding season condition and came into estrus earlier in the in conclusion the data presented here in- breeding season than later calving cows dicate that the management program imple- an interesting question is the apparent in- mented to increase reproductive performance consistency that exists between the observed in this herd has not been successful in accom- rate of decline in the mean annual pregnancy plishingplishing that objective elimination of winter rate and the continued increase in population spring grazing by domestic livestock in 1987 size 707.0 per year this herd of bison ex- may have temporarily reduced grazing pres- perienperiencesces little natural mortality has no natural sure on the island s limited forage resources predators and hunting is limited to the removal in the interim however the increased size of of 6 10 bulls per year culling practices do the bison herd and active selection for larger occur each year during the fall muster with body size have likely more than offset these the removal of approximately 10 of the ani- gains and resulted in a higher level of year mals from the herd for commercial purposes round herbivoreherbivoryherbivory supplemental feeding of 199919991 BISON reproduction illliiIII111 calves during a portion of their first year of life KIMBALL JEJ F AND MLM L WOLFE 1989 antelope island may help females achieve reproductive condi- bison studies buffalo 172817 28 29 kirkpatrick J F DFD F GUDERMUTH RLR L FLAGAN J C tion at an earlier age 2 yr it JE JC ie however MCCARTHY AND BLB L LASLEY 1993 remote monimoni- does not adequately compensate for the under- toring of ovulation and pregnancy of yellowstone lying poor forage condition on the island in bison journal of wildlife management 5740757 407 412 order to sufficiently elevate the nutritional plane KRUEGER K 1986 feeding relationships among bison of females throughout their reproductive life pronghorn and prairie dogs an experimental analysis ecology 6776067 760 770 scale likely large range improvements are the larslabsonLABSLARSONON J S AND RDR D TABER 1980 criteriacritenaofsexandageof sex and age only measure appropriate for this purpose pages 143 202 in SDS D schemnitz editor wildlife techniques manual ath4th edition the wildlife soci- acknowledgments ety washington DC LOTT DED F AND JCJ C GALLAND 1987 body mass as a factor influencing dominance status in american bison we are grateful to personnel of the utah cows journal of mammalogy 6868368 683 685 division of parks and recreation in particular MEAGHER MMM M 1973 the bison of yellowstone national M larsson J filpot T smith and K sherman park national park service monograph series 1 for assistance and logistical support provided 161 appp pages throughout the of the study numerous 1978 bison 123 133 in JLJ L schmidt and course DLD L gilbert editors big game of north america faculty members and graduate students of the stackpole Harnsharrisburgburg PA departments of fisheries and wildlife and novakowski NSN S 1965 cemental deposition as an age rangeland resources utah state university criterion in bison and the relation of incisor wear provided valuable assistance with data collec- eye lens weight and dressed bison carcass weight to age canadian journal of zoology 4317343 173 178 efforts finally the advice tion we appreciate on POPOV BHB H AND JBJ B low 1950 game fur animal and statistical analysis provided by drs DV sisson fish introductions into utah utah state department and R canfield as well as thoughtful reviews of fish and game miscellaneous publication 4 salt of the manuscript by drs T deliberto and T lake city UT 85 appp morton RACHLOW JLJ L AND RXR T BOWYER 1991 interannualInterannual variation in timing and sychronysychsynchronyrony of parturition in dall s sheep journal of mammalogy 7248772 487 492 literature CITED RUTBERG AT 1986 lactation and fetal sex ratios in american bison american naturalist 12789127 89 94 BERGER J 1989 female reproductive potential and its 1987 adaptive hypotheses of birth synchrony in apparent evaluation by male mammals journal of ruminantslummants an interspecific test american naturalist mammalogy 7034770 347 358 130692130 692 710 1992 facilitation of reproductive synchrony by SHAW JH AND TS CARTER 1989 calving patterns among gestation adjustment in gregarious mammals a new american bison journal of wildlife management hypothesis ecology 7332373 323 329 5389653 896 898 BERGER J AND C cunningham 1994 bison mating and SHIPKA MPM P MLM L WOLFE AND JEJ F KIMBALL 1995 popu- conservation in small populations columbia univer- lation structure and reproductive patterns in an sity press new york island bison population page 123 in second annual BRONSON FH 1989 mammalian reproductive biology conference of the wildlife society portland OR university of chicago press chicago 12 17 september 1995 COCHRAN WG AND GMG M cox 1957 experimental SHORT RE RAR A BELLOWS RBR B staigmiller JGJ G designs john wiley & sons inc new york berardeinelliBERARDE INELLI AND EEE E CUSTER 1990 physiolog- DELIBERTO TJ 1993 comparative digestive physiology ical mechanisms controlling anestrus and infertility of american bison and hereford cattle unpublished in postpartum beef cattle journal of animal science doctoral dissertation utah state university logan 6879968 799 816 ESTES RDR D 1976 the significance of breedingofbreeding synchrony VANVUREN D AND MTM P BRAY 1986 population dynamics in the wildebeest east african wildlife journal 14 of bison in the henry mountains utah journal of 135 152 mammalogy 6750367 503 511 GREEN WCHWC H AND A ROTHSEIN 1993 asynchronous WAGGONER V AND M HINKES 1986 summer and fall parturition in bison implications for the hider fol browse utilization by an alaskan bison herd journal lower dichotomy journal of mammalogy 7492074 920 925 of wildlifeofwildlife management 5032250 322 324 HAUGEN a0aaA 0 1974 reproduction in plains bison iowa WOLFE MLM L AND JFJ F KIMBALL 1989 comparison ofbisonorbisonofbison state journal of research 49149 1 8 population estimates with a total count journal of HOLLANDER M AND DAD A WOLFE 1973 nonparametricnonparametncNon parametric wildlife management 5359353 593 596 statistical methods john wiley & sons new york JONES C D 1985 A manual of the vascular flora of ante- received 20 july 19919977 lope island state park davis co utah unpublished accepted 28 april 1998 thesis brigham young university provo UT 110 appp cleatgleatgreat basin naturalist 592 019991999 appp 112 126 remarkable WAXING WANING AND WANDERING OF populations OF MIMULUS GUTTATUS AN unexpected EXAMPLE OF GLOBAL WARMING

robert K vickeryjr1vickery jr 1

ABSIRAC 1 the purpose of this study was to observe the dynamics of a meta population of mimulus guttatusgutgustatustatus changes in size and location of 16 original populations and the new populations established in their vicinities in big cottonwood canyon salt lake county utah weiewelewere observed for 25 yr twenty three new populations appealedappeared seven 01 iginaloriginal populations and 13 new populations had become extinct by the end of the observation period in 1996 many populations died out and weiewelewere establishedreestablishedle often repeatedly during the observation period altogether there were 54 population disappearancesdisappeaidisappear ancesanees and 34 reappearances many populations changed size as much as 100 fold or more from yeaiyealyear to yeaiyealyear there were spectacular examples of populations expanding to fill newly available large habitats frequentfleFiepiequent extinctionsextinctions were due overwhelmingly to the canyon drying trend which led to the drying up of most mill D noinorthth drainage springs creeks and ponds precipitation and minimum temperatures increased moderately during the observation period therhe growinggi owing season lengthened almost 50 a typical consequence of global warming the drying trend lengthened growinggi owing season and disappearance of mimulus populations in big cottonwood canyon appear to be a cleaiclearelear local example of global waiwalwarmingming

key words mimulus guttatusgustatusguttatus meta population global warming growing season colonization recolonization extinction populationpopulationfluctuationsfluctuations

populations of mimulus guttatusgustatusguttatus fischer ex the selective pressure versus gene exchange DCD C in the upper reaches of big cottonwood study alluded to above vickery 1978 for that canyon form a meta population iei e an inter- investigation I1 selected 4 side canyons of big acting group of local populations van der cottonwood canyon fig 1 table 1 each meijden et al 1992 this conclusion is based side canyon or drainage is separated from the on my earlier study of gene exchange and others by high ridges of up to 3000 m or more selective pressures in these populations vick- each contains a creek that runs from the side ery 1978 the purpose of the present longitu- canyon headwaters at ca 2750 m down to its dinal study was to observe for a long enough confluence with big cottonwood creek at period 25 yiyr the dynamics that is speed of 2250 m I1 selected 4 study sites as equally turnover of the local M guttatusgustatusguttatus populations spaced as possible along each of the creeks constituting the meta population how quickly each study site contained 1 population ofofmM and frequently do they increase or decrease in guttatusgustatusguttatus the original study population table size die out colonize new habitats or recolodecolo 1 most study sites had I1 to 6 or more suitable azenizemze old ones how do these changes relate habitats for local M guttatusgutgustatustatus populations al- to the environment precipitation insolation though there was only the original study pop- temperature and growing season ulation at the beginning of the observation period STUDY SITES study sites for this moisture loving species were in habitats by streams or springs or foifor this research I1 studied a sample of 16 of along pond or lake shores some were in full the approximately 170 local populations of M sun but more were in partial shade of willows guttatusgustatusguttatus in the upper reaches above the ter- small alders or young aspen trees while a few minal moraine at 2150 m of big cottonwood were in open areas or light gaps in the denser canyon salt lake county utah fig 1 I1 ob- shade of the spruce forest habitats tended to served the same set of populations studied in be unstable and transitory that is the stream

iiiliiiliology iepartmentimpartmentdepal tineitineltinoi it universityuniveiuniver oty ofutdiot utalibutali salt lake city UTU r 84112

112 1999 META population OF mlmulusMIMULUS GUTTATUSCUTTATUS 113 gobblers knob I1 D north 2923m 95001 PNC N 2615m SW8500 DIVA 2297mv7mvam 7097500

2461m8000 ig cottonwood mill south 17reek 2769m 9000 mill F east DSB FEApea ailltmillt south 7feb 3076m 10000 fs44 FEC DSC FSB PSC FED FSD si vervey lake toatox 3385m 1I10nlom clayton peak mt superior 0 0 iamikm mt millicent

fig 1 map of the upper reaches of big cottonwood canyon in the wasatch mountains salt lake comitycounty utah side canyons drainages and locations of study sites areai e indicated based on maps of the area of the USU S geological survey washington DC 20242 would erode them ponds or lakes would scattered often isolated local populations and change level and bushes and trees would only occasionally meta populations M gutta grow thus shading the populations or most tus consists of at least 6 ecoecotypestypes personal importantly streams would decrease or dry observation including the wasatch moun- up thereby killing the plants also in some tains one to which the constellation of local cases deer or moose trampled the populations populations of big cottonwood canyon or pikas ate them belongs this ecotype is distinguished by its M guttatusgutgustatustatus local populations were sepa- numerous rhirhizomeszomes but otherwise it is rather rated one from another by spaces of several typical of the species the species may be meters often as much as 050.5os km in which annual or rhizomatous perennial stems are there were no mimulus plants these spaces erect 2 60 cm high leaves arearc in opposite typically were unsuitable for monkey flowers pairs glabrous to pubescent rounded ovate and were often filled with bushes trees 1 10 cm long X 1 6 cm wide with dentate andor rocks margins lower leaves are petiolate upper leaves sessile inflorescences are racemose MIMULUS CUTTATUSGUTTATUS often few flowered or solitary on taller plants the flowers are mostly in pairs calyces are mimulus guttatusgutgustatustatus fischer ex DC ranges campanulate 6 12 mm long much inflated in from northern mexico to the aleutian islands maturity the upper tooth longest corollasCorollas are type locality and from the pacific coast to 1 2 cm long the throats spotted with red and the rocky mountains it has gone wild in the palate ridges hairy upper lips of the corol europe A moisture loving species that forms las are reflexed and much shorter than the 114 GREAT BASIN naturalist volume 59

TABLE 1 locations of study sites by elevation latitude and longitude and characteristics of habitats within the study sites of the original and new populations by location relative to the original population area insolation average maxi-maxi mum and minimum temperatures foiforhorbor the 50sod d heart of the season 4 july 23 august 1973 moimorfoifor the original populations soil type and source of moisture

MILL D NORTH DRAINAGE southwest facing DNA study site elev 2350 m 403857n4003857n 1h389w111389v DNA 01 igbaligmaloriginal population 050.5os0 5 m2ma partial shade 180531805.318 05 3 sandy gravel streamside DNA new population 8 m down and across stream 101.0iolo1 0 m2ma full sun sandy gravel streamside DNB study site elev 2520 m 403943n4003943n lir3740w1113740v DNB original population 30 m2ma full sun 1332613.32613 32 6 moss and gravelly soil by large spring DNC study site elev 2670 m 403940n4003940n nr374w111374w DNC original population 2 m2ma partial shade 153431534.315 34 3 grassy soil by beaver pond DNC new population 20 m downstream ca 350 m2ma deep loam moist pond bottom DNC new population 40 m downstream ca 150 m2ma deep loam moist pond bottom DND study site elev 2760 m 403935n 1113638v11p3638w DND original population 252 5 m2ma open shade ofaspenof aspen 1434014 34 0 sandy gravel along a nilniirill DND new population 30 m downstream 6 m2ma sandy gravel stream delta

MILL D SOUTH DRAINAGE CARDIFF CANYON northeast facing DSA study site elev 2277 m 403827n IHTW DSA original population 151.5isls1 5 m2ma light gap in dense shade 1926819 26 8 sandy gravelly soil strearstreanstreamsidestrearasideaside DSA new population 100 m upstream ca 24 m2ma light gap in dense shade loamy soil boggy spring by stream DSB study site elev 2474 m 403734n 113914w1113914v DSB original population 12 m2ma full sun 1434114 34 1 moss gravel and pebbles streamside DSB new population 3 m across stream 14 m2ma full sun moss gravelly soil streamside DSB new population 10 m downstream 1 m2ma partial shade sandy soil streamsidestrearnside DSC study site elev 2628 m 4037tn40371n4037TN 13914v11103914wI1 I1 DSC original population 2 m2ma partial shade 188421884.218 84 2 loamy soil by stream DSC new population 8 m downstream 010 1 m2ma partial shade loamy soil by stream DSD study site elev 2763 m 403625n 1h3919w1113919v DSD original population 18 m2ma full sun shade at edges 1414514.14514 14 5 deep loam in boggy spring DSD new population 20 m upstream 3 m2ma partial shade loamy soil by small rill DSD new population 15 m downstream 3 m2ma full sun gravelly soil by small rill

spreading lower ones capsules are oblong of plants of one canyon population in the phy seeds are football shaped ca 050.5os mm wide X totroncotron at cal tech vickery 1974 this plastic- 101.0lo mm long and longitudinally striate for ity was confirmed twice by growing sets of the fuller descriptions see grant s monograph 16 original study populations in uniform grant 1924 or the treatment in the jepson greenhouse environments for the earlier set manual thompson 1993 vickery 1978 plants were scored for over 100 flowers which are pollinatedpollinated by bumble traits and compared statistically there were bees and occasionally by other insects produce no significant differences for the more recent numerous seeds 50 to 250 per capsule set appendix A plants were scored for a dif- reproduction is from seeds but in this eco- ferent smaller set of traits and compared visu- type it is commonly from underground rhi ally there were no apparent differences zomes as well M guttatusgustatusguttatus plants of the canyon vary mor- METHODS phophologicallylogically both within and among populations see voucher specimens in the garrett I1 counted or estimated the number of plants herbarium UT university of utah canyon in each of the original study populations each plants range in height from I1 cm to 30 cm in year beginning in 1972 and concluding in 1996 leaf length from 1 cm to 5 cm and in number actual counts were made for small popula- of flowers from none to a dozen or more dif- tions estimates made for medium sized and ferencesferences among plants apparently are plastic large populations were based on counting 50 responses to the varied environments of the or 100 plants in an area and then multiplying canyon much as I1 showed for clone members by the number of such areas in the population 1999 META population OF MIMULUS GUTTATUSCUTTATUS 115

TABLE 1 continued

MILL F EAST DRAINAGE GUARDSMAN PASS west facing FEA study site elev 2524 m 403724n 1113525w FEA original population 24 m2ma deep shade 135601356.013 56 0 moss and sand bars in and beside stream FEA new population 100 m downstream 1 m2ma full sun gravelly soil streamside pebFEEFEBpee study site elev 2615 m 403720n 111353v11p353w FEB original population 2 m2ma open shade 169311693.116 93 1 grassy soil streamside FEEFEB new population 12 m downstream 1 m2ma open shade sand bar in stream FEB new population 3 m across stream 5 m2ma open shade loamy bank edge of stream FEEpebFEBpee new population 5 m downstream 151.5lsis1 5 m2ma open shade sand bar in stream FEC study site elev 2670 m 403656n 1113733w11p3733w FEC original population 6 m2ma partial shade 144391443.914 43 9 loamy soil by stream FEC new population 10 m upstream 16 m2ma open shade sand bar in stream FEC new population 30 m upstream 24 m2ma sunny grassy loam by stream FEC new population 7 m upstream 10 m2ma sunny grassy loam by stream FED study site elev 2830 m 403634n I113339v11p3339wI1 FED original population 0500.5os 5 m2ma partial shade 166511665.116 65 1 gravelly loam streamside FED new population 5 m upstream 15 m2ma sunny loamy soil by stream FED new population 15 m downstream 1 m2ma sunny sandy gravel streamside

MILL IT SOUTH DRAINAGE SOLITUDE CANYON northeast facing FSA study site elev 2528 m 40374n 1113530v1113530w FSA original population 2 m2ma sunny 135601356.013 56 0 grassy soil streamside FSA new population 5 m upstream 8 m2ma full sun loamy soil and gravel streamside and min stream FSB study site elev 2560 m 40370 N 1113540v1113540w FSB original population 15 m2ma open shade 17653o17653017 65 3 sandy soil streamside FSB new population 20 m south along new channel 12 m2ma full sun pockets of sandy soil amongst cobbles in stream FSC study site elev 2708 m 403647n 1113551v11p3551w FSC original population 8 m2ma sunny 132361323.613 23 6 loamy soil by small spring FSC new population 20 m downstream ca 600 m2ma full sun in new grassy loamy meadow near stream FSD study site elev 2770 m 403629n 1114056v1114056w FSD original population 10 m2ma sunny 136371363.713 63 7 deep loam edge of lake solitude FSD new population 25 m north along lake shore 12 m2ma sunny deep loam lake shore FSD new population 50 m west along lakeshorelakesborelakelakes borehoreshore ca 12000 m2ma full sun deep loam exposed lake bottom to obtain the estimated population size num- air temperature of each study site in 1973 bers were rounded off A rosette was consid- table 1 was measured using temp scribe ered to be a plant in some cases I1 found that recorders vickery 1978 temperature pre- 2 or 3 rosettes were connected by under- cipitation snovtacksnowsnowpackpaekpack and growing season data ground stems stems would decay as the sea- for the entire 25 yr observation period were son progressed so that rosettes became inde- obtained from US climatological data for pendent plants although some would share utah environmental data services 1972 1996 the same genotype as observed by waser et for silver lake 403348n4003348n 111354wellsw at al 1982 for other populations of mimulus the head of big cottonwood canyonlllswfig 1 guttatusgustatusguttatus population counts were made in late summer or early fall as populations were at RESULTS their maximum toward the end of the growing season random duplicate counts were made populations ranged widely in size from a to check the accuracy of counts and estimates single plant to 37000 plants appendix B they varied by 10 to 20 which is modest some populations did not vary in size from considering the very large differences noted year to year while others waxed or waned with among different local populations the same time still others varied dramatically over the year or among different years of the same pop- period of a few years figs 2 5 ulationulation the vicinity of each population was of the 16 original study populations 7 had checked to see if new populations had become disappeared by the end of the 25 yr observation established if they had their locations were period appendix B not only were they gone noted and they too were scored as above but each one had disappeared and reappeared 116 GREAT BASIN naturalist volume 59

DNA DNADAM DNB DNC DNC DND DND ign1972

M1976 i

ism1981

19861996

1991

1996

1 19 0 100 300 1000lom 4300 0 20 95 0 350 900 0500015000 40000 Ffigig 2 mill D northnoi th drainagediadladidinage representation of changing sizes of the original study populations and new populations ovelover 25 yiyr of observationabseiobsei vation at least once prior to its final disappearance the number of populations climbed from of the 9 original populations still present at the original 16 to an average of 23 table 2 the end of the observation period 4 had disapdigap once the higher value was attained there was fearedpearedpeaipeal ed and reappeared only 5 of the original a dynamic equilibrium of populations appear- populations had been present throughout the ing disappearing and reappearing numbers study figs 2 5 climbed as high as 30 and declined to as low twenty three new populations almost I1 per as 15 table 2 fig 6 the establishment of year appearedappeal ed in study sites in the vicinity of populations depended on availability of suitable the original populations figs 2 5 appendix moist without too much shade or plant com- B they disappeared and reappeared like the petition habitats within each of the 16 study original populations at the end of the obser- sites vation period only 10 new populations were the disappearance of populations reflected still present and only 4 had been consistently loss of suitable habitats the most common present since they appealedappeared cause being loss of moisture drying could be 199919991 META population OF MIMULUS CUTTATUSGUTTATUS 117

DSA DSA DSB DSB DSB DSC DSC DSD DSD DSD 1972

1976

iggi19912982 4 1

1986

T 1991iggy

1 1996 I 10002000 45004 1 19 100 300 0 W 500015000 40000 & 20 95 A 350 900 0

fig 3 mill D south drainage cardiff canyon representation of changing sizes of the original study populations and new populations over 25 yr of observation

transitory ie short term 1 3 yr or pro- discussion longed ie long term more than 3 yr altogether there were 54 cases of populations dis- the most striking results of my observa- appearing table 2 twenty six were due to tions were fourfold 1 the remarkably large short term drying and 12 to longtermlong term drying changes in population size from one local pop- in addition to drying out 7 populations disap- ulation to the next and from one year to the peared because erosion removed their habi- next for the same population 2 the high fre- tats another 4 were lost apparently due to quency with which populations appeared competition and 5 more to shading from a both new ones and old ones after an absence dense overstory table 2 3 the high rate at which both new and old 118 GREAT BASIN naturalist volume 59

FEA nnaFEA FEB FEB FEB FEB FEC FEC FECEECpec FEC FED eedFED eedFED 1972

1976 I1

1982

198629861996

iggi19911992 4 0

1996 i 1 19 0 100 300 0 1000 4500 20 95 0350350 900 0 5000 40004000040.000 fig 4 mill F east diadladrainageinage representation of changing sizes of the original study populations and new populations over 25 yr of observation populations disappeared and 4 probably the be limiting eg dense meadow grasses but most important the implications of population usually it was not a determining factor insola- disappearance for global warming tion the amount of sunlight rarely was limiting except in the case of deep shade precipi- population sizes tation rain and snow at the actual habitat was the size of a population reflects available irrelevant because this moisture loving species habitat area which in turn reflects soil type required habitats to begin with that were plant competition and climatic parameters of steadily moist eg stream banks however insolation precipitation and temperature soil precipitation for the canyon as a whole was type did not appear to be limiting inasmuch as relevant for recharging the aquifers that sup- M guttatusgutgustatustatus grew on all substrate types from plied the springs and streams very wet years sand bars to loamy bogs competition could appeared to enhance competition at the expense 1999 META population OF MIMULUS GUTTATUS 119

FSA FSA FSB FSB FSC FSC FSD FSD FSD 1972

1976

1981

1986

iggi19912992

1996299619

21 2919 0 loo100200 300 10002000 4500 20 95 0 350 900 0150005000 40000 fig 5 mill F south drainage solitude canyon representation of changing sizes of the original study populations and new populations over 25 yr of observation

ofmof M gutguttatusgustatustatus population size temperature of 1994 the hottest year of the study with those the actual habitat was relatively unimportant in 1993 the coldest year of the study see as mimulus guttatusgutgustatustatus has a wide norm of reac- appendix B tion for temperature vickery 1974 however population sizes could be roughly grouped for the area as a whole temperature was into 3 categories small populations 1 to 20 important particularly because it shortened or were usually part of a sequence in the process lengthened the growing season menenimyneni et al ofpopulation decline toward extinction in many 1997 warmer temperatures did appear to cases these populations would grow smaller lead to larger mimulus populations as seen and smaller disappear reappear and finally dis- for example in comparing population sizes in appear for the remaining observation period 120 GREAT BASIN naturalist volume 59

taineTAISIL 2 population appearancesappeal ances disappearances numbers and parameters of the environment from US weather appealappearancesancesanees disappearancesdisappeaiances new popula- lecolrecolgrecol drying drying compe- yeaiyealyear tions omzedonizedionized short termteitel m long term tition shading erosion 1972 1973 0 0 1974 2 0 1975 4 0 FEB 1976 3 0 DNA FEC 1977 1 0 DNA DND FEA 1978 1 2 DNC 1979 1 2 1980 1 0 DNA DSD

totals 13 4 5 1 1 0 2 1972 1980

1981 1 1 FEA 1982 2 3 DND 1983 0 0 FEC FEA FEA FED FSB 1984 0 1 FEC 1985 1 1 DSA DNA FEBFEE DSC 1986 2 4 1987 1 1 1988 1 0 DNA DND DND DSC FEB 1989 1 3 FED DSC 1990 1 2 FEA DNC DSC FEB FEC FSB 1991 0 1 DSD FEC DSB FED 1992 0 0 pebFEBFEEpee FEC FED DNC FSA FSA DSD 1993 0 4 FEC 1994 0 2 DNA FSA DSD 1995 0 4 FSA FSCFSQ FEC 1996 0 3 FEB DSA

totals 10 30 21 11 3 5 5 1981 1996 totals 23 34 26 12 4 5 7 1972 1996

medium sized populations 20 to 900 usually improved the ski run but opened a large reflected stable habitats sizes of these popu- sunny moist meadow with little plant compe- lations were limited by the carrying capacity tition mimulus immediately invaded via seeds of their habitats and often fluctuated around a from FSC and upstream populations where value well below the observed maximum there had been no mimulus in 1978 in 5 yr number of plants large populations 1000 or time there were 37000 plants in this new more grew inm large favorable habitats large excellent habitat fig 5 appendix B the close relationship between population in time the original FSC population declined size and habitat area was vividly demonstrated as its spring source wandered off another ex- in several cases for example solitude ski ample of a population explosion occurred at resort bulldozed a large 30 m X 50 m thicket the DNC study site the adjacent beaver pond of willowsofwillows at the FSC study site this not only dried up in 1977 exposing a large 30 m X 40 m 199911999 META population OF MIMULUS GUTTATUS 121 service records for silver lake big cottonwood canyon etabutahutab july augustau ust avgaag winter season temperaturestempel aturesagures july 222 2cac C crowinggrowing seasonn total aug total snow num- popula- max minmm precipprecia precipprecia pack ber of I I tions 1 QC 1 QC mm mm m start end days 16 226 53 27 1123 115 912 16 208 55 104 1131 134 65 18 218 53 64 1073 136 69 713 23 21 211 53 31 1258 122 626 828 62 22 221 62 32 434 70 626 828 62 20 211 62 136 669 51 530 918 110 22 214 52 63 1246 117 616 817 61 25 213 58 72 950 100 69 914 96 24 220 64 47 1162 126 616 820 64

Tx 20 216 57 64 1005 108 614 824 68

25 227 76 55 905 70 615 926 102 29 212 66 127 1422 139 69 912 94 24 1500 138 614 920 97 24 209 64 132 1529 141 611 922 102 22 225 61 66 1294 125 530 913 105 28 211 69 140 1401 118 523 916 115 30 199 56 155 968 75 526 917 113 26 232 74 18 794 67 531 911 102 28 218 76 94 1021 102 622 825 63 25 216 70 71 880 83 107

22 215 74 132 967 93 528 915 109 15 211 64 137 847 70 510 826 107

18 193 43 131 1345 127 624 830 66 17 238 81 99 621 107 138 18 216 66 57 622 922 91 19 232 76 47 531 917 108 x 23 217 68 97 1144 104 65 914 101

x 22 217 64 85 1087 105 610 98 90

expanse of moist pond bottom into which except inm DNB with its large x 6500 origi- mimulus seeds invaded from the DNC popu- nal study population that effectively filled all lation the new DNC population steadily suitable habitats in the vicinity of its large increased for a decade from 0 to well over spring 8000 plants fig 2 appendix B populations are easily established or reestablished from seeds floating downstream populations appearing from upstream plants that often overhang the populations appeared and reappeared in stream and shed their seeds into the water suitable moist habitats along stream banks by lindsay and vickery 1964 waser et al 1982 ponds along lake shores and by or in tributary also populations may be established by wind- springs such habitats were colonized in all 16 blown seeds vickery et al 1986 windblown study sites near the original study populations seeds rarely travel more than 5 m whereas 122 GREAT BASIN naturalist volume 59

1972 1976 1981 1986 1991 1996 30- 1 1 27- 2 24 MA 8 21- 0 18

15 4 T 1600

cc lowlom CL ES 2 E 1000

700

400 82-

D iai3 62 CL 525.2

4o2m

140-

0 iioilo110 2 C 80- 0 ca 9 R 3 50- 2 20 1972 1976 1981 1986 1991 1996

fig 6 plot of population number annual precipitation minimum average temperatures for july august and length of growing season against years the growing season is the period between the last spring minimum temperature of 22c2 2cac 28f and the first fall minimum of 22c2 2cac 28f water borne seeds can travel for hundreds of seeds in the seed bank at that site thompson meters 1987 leek et al 1989 occasionally I1 have the actual source of the raitrairrain of floating observed dormant rhirhizomeszomes under rocks mimu- seeds for population establishment or relstabreestab lus gutguttatusgustatustatus seed bank seeds can readily sur- lishmentlishment could be identified in a few cases vive for 2 or 3 yr but not in large quantities for example for the new dsdpopulationDSD population the vickery unpublished study have been the original study seed source must populations disappearing population DSD because there were no other upstream populations windblown seeds can disappearances of populations both origi- be inferred as the source for some populations nal study populations and newly established eg for the new FED population where seeds ones were due to habitats becoming too shady must have come from the only nearby but or too crowded with competitors or to their downstream population FED being eroded away by the stream or trampled mimulus gutguttatusgustatustatus populations that reappear by animals however the disappearance of in 1 2 or even 3 yr at the same site probably do populations was due overwhelmingly to habi- so from dormant rhirhizomeszomes or more likely from tat drying table 2 199919991 META population OF MIMULUS GUTTATUSCUTTATUS 123

the disappearance of several populations global warming was caused by shading of habitats for exam- what caused the drying trend counter increasingly ple FEA was shaded out by an intuitively decreased precipitation was not the dense stand of DSC popula- spruce trees the cause fig 6 beginning in 1981 precipitation tion was first partially eaten then trampled actually increased from an average of 1000 mm and finally shaded out by the growth of willow to an average of almost 1150 minmm table 2 thickets the pattern of decline due to shading insolation presumably remained constant was a gradual decrease in population size increased temperature probably played a role accompanied by a decrease in plant size and however the july august average maximum amount of flowering finally resulting in no temperature was not higher but the average monkey flower plants increasing competition minimum temperature was PC higher these led to much the same pattern of decline for ex- temperatures follow the same pattern noted ample DNC declined to extinction as meadow by karl in his global warming study of the arc- grasses of the site increased and formed a tic slope of alaska also beginning in 1981 dense turf the stream swept away some pop- waring and running 1998 in big cotton- ulatulationsions particularly ones established on sand wood canyon the growing season fig 6 bars such as FEB and FEB much as lengthened by almost 50 from an average of porath et al 1987 observed for the fugitive 68 d 1971 1980 to an average of 101 d species verbascum sinaiticumsinaiticum on the gravel 1981 1996 see table 2 lengthening of the bars ofofladisofwadiswadis in the negev desert growing season is the hallmark of global canyon drying trend warming according to a satellite study of the northern latitudes menenimyneni et al 1997 the drying of population habitats was part of a drying trend in big cottonwood canyon pronounced drying trend in big cottonwood appears to be a specific local example of canyon that began in the late 1970s and early global warming 1980s in 1981 on average the drying trend how does a lengthened growing season strongly affected mill D north and mill F translate into the observed drying trend the east drainages both of which are southwest immediate result of a longer season is increased facing meaning they have high insolation evapotranspiration from the aspen forest and also their aquifers are shallow D scheck its understory of shrubs herbs and grasses J personal communication the whole of the ehleringer personal communication the con- initially flowing mill D north creek dried up sequence of increased evapotranspiration is except for the moist beaver pond bottom at the drying out of surface soil which may lead DNC and the large spring and stream imme- to a failure to recharge underlying aquifers diately below it at DNB see fig 1 the 4 which are shallow anyway D sheck personal original populations had increased to 7 in the communication depletion of the aquifers early 1980s but as the drying trend set in results in springs drying up which in turn populations declined to just 2 by the early leads to streams and ponds drying up since 1990s and those 2 were rapidly waning by the there is considerable normal variability in pre- end of the observation period appendix B cipitation temperature and length of growing in the mill F east drainage the top spring season and hence of spring and stream flows and ca 100 in of stream below it dried up as mimulusMimultis populations often dry out and die only well as the bottom ca 200 in of stream the 4 to be briefly reestablished as the drying trend initial populations had increased to 13 in the proceeds the drying out and disappearance course of the observation period by the end of so many local populations of the extensive of the period 5 populations had dried out and meta population ofofmM guttatusgutgustatustatus in big cotton- disappeared all 3 of the top and both bottom wood canyon was due to the pronounced populations four intermediate level popula- canyon drying trend the root cause of which tions had disappeared due to erosion the appears to be global warming original FEB populations appeared to be on the way to extinction however 3 of the FEC acknowledgments site populations were flourishing altogether over 13 of the population had disappeared I1 thank yaw en chu stephen purcell and due to the drying trend david vickery for their help with the weekly 124 GREAT BASIN naturalist volume 59

hiking to change the temp scribes I1 thank THOMPSON DMD M 1993 mimulus monkey flower in JCC for much and hickman editor the jepson manual higher plants jeanette stubbe patient typing press young for his of california university of california berke- richard expert photography of ley 1400 appp the illustrations I1 thank mr dan sheck salt THOMPSON K 1987 seeds and seed banks new phytolo- lake city hydrologist and drs mary price gist 106log Supplement 23 34 and jim Ehlenehleringernger for their helpful advice VAN DER MEIJDEN E PGLPG L klinkhamer TJ DEJONG AND CAM VAN WIJK 1992 meta population dynam- lesiesics of biennialofbiennial plants how to exploit temporary habi- CITED literature tats acta biologicabiologicalBiolo gica Neerlandneeilandicaneerlandicaleaicaiea 4124941 249 270 VICKERY RKR K JR 1974 growth in artificial climates an environmental dalaDAIADATA SERvicservicesSLRVICESrs 1971 1996 utah climachmacilmaachma indication of mimulus ability to invade new habitats tological data volumes 73 98 national oceanic and ecology 5579655 796 807 atmospheric admimstiadministrationatlon US department of 1978 case studies in the evolution of species Commecommercecommereeieeicelee asheville NC complexes in mimulus evolutionary biology 11 GRANGRANT 1 AL 1924 A monograph of the genus mimulus 405 507 annals of the missouri botanical garden 119911 99 389 VICKERY RKR K JR DRD R PHILLIPS AND PR WONSAVAGE lekleokLLKLLCK MA VT PARKER AND RL SIMPSON EDITORS 1986 seed dispersal in mimulus guttatusgustatusguttatus by wind 1989 ecology of soil seed banks academic press and deeldeer american midland naturalist 116206116 206906gog 208 san diego CA 462 appp WARING RRHH AND SWS W RUNNING 1998 forest ecosys- LINDSAY DW AND RK VICKERY JR 1964 natural dis- tems analysis at multiple scales and2nd edition acad- persal of mimulus guttatusgutgustatustatus proceedings of the utah emleemicemie press harcourt brace & co san diego CA academy of science aitsaltsarts and letters 4123741 237 241 370 appp menenimyneniMYNI NI RBR B CDC D KEELING CJC J TUCKER G ASRAR AND WASER NMN M RKR K VICKERY JR AND MXM V PRICE 1982 RR NEMANINLMANI 1997 inclinciincreasedeased plant glowthgrowth in the patterns of seed dispersal and population differentia- northernnornonthein high latitudes from 1981 to 1991 nature tion in mimulus guttatusgutgustatustatus evolution 3675336 753 761 386697386 697 702 PORAIIIPORATH D M AGAMI AND G ORON 1987 compliance received 11 august 1997 of a biennial mullein verbascum sinaiticumsinaitic um benth accepted 22 june 1998 to enaticerratic desert floods in the negev highlands israel journal of andaridarld environments 1312913 129 136

appendixAPPLNDIX A characteristics of the 16 original study populations when grown under standard conditions in the biology department greenhouse university of utah

plant days to drydrywtdrywatwt drywtdrydrywatwt largest leafittafsafsat at ist healthy typical popula- height flower- no of stems runners flower cm flower cm tiontpionttion cm ing flowers 9 g length width length width DSA 24 69 9 24 59 61 43 41 25 DSB 30 86 15 11 32 77 31 31 25 DSC 26 69 24 31 14 68 48 29 26 DSD I1L DNA 38 67 21 15 46 78 49 28 25 DNB 49 52 38 26 24 52 39 23 19 DNC 34 44 21 07 14 47 34 22 24 DND 33 56 20 12 15 56 44 24 19 FSA 65 56 37 24 22 FSB 30 62 20 23 46 56 44 27 23 FSC FSD 50 66 63 26 23 FEA 35 59 35 29 35 90 53 29 25 FEB 40 65 63 24 18 72 41 26 21 FEC 30 74 20 18 64 59 37 28 25 FED 62 93 45 26 24 T 34 63 26 20 33 66 43 27 23 till1113ased1113Based on ththe ineansindansme ins 45of 5 plantspi mts fccsceseescc1ab1cscc 1 ibleiblc I1 toifoifol populationpopulkopul ution sitsite datacl it t virwirtisvii us killed piplantsints 1999 META population OF MIMULUS CUTTATUSGUTTATUS 125

APPENDIX B sites and sizes of original and new populations site abbreviations are described in table 1 year DNA onaDNA DNB DNC DNC DNDD ND DND

MILL D NORTHnoRTH DRAINACDRAINAGE E 1972 100 2000 500 200 1973 700 5000 200 550 1974 30 2500 350 400 1975 2 1 6000 250 300 1976 0 450 20000 1250 2000 15001 500 1977 0 0 1000 12 200 0 1978 0 100 10000 0 50 1 1979 1 150 7000 0 1500 35 1980 0 50 3000 0 600 60 1981 0 35 8500 1100 1700 6 1000 1982 4 146 27800 50 350 0 260 1983 3 36 19000 200 1500 0 100 1984 7 7 3500 350 8500 100 150 1985 0 8 6000 1200 8000 25 500 1986 0 10 2900 200 3000 1 5 1987 0 6 1000 2 6000 40 800 1988 0 0 5000 250 4100 0 0 1989 0 0 12000 10 3000 0 0 1990 0 0 5000 0 2000 0 0 1991 0 0 3000 500 4000 0 0 1992 0 0 2400 0 300 0 0 1993 8 0 2500 0 600 0 0 1994 0 0 9000 0 250 0 0 1995 0 0 1500 0 50 0 0 1996 0 0 120 0 15 0 0

year DSA DSA DSB DSB DSB DSC DSC DSD DSD DSD

MILL D soisotSOUTHJTH DRAINAGDRAINAGE E 1972 20 100 100 30 1973 60 500 300 200 1974 55 176 200 1100 1975 45 50 200 150 2400 1050 1976 50 100 200 50 150 3000 100 1977 100 400 150 30 280 250 50 1978 850 350 300 1000 2700 2200 4400 1979 600 300 250 800 200 3000 400 1980 700 620 600 700 100 3000 0 1981 600 1000 100 600 100 2000 0 1982 81 700 95 265 16 1015 0 1983 23 250 250 230 4 1250 0 1984 55 12 110 200 5 475 0 1985 400 0 300 5 0 1200 0 1986 200 0 100 25 150 6 900 0 300 1987 60 12 6 120 50 4 235 0 1000 1988 12 40 30 400 300 0 15 750 0 600 1989 60 30 6 850 50 1 0 1000 0 100 1990 3 80 40 1200 40 0 0 650 200 150 1991 3 250 50 100 0 0 0 400 15 0 1992 35 5 100 1600 0 0 0 300 0 0 1993 24 150 30 50 0 0 0 200 0 50 1994 130 350 24 75 0 0 0 575 0 0 1995 120 200 12 1000 0 0 0 200 750 0 1996 250 0 100 500 0 0 0 100 150 100 126 GREAT BASIN naturalist volume 59

APPENDappendix ix B conticontinuednuedaued year FEA FEAFIpeaA FEB FEEFEB FEB FEB FEC FEC FEC FEC FED fenFEDFED FED

MILL F EASTE ST DRAINADRAINAGE GE 1972 70 25 1500 120 1973 24 150 500 200 1974 6 30 100 500 500 300 1975 11 5 50 0 200 600 100 1976 12 loo1001130 50 0 100 0 1500 600 1977 6 0 75 0 300 0 500 75 1978 15 0 30 0 70 0 150 70 1979 25 5 150 0 220 0 60 450 50 1980 10 6 8 0 450 0 20 500 200 S200 1981 12 0 18 0 40 0 20 135 150 10 1982 3 5 36 0 220 3 120 70 110 30 1983 0 0 50 0 50 0 4 6 0 50000 1984 0 0 20 0 95 0 0 100 0 160 1985 0 0 0 0 105 0 35 100 0 45050 1986 0 0 20 0 30 100 40 100 5 m800 1987 0 0 30 0 150 250 200 3 200 10 300WO 1988 0 0 120 0 25 0 140 60 200 40 250 1989 4 0 175 0 30 120 10 10 55 25 0 24 1990 0 0 475 0 20 0 30 1 100 0 0 50 50 1991 0 0 150 0 150 0 50 0 150 0 0 200 0 1992 0 0 250 0 0 0 0 0 80 0 0 0 0 1993 0 0 50 0 0 0 0 0 0 0 0 0 0 1994 0 0 150 0 0 0 150 20 0 0 0 0 0 1995 0 0 200 0 50 0 600 0 40 0 0 0 0 1996 0 0 15 0 0 0 1000 1000 50 0 0 0 0

year FSA FSA FSB FSB FSC FSC FSD FSD FSD

MILL F SOUTH DRAINAGE 1972 50 100 30 500 1973 60 150 50 450 1974 120 220 150 1000 1975 240 40 150 600 1976 20 100 200 300 1977 350 150 100 20 2 1978 500 250 600 30 50 150 1979 250 200 400 100 2000 600 1980 250 200 600 60 20000 600 1981 20 30 100 50 21500 4000 1982 2 5 1 500 37000 1500 50 7500 1983 1 20 0 150 10000 1250 100 12650 1984 2 30 0 80 10750 1400 100 14500 1985 3 350 0 255 150 9000 1200 100 3500 1986 12 800 0 18 250 5650 1100 200 9700 1987 2 100 0 150 120 26700 7000 1200 20000 1988 1 25 0 60 20 2900 2500 1500 5000 1989 30 12 0 350 2 1000 800 200 12000 1990 4 12 50 0 40 1200 3700 1600 14000 1991 3 5 70 0 450 2400 6000 1500 8800 1992 0 0 220 0 100 4200 3000 2000 18000 1993 20 1 60 0 200 6500 1000 500 4000 1994 8 0 30 0 100 4300 3000 2000 11000 1995 0 40 175 0 0 2950 750 3000 5000 1996 1 11 60 0 0 3850 1500 2300 3500 gleatgreat basin naturalist 592 199901999 appp 127 135

biogeography AND physiological adaptations OF THE BRINE FLY GENUS EPHYDRAEPHVDRA DIPTERA ephydridae IN SALINE WATERS OF THE GREAT BASIN

david B herbstlherbstsherbst1

ABSTRACT four species of the genus ephydra are commonly found in saline waters within the hydrologic great basin E hians E gracilis E packardipackpackardsardi and E aurilesaundesauripesauaunpesripes though none of these brine flies is endemic distributions also occur outside the great basin they all inhabit distinctive habitat types and form the characteristic benthic insect fauna of inland saline water habitats the affinities of each species for different salinity levels and cherchemicalnical compositions and ephemeral to perennial habitats appealappear to form the basis for biogeographic distribution patterns within any habitat changing salinity conditions over time may impose physiological or ecological constraints and further alteraltel patterns of population productivity and the relative abundance of co inhabiting species based on the physiology of salt tolerance known for these species high salinity conditions favor E hians in alkaline water and E gracilis in chloride water at lower salinitiessalinities based on limited habitat data E aurilesauripesaunpes and E packardipackpackardsardi aiealeare often more common again showing respective preferences for alkaline and chloride chemical conditions specialized adaptations for alkaline carbonate waters are found in the larval malpighian tubule lime gland of the alkali fly E hians while high salt tolerance in E gracilis appears to be conferred by high hemolymph os molality adaptation to ephemeral and low salinity conditions may be accomplished by swift adult colonizing ability and lapidrapid larval development latesrates it is hypothesized that adaptive specializations in both physiology and life history and varied geochemistry of saline water habitats across the great basin produce the biogeographic pattern of distributions foibrifor species in this genus this perspective on the genus ephydra and possibly other biota of mineral rich great basin waters suggests that interconnections among pluvial lakes may be less relevant to aquatic biogeography than chemical profiles developing in remnant lakes and ponds with the progression ofaridofariaof and post pluvial climatic conditions

key words ephydra saline lakes great basin osmoregulation biogeography salt tolerance

ever since the studies of hubbs and miller within the great basin have been only incom- 1948 biogeographic studies of the aquatic pletely described habitat affinities are poorly fauna of the great basin have been dominated known and collection records typically have by a search for vicariancevicari ance patterns distribu- no associated physical or chemical data tions of fish species often revealed patterns in addition to geographic barriers differen- suggesting pluvial interconnections among tiation of populations may also result from dif- lake basins and provided evidence for post ferencesferences in the physical and chemical features pluvial hydrographic changes including isola- of aquatic habitats without producing geo- tion leading to extinctionsextinct ions and species differ- graphic isolation and endemism selection for entiationentiation miller 1946 smith 1978 in addition physiological adaptation may restrict species to fish other obligate aquatic species eg distributions to certain habitat types great spring snails leeches molluscsmollusksmolluscs have received basin aquatic habitats often include waters an inordinate amount of attention because of with varied chemistry derived from high min- the potential for endemic distributions arising eral content the closed basin drainages that out of the vicariancevicari ance events of pluvial lake define the great basin collect and evaporate drying and recession and the example set by water in saline lakes ponds and wetlands hubbs and miller taylor 1960 hershler 1989 many springs geothermal and otherwise con- hovingh 1995 insects with poor dispersal tain high concentrations of dissolved soluteskolutes ability have also been studied in some depth and trace minerals streams in lower eleva- in the great basin eg naucoridae polhemus tions of the basin and range also typically have and polhemus 1994 but biogeographic pat- relatively high conductivity and alkalinity terns for most other aquatic invertebrates physical conditions also may be quite varied

ienalena nevada aquatic research laboratory universityumveisity of california route 1 box 198 mammoth lakes CA 93546 and marine science institute university of california santa barbara CA 93106

127 128 GREAT BASIN naturalist volume 59

in terms of thelmalthermal regime hot springs and that some ephydra had particular chemical ephemeralephemeldimeidl and episodic filling of playasolayas and preferences the objective of this paper is to intermittent streams gradients of chemical present evidence for the hypothesis that affini- and physical nationvanationvenationvariationva form a habitat template ties of each species for different salinity levels that sets the stage for adaptive syndromes and chemical compositions and ephemeral to determining the distribution and abundance perennial habitats form the basis for biogeo- of species gensusensu southwood 1977 graphic distribution patterns geochemical evolution of major soluteskolutes in closed basin saline lakes ege g eugster and METHODS jones 1979 may be a useful model for following progression and pattpatternseins in the biological contrasts of physiological specializations evolution and distribution of saline water biota for habitat chemistry are presented here as thithl oughoutthroughout the great basin water chemistry is evidence of adaptations that determine bio- determined primarily by the following factors geographic patterns data for comparisons of drainage basin lithology of inflowing osmoosmoregulatoryregulatory physiology and salt tolerance waters especially igneous vs sedimen- in ephydra were derived from published tary research nemenz 1960 herbst et al 1988 mineral precipitation pathways depend- calculations of relative costs of osmoregula- ing on the initial ratio of bicarbonate to tion were determined by applying the known gradients of calcium and magnesium and resulting osmotic to calculations the rela- energy required foifor work of in lake brines with differing contents of tive the active transport data on the range of known devel- the major cationscanions naca2mg2na ca Mgmg4242 and 0opment in ephydra were also summa- major anions cl s04 2603 2 high bi- times in 2c03 rized from published sources ping 1921 for E carbonate generally evolves into alka- packardipackardspackardi reported as E subosubopacapaca collins line brines and low bicarbonate into 1980a for E gracilisgraciks E chloride sulfate brines and low salinity reported as cinerea and herbst 1986 for E hians distribution conditions may retain similar content of data were derived from wirth 1971 and fur- bicarbonate and caegcamg without forming ther supplemented by collection records and precipitates habitat chemistry data of the author fractionation processes which enrich highly soluble such as conservative ions RESULTS na and cl and deplete others through carbonateeaicaieal bonate mineral formation release of ecological and physiological limitations carbon dioxide gas and biogenicdiogenicbio genic reduc- under changing salinity conditions in salt lakes of example tion sulfate for appear to result in varied distribution and pro- progressive enrichment from carbonate to ductivity of ephydra sppapp distribution maps sulfato sulratochlondesulfatochloridechloride to chloride waters appears to figs 1 2 and limited habitat data suggest occur with the concentration of inflowinginflowinginblowing high salinity conditions usually 25 glg L 1 waters ovelover time and distance along intercon- favor E hians in alkaline water and E gracilis nected evaporation basins of varied volume in chloride water in low salinity 25 glg L 1 hutchinson 1957 spatial and temporal com- and ephemeral waters E aurilesaundesauripesaunpes and E ponents of variation in water chemistry form a packardipackpackardsardi are often more common again show- habitat template that may be the foundation ing respective preferences for alkaline and foifolfor shaping the distribution of saline water chloride chemical conditions the general organisms prevalence of E gracilisgracihsgracios in the eastern and E four species of the genus ephydra are hians in the western great basin correspond commonly found in saline waters within the to the trend in lake geochemistry being derived hydrologic great basin E hians E gracigraciasgracihsgracifishsfishis from eastern sedimentary vs western igneous E packardipackardspackardi and E aurilesauripesauaunpesripes though none of lithology in the generation of chemical profiles these brine flies is endemic distributions also jones 1966 fiero 1986 habitats in which occur outside the great basin they inhabit I1 have collected E aurilesaundesauripesaunpes and E packardipackardspackardi distinctive habitat types and form the charac- are typically small shallow seep and spring teristicte benthic insect fauna of inland saline outflows onto saline playasolayas whereas E hians water habitats wirth 1971 first suggested and E gracigraciasgracihsgracifisfishis habitats are often large deep 199911999 biobiogeographyBlO GEOGRAPHY OF EPHYDPAEPHYDRA 129 perennial lakes or ponds within the owens tiai difference As a first approximation the death valley system of the southwest great osmotic concentration gradient can be used to basin E aurilesauripes is common in alkaline seeps estimate energetic costs assuming cuticular around owens lake while at the death valley permeability P and surface area A and elec- terminus of these onceonee interconnected basins trical gradient E are similar for both species E packardipackpackardsardi dominates the chloride waters of and chemical conditions and all other units small seeps are constants the relative cost becomes pro- comparisons of osmoregulation in E gra- portional to the natural log of the osmotic gracilis and hians fig 3 show that E gracilis dient chclCCchal fig 4 the cost functions shown regulates hemolymph molalityososmolality at an unusu- indicate that E gracilis may inhabit more saline ally high level for insects 700 1000 cosmmosm waters because high hemolymph molalityosmolalityos nemenz 1960 and about 3 times that found reduces the gradient and relative cost is only in E hians herbst et al 1988 the energetic about half that for E hians at an equivalent cost of osmoregulation against this gradient external salinity chloride waters tend to be may be calculated as shown by potts and parry more hyperostotichyperhyperosmoticosmotic than carbonate because 1964 the osmotic concentration is higher for an equivalent TDS glg L 1 concentration chfi while E gracilis possesses physiological calories laii PA RT inlillii ln soluteskolutesutes 7c afenfe adaptation for life in high salinity chloride water E hians shows much greater tolerance where P and A refer to integumentary perme- for life in carbonate waters alkaline soda ability and area R is the gas constant T is lakes than in chloride waters LC 50 toxicity temperature chC and clC are the high and low values occur at lower concentrations in chlo- concentrations of the concentration gradient ride water than carbonate water figs 5aaa ab5b external or internal n is ion valence F is E hians is adapted to alkaline carbonate lakes faraday constant and E is the electrical poten by virtue of the lime gland herbst and bradley

eu 1 13

c313 U

tat3

4pap

Ephydr g M ephydi p avdikvdi

Q ephydra hians U ephydra auricaauripa

fig 1 distribution of ephydra gracigracilisfishis and ephydra fig 2 distribution of ephydra packardipackpackardsardi and ephydra hians within the hydrographic great basin based on aurilesauripesaunpes within the hydrographic cleatcreatgleatgreat basin based on wirth 1971 and supplemental collections wirth 1971 and supplemental collections 130 GREAT BASIN naturalist volume 59

1000 r iso osmotic line 900

& 800

700 ephydra gracigracilisfisbisbbs 0 M 600

500

400 4 0 300 4 ephydra hians 200 0 100

0 i i i i i i i I1 0 500 1000 1500 2000 2500 3000 3500 4000 4500 5000

osmotic concentration of lake water cosmmosm

fig 3 osmotic regulation in ephydra hians from herbst et al 1988 and ephydra gracilis from nemenz 1960 iso osmotic line indicates where internal equals external concentrations

0 0E 25 E ephydra hians 0 2 0

ephydra gradfisgraddis 0 E

0 05

0 i i 0 500 1000 1500 2000 2500 3000 3500 4000 4500 5000 osmotic concentration of lake water cosmmosm

fig 4 relative cost of osmoregulation in ephydra hians and ephydra gracilis based on osmotic gradient cost is mini-mini mum where hemolymph and lake water are iso osmotic

1989 the lime gland is a modification of one resting stages or diapause colonizing ability pair of the malpighian tubules in this species and rapid development williams 1987 A wherein calcium is concentrated and bicar comparative summary of ephydrid develop- bonatecarbonatebonate carbonate are removed from the blood ment rates in ephemeral vs perennial habitats by forming a calcium carbonate precipitate is given in table 1 E packardipackpackardsardi has a short within the tubules lime the intolerance development time comparable to scatellascafellaScatella shown by E hians for chloride waters may picea from ephemeral ponds this contrasts indicate reduced capacity for regulating this with the longer and more flexible develop- anion though this species can survive in lower ment times seen in E gracilis and E hians salinitiessalinities of chloride water as suggested by the which may extend larval growth over long convergence of toxicity data for longer expo- periods when exposed to the stress of reduced sure times and lower salinitiessalini ties fig 5 food availability or increased salinity species completion of a life cycle in ephemeral of temporary waters without resting stages aquatic habitats involves such adaptations as do not have this option and must develop 199911999 biogeography OF EPHYDRA 131

TABLE 1 duration of larval development in selected ephydridae range of larval development time for instars 1 3 days habitat type reference ephydra packardipackpackardsardi 11 13 ephemeral saline pools ping 1921 ephydra gracilis 16 30 saline lakes collins 1980a 100 ephydra hians 15 58 saline lakes herbstheibst19861986 120 Scatellascafella picea 4 11 ephemeral ponds connell and scheiringScheinng 1982

under food limitation or salinity stiesstiegstress range for populationpopulations from mono and abertahert likeslakes at 50 glg L 1 salinity and eeessexcessess food

rapidly to reach maturity and disperse to new 300 habitat E packpackardipackardsardi exhibits this type of life 250 history in addition E packardipackardspackardi exhibits best cacarbonate wwater growth at 40 50 1 and cannot survive gl 200 above 90 glg L 1 ping 1921 well below the limits for E gracigracilisfishis or E hians 150

chloride water discussion loo100

50 f based on the physiology and expense of salt tolerance high salinity conditions favor E 0 hians alkaline E gracilisfishisbis chlo- 24 48 72 in water and graci in exposure time hours ride water at lower salinitiessalinities based on limited habitat data E aurilesauripes and E packardipackpackardsardi are often more common again showing respective preferences for alkaline and chlo- 7000 a ride chemical conditions specialized adapta- 6500 tions for alkaline carbonate waters are found in the larval malpighian tubule lime gland of 1p 600000 0 alatefhateulateftatecarbonate water the alkali fly E hians while high salt tolerance E 5500 in E gracilis appears to be conferred by high 7 y hemolymph osmolality adaptation to ephe- 5000 meral and low salinity conditions may be 4500 chloride water achieved by rapid larval growth rates as seen 4000 i i in E packardipackardspackardi biogeographic patterns of asso- 24 48 72 ciation with chemical and regional differences exposure timerimenimemime hours in habitat types correspond with differing abil- ities of these ephydra species fig 5 salt tolerance in third instar larvae of ephydra while water chemistry may broadly define hians exposed to saline waters where either carbonate or chloride are the dominant anions from herbst et al the distribution of ephydra sppapp among differ- c9881988 LC 50 values are lethal concentrations at which ent habitat types salinity changes within a half the exposed population dies as determined in mortal- habitat appear to control population produc- ity bioassays A expressed as total dissolved solids exter- 1 tion of species and the relative abundance of nal salinity glg lri B expressed as external osmotic con- centrationcentration cosmmosm milliosmolalmilliosmolal existingcoexistingco species the intermediate salinity hypothesis herbst 1988 proposes that abundance of salt tolerant organisms is limited by in fluctuating salt lakes increased abundance physiological stress at high salinitiessalinities and by of E hians at mono lake california and E ecological factors such as predation and com- gracilis at great salt lake utah occurred petition in more diverse communities at low during periods of salinity dilution winget et salinitiessalinities fig 6 field observations have pro- al 1972 herbst 1988 during a period of vided evidence of changing population dynamics dilution of already low salinity at abert lake 132 GREAT BASIN naturalist volume 59

111zik 1 salinity hypothesis saline lake productivity herbst 1988

A A

F limitations due to biotic limitations due to interactions in a diverse physiological stress community at low salinity of increasing salinityZ W0 competition predation 0

SALINITY

fig 6 intermediateintel mediate salinity hypothesis herbst 1988

Oioregonexonegon E hians abundance was reduced in these data also might be applied to under- the piepresenceseneesence of increasing numbers of other standing other zoogeographic patterns in the benthic invertebrates herbst 1988 prelimi- great basin brine flies play a central trophic nary results of mesocosmmeso cosm salinity gradient role in saline lakes as consumers of benthic experiments also support the intermediate algae in turn providing one of the principal salinity hypothesis herbst in preparation food sources used by migratory and breeding dilution of the chloride nchrich waters of the shorebirdsshorebirds and waterfowl jehl 1994 rubega creatgleatgreat salt lake has been followed by colo- and inouye 1994 the historical development nizanizationtion and inciinclincreasedeased abundance of E hians ofofalianofavianavianavlan migratory routes and breeding colonies where otherwise only E gracilis was present along the pacific flyway within the great basin welkerweiker and havertz 1973 collins 1980b during the holocene may therefore be linked this last example suggests coexistence can to salinity related changes in the population occur within transitional regions of the physio- production of ephydra in different saline lake logical tolerance zones of ephydra species basins assuming that optimum production for saline water habitats appear to be parti- E hians is in the 25 100 gl 1 range and for tioned among the 4 species of great basin E gracilis in the 100 200 gl 1 range proba- ephydra along gradients of chemical composi- ble historical locations and changes of feeding tion salinity and stability fig 7 this habitat grounds may be reconstructed from paleo template model gensusensu southwood 1977 pre- salinity records alone dicts expected species distributionsdistl ibutions and poten- physiological and life history traits related tial changes in range and coexistence in space to the selection regime of saline water habitats and time under varied environmental condi- could provide distinctive characters for cladis tions As a test of this conceptual model fur- ties analysis and permit further investigation ther seaichsealchleresearch on ephydra sppapp should include of the phylogenetic relationships in the genus i elatingrelating zoogeographic distributions to ionic ephydra and other ephydrinaeephydridae mathis 1979a tolerance and habitat chemistry using triangu- mathis separates haloscatellaHaloscatella from other sub lar anionanlonamon diagrams and comparisons of colo- genera of lamproscatellaLamproscatella partly on the basis of nizing ability life history traits and population saline water habitat preferences and it would abundance along environmental gradients for be useful to examine further whether chemical E auaurilesauripesaunpesripes for which little data are available tolerances among the species of the subgenus the conceptual model predicts that this species correspond to the dogramclacladogram presented mathis has sholtshort development time rapid coloniza- 1979b the 5 nearctic species of lamprosca tion of new habitats lower salt tolerance than tella haloscatellaHaloscatella sppapp can be found within E hians greatergi eater survival in carbonate than the great basin and mathis suggested that chloride chemistry and poolpooipoor competitive abil- this is the zoogeographic origin of this group ity in coexistence with E hians isolation through physiological adaptation to 1999 biobiogeographyBlOGEOGRAPHY OF EPHYDRA 133

high perennial lakes E gracifishisbis E hians

regions of overlap SALINITY potential existencecoexistenceco HABITAT STABILITY

E packardipackpackardsardi E aurilesauripes ephemeral low pools seeps

freshwater HABITATS

CHLORIDE CARBONATE

fig 7 saline water habitat template foiformeiemi the distribution ofgieatof creatgreat basin ephydra each coinercolnercorner of environmental template inhabited primarily by a single species

varied water chemistry may in fact be a more alkaline water but can live in moderate salini likely explanation for speciation and distribution ties of sulfate water these distributions suggest in this group than through repeated vicariancevicariadcevicarianeeance adaptations to different chemical conditions through glaciation events as proposed by allow habitat partitioning fossil preservation of mathis ostraostracodescodes permits their use as paleopaleosalmitypaleosalinitysalinity other species in the genus ephydra exhibit indicators othelother taxa with probable specific thermal adaptation E geodeni inhabits hot chemical habitat affinities and potential for springs in freshwater to moderate salinitiessalinities biogeographic interpretation through geochem- barnby 1987 E thermophilathermophilethermophila and E bruesibrueni istry include diatoms blinn 1994 corixconxidscorixidsids are endeendemicsmics of acidic and alkaline thermal scudder 1976 and branchiopod crustacea springs respectively in yellowstone national bowenetalbowen etalet al 1985 resh and sorg 1983 have park collins 1977 physiological adaptation shown that lithium tolerance in the shore bug and habitat partitioning along thermal con- saldulasaldala usingennausingerinausingerinagenna permits survival in certain ductivity and ph gradients may contribute to thermal springs and can be used to predict the origins and zoogeographic distribution of local habitat distribution these species the remnants of pluvial great basin lakes other ephydridae found in athalassic saline are primarily saline bodies of water only a water habitats of the world may also exhibit few perennial saline lakes remain including biogeographic associations with geochemistry mono lake abert lake great salt lake and physiological adaptation to varied habitat geo- pyramid and walker lakes and many ephe- chemistry may exist in homologous ephydrid meral ponds wetlands and spring seeps are fauna in isolated desert regions such as the also disappearing habitat loss has resulted dimecoeniaDimecoenia sppapp of the south american alti- primarily from stream diversion and spring plano ephydrellaEphy drella sppapp of the australian inte- development perennial lakes and ponds and rior and ephydra sppapp of the old world other habitat refugia arearc threatened and must wirth 1975 become part of a program of aquatic systems use of biological proxies as indicators of protection in the great basin to ensure the water chemistry of closed basin lakes is well mosaic of habitats is available that has permit- illustrated by the distribution of certain ostra- ted the diversification of life such as is repre- code taxa in saline waters forester 1986 sented in the ephydridae species in the genus limnocythere appear to in conclusion the following general syn- have varied anionic preferences with L stapstaplinistaplinslini dromes of biogeographic patterns in the great found in chloride and sulfate waters L sap basin are presented to reflect the importance paensis in alkaline waters and L ceriotuberosaceriotuberosa of both vicanancevicariancevicavieavicarianceaneenancenanee and geochemical variation in waters of mixed chemistry another species 0 obligate aquatic species or those with candonacacdona raisonirawsonirawsoni is found only in low salinity poor dispersal ability having restricted 134 GREAT BASIN naturalist volume 59

or endemic distributions habitat isola- meadows drainages death valley system califor tion through vicariancevicariance nia nevada proceedings of the biological society of washington 102176102 176 248 obligate aquatic species with widespread HOVINGH P 1995 zoogeographical studies of rivers in the distributions wide tolerance great basin USA amphibians molluscsmollusksmoll uscs and leeches vagile species with widespread distribu- bulletin of the north american Benthbenthologicalological soci- tions and wide habitat tolerance ety 1219212 192 HUBBS C AND R R MILLER 1948 correlation between vagile with distribu- CLL RR species restricted fish distribution and hydrographic history in the tions due to specialized adaptations for desert basins of western united states pages 17 166 certain chemical or physical habitat in the creatgreat basin with emphasis on glacial and conditions eg ephydra others postglacial times bulletin of the university of utah 38 biological series 10 hutchinson GEG E 1957 A treatise on limnology volume literature CITED 1 part 2 chemistry of lakes john wiley and sons new york 496 appp BARNBY MA 1987 osmotic and ionic regulation of two JEHL JRJ R JR 1994 changes in saline and alkaline lake brinebi me fly species diptera ephydridae from a saline avifaunalavifaunas in western north america in the past 150 hot spring physiological zoology 6032760 327 338 years studies in avian biology 1525815 258 272 BLINN DW 1994 diatom community structure along JONES BFB F 1966 geochemical evolution of closed basin physicochemical gradients in saline lakes ecology water in the western great basin pages 181 200 in 74124674 1246 1263 JLL rau editor second symposium on salt north- BOWEN STS T EAE A FOGARINO KNK N FITCHNER GLG L DANA ern ohio geological society VHSVH S CHOW MRM R buoncristianibuoncbistianiBUON CRISTIANI AND JRJ R CARL MATHIS WN 1979a ephydnnaeephydrinaeephydridae diptera ephydridae 1985 ecological isolation in artemia population dif- a new perspective pages 47 60 in DLD L deonier ferencesferences in tolerance of anionanlon concentrations jour- editor first symposium on the systematics and ecol- nal of crustacean biology 5106slog5 106log 129 ogy of ephydridae diptera north american ben COLLINS NCN C 1977 mechanisms determining the rela- thological society tive abundance of brine flies diptera ephydridae 1979b studies of ephydnnaeephydrinaeephydridae diptera ephydri- in yellowstone thermal spring effluents canadian dae II11 phylogeny classification and zoogeography entomologist 190415igo190 415 422 of nearctic lamproscatellaLampros catella hendel smithsonianSmithsoman 1980a developmental responses to food limitation contributions to zoology 2951295 1 41 as indicators of environmentalenanenvn onmental conditions for ephy- MILLER RRR R 1946 correlation between fish distribution dra cinerea jones diptera ecology 6165061 650 661 and pleistocene hydrography in eastern california 1980b population ecology of ephydra cinerea and southwestern nevada with a map of the pleis- jones the only benthic metazoan of the great salt tocene waters journal of geology 544354 43 53 lake USA hydrobiologia 689968 99 112 NNEMENZE M E nzanz H 1960 on the osmotic regulation of the larvae CONNELL TD AND JXJ F SCHEIRING 1982 demography of of ephydra cinerea journal of insect physiology 4 the shore fly Scatella picea walker diptera ephy- 38 44 drdridaeidae environmentalenanenvn onmental entomology 1161111 gli611 617 PING C 1921 the biology of ephydra subopacasubopaca loew EUGSTER HPH P AND BFB F JONES 1979 behavior of major cornell university agriculture experiment station soluteskolutes duangdunngduring closed basin brine evolution ameri- memoirs 4956149 561616561 616gig can journal of science 279609279 609gog 631 POLHEMUS JTJ T AND DAD A POLHEMUS 1994 A new species fleFIEpieFIEROro B 1986 geology of the great basin university of of ambrysusAmbryofambrysussussms stal from ash meadows nevada het- nevada press reno 197 appp eroptera naucoridaenaucondae journal of the new york FORESTER RMR M 1986 determination of the dissolved entomological society 102261102 261 265 anionanlonamon composition of ancient lakes from fossil ostra POTTS WTW AND G PARRY 1964 osmotic and ionic regu- codes geology 1479614 796 798 lation in animals pergamon press oxford 423 appp HERBST D B 1986 comparative studies of the population RESH VH AND KLK L SORG 1983 distribution of the wilbur ecology and life history patterns of an alkaline salt springs shore bug hemiptera saldidaeSaldsalpidaeidae predicting lake insect ephydra hydropyrusHydropyrus hians diptera occurrence using water chemistry parameters envi- ephydridae unpublished doctoral disseitationdissertation ronronmentalmental entomology 12162812 162816351628 1635 oregon state university corvallis RUBEGA M AND C INOUYE 1994 prey switching in red- 1988 comparative population ecology of ephy- necked phalaropesPhalaropes Phalaphalaropusropus lobatus feeding dra hians say diptera ephydridae at mono lake limitations the functional responselesresponse and water man- california and abert lake oregon hydrobiologia agement at mono lake california USA biological 158145158 145 166 conservation 7020570 205 210 HERBSTHLRBST DBD B AND TJ BRADLEY 1989 A malpighian SCUDDER GGEG G E 1976 water boatmen of saline waters tubule lime gland in an insect inhabiting alkaline salt hemiptera corixidae pages 263 289 in L cheng lakes journal of experimental biology 14563145 63 78 editor marine insects north holland publishing HERBSTHCRBST DBD B FPEP gonteCONTECONTC AND VJ BROOKES 1988 osmo- company regulation in an alkaline salt lake insect ephydra SMITH GRG R 1978 biogeography of intermountain fishes hydropyrusHydropyrus hians say diptera ephydridae in great basin naturalist memoirs 217291717 42 relation to water chemistry journal of insect physi- SOUTHWOOD TRETR E 1977 habitat the templet for ecolog- ology 3490334 903 909 ical strategies journal ofanimalofanimal ecology 4633746 337365337 365 HERSHLER R 1989 Springspnngsnailsspringsnailssnails gastropoda hydrobi TAYLOR DWD W 1960 distribution of the freshwater clam idae of owens and amargosaamargoso river exclusive of ash pisidium ultramontanum a zoogeographic inquiry 199911999 biobiogeographyBlO GEOGRAPHY OF EPHYDRA 135

american journal of science 258a325258a 325 334 WIRTH WW 1971 the brine flies of the genus ephydra WELKER MCM C AND DSD S HAVERTZ 1973 A study of species in north america diptera ephydridaeephydndae annals of diversity of the genus ephydra on the great salt lake the entomological society of americaofamerica 6435764 357 377 utah academy proceedings 507850 78 80 1975 A revision of brine flies of the genus ephy- WILLIAMS DDD D 1987 the ecology of temporary waters dra of the old world diptera ephydridae ento timber press portland OR 205 appp mologicamologica scandinavicaScandinavica 611gli6 11 44 WINGET RN DM REES AND GC COLLETTCOLLETE 1972 insect problems associated with water resource received 18 march 1998 development of great salt lake valley pages accepted 15 june 1998 156 166 in the great salt lake and utah s water resources utah water research lab logan gigreateat basin naturalist 592 199901999 appp 136 143

STREAM temperatures AND THE elevational distribution OF REDBAND TROUT IN southwestern IDAHO

bruce W zoeiZoelZoellicklickl1

ABSIKACABSTRACT 1 duringdm ing july to septemberSeptembeiber 1994 1996 I1 examined water temperatures at the lower end of the elevational distribution of redband trout oncorhynchus mykiss gairdnergairdnengairdgairdnerinerinerlnen in 4 streams in the owaheeowyhee mountains in southwestern idaho maximum water temperatures in castle shoofly little jacks and big jacks creeks during low flows during a di oughtdrought in 1994 ranged from 267c26 7cac to 290c29 0cac water temperatures fluctuated 959.59 5 11cilc during the 24 h period maximum tempeltemperaturesatul es weiwelwere e abseiobservedobsei ved stream flows at the lower end of big jacks and little jacks creeks in 1994 were 00030 003 m3rn3rna s 1 and subsided underground 50 130 m downstream of pools inhabited by trout trout were distributed to lowerlowel elevations where drainage basin aleaarea was larger inm 22of3yrp0of 3 yr P 0030.0303 lower elevational limits of redband trout distributiondistl ibution in big jacks little jacks castle and shoofly creeks were 920 934 972 and 1090 m above sea level respectivelyi min 1994 with higher stream flows inm 1995 96 trout were found 3 6 km faitherfartherhartherbarther downstream in castle big jacks and shoofly creeks at elevations of 860 891 and 998 m respectively and tolerated maximum temperatures of 25525.525 5 290c29 0cac trout weiewelewere not distributed farther down little jacks creek because of poor channel conditions maximum daily watelwater temperaturestempel atuiesaluies of 290c29 0cac may have limited trout distribution in big jacks creek as flows and suitable channel conditions but higher temperaturestempel atuiesaluies continued 5 km downstream of the lowest pool inhabited by trout in 1995 96

kenkeyku words redbandredhand trout oncorhynchus mykiss gaiidnengairdgairdnerinerinerl desert streams water temperature elevation southwest idaho distribution

redband trout oncorhynchus mykiss gaird more streams or extend their downstream dis- neri in desert basins of western north amer- tributiontribution in occupied streams studies of the icaleaiea are thought to have evolved adaptations to elevational distribution of redband trout in rela- live in harsh environments characterized by tion to water temperatures and stream flows are extremes in water temperature and flow needed to determine if or how these parame- behnke 1992 however little is known about ters limit distribution stream flows and conse- the environmental extremes redband trout tol- quently trout distribution may also be influenced erate in desert streams redband trout were by watershed characteristics such as drainage observed feeding at water temperatures of basin area and elevation of source springs 283c in chino creek a tributary to the the objectives of this study were 1 to owaheeowyhee river in northern nevada behnke determine the lower elevational distribution of 1992 in contrast rainbow trout oncorhynchusoncorhynchtis redband trout in 4 drainages typical of streams mykiss irideus are typically stressed by water on the northeast slopes of the owaheeowyhee moun- temperatures 22c behnke 1992 redband tains that are tributariestributaries to the snake river trout inhabit desert streams in the snake river 2 to determine the maximum water tempera- basin in southwestern idaho many of these tures trout were tolerating at the lower end of streams become intermittent at lower eleva- their distribution in these streams and 3 to tions and flows fluctuate greatly year to year examine whether the size of the drainage basin depending on winter snowsnowpackspacks affected the lower elevation to which trout knowledge of the maximum temperatures were distributed in a stream additionally I1 redband trout tolerate would assist fisheries compared the distribution of redband trout managers in determining potential distribution and maximum water temperatures at the lower of redband trout in streams in the snake river limit of their distribution between years of low basin in southwestern idaho with watershed stream flows in 1994 and average flow condi- lestiestrestorationoration redband trout may be able to inhabit tions in 1995 and 1996

luUSS burbuibinuuieflandoflandof lcincl managmanagementet lower slakesnake riverrivel district3948District 3948 development ave boise ID 83705

136 199919991 STREAM temperatures AND REDBAND TROUT 137

STUDY AREA in castle creek are primarily gravels and finer sized particles as are the lower portions I1 conducted the study on big jacks little of shoofly creek streamside vegetation is jacks shoofly and castle creeks which flow dominated by willows salix lasiolepsis S northward from the owaheeowyhee mountains to the lasiandralashandralasiandra S exigiaexigua and S lutealuted in areas of snake river near the town of bruneau in historical heavy livestock grazing and where southwestern idaho fig 1 redband trout flows become intermittent streamside vegeta- occupy the upper reaches of these streams all tion is composed of mesic forbs such as gold- 4 of which are usually intermittent at the enrod solidago sp and grasses lower end of the drainages and are moderately little jacks creek is a tributary to big jacks confined by side valley slopes with gradients creek but it rarely flows aboveground past a of 151.5lsis 4 B channel types rosgen 1994 highly degraded braided stream channel D table 1 big jacks and little jacks creeks and channel type rosgen 1994 starting 696.9gg km the upper portion of shoofly creek flow upstream of its confluence with big jacks through 30 to 210 m deep canyons with nar- creek big jacks creek flows to stream km row floodfloodplainsplains and stream substrates domi- 20320.3 site of a US geological survey gaging nated by cobble sized rocks stream substrates station 60 of the time primarily during the

0 bruneau

watershed boundary

co perennial 0 stream intermittent stream avdoaveo

P A

KIUIE f 0 10 km area map

fig 1 location of castle shoofly little jacks and big jacks creeks in southwestern idaho their watershed bound- ariesarles lower distributional limits of redband trout in the streams in 1994 square symbols 1995 circles and 1996 trian- gles and area map inset 138 GREAT BASIN naturalist volume 59

TABLETABLL 1 charaeCharaccharacteristicscharacteicharacterteitel isticsristics of 4 drainages in which redband trout were studied in southwestern idaho 1994 1996 drainage geologic elevation m stream basin area parent stream orderordel km2 material headwaters Confluence a1

big jacks 3 633 rhyolite lava 1670 748 little jacks 2 260 rhyolite lava 1673 748 shoofly 2 55 rhyoliterhyrbyolite lava 1639 724 castle 3 790 granitgranitelavaelava 1774 710

confluence of drainageclicil linage with snake riverhivel big jacks and little jacks join to formhormborm jacks creek

months of january through june kjelstrom et trout density was estimated in july by elec al 1995 and occasionally flows aboveground fishingtrotrofishing representative sites within 3 km of to the snake river in early spring and during the lower limit of trout distribution two or 3 storm events in 1992 big jacks creek did not electrofishingelectrofishing passes were made and popula- flow down to the gage site shoofly creek also tion sizes estimated for 60 to 80 km long seg- rarely flows to the snake river flows in cas- ments using the zippin capture removal model tle big jacks and shoofly creeks are diverted zippin 1958 for irrigation at the lower end of the drainages I1 placed a thermograph in or near the far- where the streams enter broader valleys with thest downstream pool where redband trout associated low gradient unconfined channels were encountered pools were shallow enough C channel type rosgen 1994 that flows thoroughly mixed resulting in no thermal stratification as determined from mea- METHODS suresurementsments with a hand held thermometer Thermothermographsthermographygraphs were placed in the streams in I1 visually observed redband trout at the late june 1994 1996 and monitored water lower end of each drainage in late june temperatures through september water tem- 1994 1996 to determine their approximate peraperaturestures were recorded every 2 h or the daily elevational distribution I1 then used a smith maximum minimum and average tempera- root model 12 A backpack electrofisherelectro fisher to tures were recorded temperature readings of collect fish to determine the lower limit of dis- thermographsthermothermographygraphs were checked against hand held tributiontribution of redband trout in each stream pools thermometers when they were set in the stream that provided relatively high quality habitat and again when they were retrieved depths of 020.2ob0 2 030.30 3 m with some cover for red- in 1995 maximum registering thermome- band trout were electrofishedelectrofished to determine if ters were placed at the lower end of the distri- trout were present I1 sampled downstream to bution of trout in little and big jacks creeks where flows subsided underground or to 200 Thermothermographsthermographygraphs were placed in these creeks in m downstream of the last pool where trout 1995 prior to electrofishingelectrofishing and the thermo were encountered by electroelectrofishmgelectrofishingfishing excep- graphs were 50 2700 m upstream of the lower tions were shoofly creek which was sampled end of the trout distributions therefore ther- only as far downstream as a private land mometersmometers were used in addition to thermo boundary in 1994 and similarly castle creek graphs to measure maximum temperatures in 1996 trout may have been distributed far- I1 measured stream flows with a pygmy flow ther down castle creek in 1996 and probably meter using standard discharge measurement were distributed farther down shoofly creek methods US geological survey 1977 dis- in 1994 charges were measured in late summer I1 visually checked the presence of trout at august september to determine base flows the lower end of their distribution in little I1 also used US geological survey USGS jacks creek every 2 wk during the summer in gage data from a station at stream km 20320.3 of 1994 95 and also on this schedule in big jacks big jacks creek to determine stream flows creek in 1994 little jacks creek was observed elevations at the lower end of the trout dis- monthly in 1996 all sites were rechecked for tributiontribution on each drainage were calculated trout in september by electrofishingelectrofishing from USGS 75 minute topographic maps 1999 STREAM temperatures AND REDBAND TROUT 139

using least squares linear regression I1 exam- the last pool inhabited by trout however ined the relationship between the lower eleva- stream flows of about ooi0010.01 m3ma s 1 continued tion of trout distribution and drainage basin another 434.3 km below the last pool inhabited size the relationship between lower elevation by redband trout in castle creek trout were of occurrence and drainage basin area was present in shoofly creek to at least an eleva- examined for each year tion of 1113 m at the lower limit of public land on the stream flows in shoofly creek of RESULTS 00050.005 m3ma s 1 continued about 2 km downstream onto private land and trout were likely I1 electrofishedelectro fished 232.3 292.9 and 141.4 km of stream present a portion of this distance to an eleva- near the lower limit of trout distribution in the tion of about 1090 m 4 streams in 1994 1995 and 1996 respectively redband trout were distributed to lower trout were continuously distributed pools pro- elevations in 1995 96 when stream flows were viding some security or resting cover were about 5 times greater than in 1994 table 2 almost always occupied by 1 or more trout fig 1 big jacks creek flowed 838.3 km farther densities were low averaging 222.2 trout100trout 100 downstream in 1995 than in 1994 with trout m2ma of stream range 020.2 to 7 trout100trout 100 m2ma recolonizing about 13 27272.7 km of this dis- n 4 within 3 km of the lower limit of trout tance trout in castle creek were distributed distribution in 3 streams in which I1 examined 494.9 and 585.8 km farther downstream in 1995 density big jacks castle and shoofly creeks and 1996 respectively than in 1994 in 1996 adult trout ranged from 146 mm to 231 mm in trout in castle creek were distributed at least length young of year trout were 50 75 mm to a diversion at a private land boundary and long all fish appeared healthy may have occurred downstream of the diver- trout were always present in september at sion similarly in 1995 96 trout in shoofly the sites of their lower limit of distribution creek were distributed 2 3 kmkrakna farther down- determined from electroelectrofishingfishing in late june stream than in 1994 to an elevation of 998 m and early july trout were also always observed where stream flows were diverted into an irri- at the lower limit of their distribution when gation canal flows in big jacks creek were monitored biweekly or monthly during july to greater in 1996 but the lower limit of trout dis- september tribtributionution was unchanged from 1995 table 2 elevational distribution elevations at trout distribution in little jacks creek in the lower limit of redband trout distribution in 1995 96 differed only slightly from 1994 with 1994 were 920 1090 m table 2 flows in big their distribution ending at the upper end of a jacks creek continued at 00030.003 m3ma s 1 just highly degraded braided stream channel even 50 m farther downstream of the last pool in- with greater stream flows in 1995 table 2 habited by trout in 1994 similarly stream surface flows continued just 080.8 km into the flows in little jacks creek of 00030.003 m3ma s 1 degraded segment and did not provide pool continued only 130 m into a highly degraded habitat more than 200 m into the segment braided stream channel D channel type below surface flows in 1996 continued about 252.5 km

TABLE 2 stream flows mam3 s 1 and elevations m at lower distributional limits of redband trout in 4 southwestern idaho streams 1994 1996 year

1994 1995 1996 stream elevation flowfloweflow1 elevation flow elevation flow castle creek 972 ooi0010.01 871 008 859 c little jacks creek 934 0030.03 934 007 934 0090.09 big jacks creek 920 0oobbobabqb 891 003b0 891 00810.0810 shoofly creek 1090 0005 998 003 998 003 astreamsstrsti earnsearms gaged late summer august Septemseptemberberbei to estimate base flow measmedMeabmeasuredmeasmersmed at USU S geological survey gage station 838 3 km downstream offish distribution limit in 1994 cnoanono data 140 GREAT BASIN naturalist volume 59

into the degraded channel and trout were dis- a 1994 tritributed 150 m farther downstream 65 27 51 65 the elevation to which trout were distrib- loo100 uted was dependent on the area of the 90 drainage basin of the 1995 fi2 stream in rar2 n 80 0950.95ogs df 12 FP 0030.03 and 1996 rar2 0960.96ogg 70 df 12 FP 0020.02 but not in 1994 rar2 0410.41 df 12 P 0360.36 generally the larger 0LL 60 drainage basins provided flows that allowed z 50 occupancy of lower elevations however cas- 0LU 40 tle creek had the largest basin area table 1 of 30 aLU but in 1994 trout extended their distribution 20 to lower elevations in 2 of 3 other drainages 10 STREAM temperatures daily maximum temperatures ranged from 179c to 290c 0 during 1994 fig 2aaa I1 may have missed sam- b 1995 pling the maximum temperature in castle 35 35 35 creek due to equipment failure the thermo- loo100 graph did not operate until 5 august 1994 90

the median daily maximum temperature was C 80 245c and the 95th percentile of daily maxi- 70 for 3 in mum temperatures streams monitored U 60 from late june through august was 0 280c 50 maximum temperatures for individual water ujz streams ranged from 267c to 290c table 40 3 because trout in shoofly creek in 1994 uj 30 a were likely distributed downstream onto pri- 20 vate land they probably tolerated maximum 10 which temperatures 267c were the maxi-maxi 0 mum temperatures measured at the lower c 1996 boundary of public land water temperatures gs 64 64 64 61 fluctuated 959.5 11cilg during the 24 h period loo100 maximum temperatures were observed in 90 1994 average daily temperatures ranged from CO 80 210c to 230c table 3 temperatures remained within 3c30cac of the maximum for 4 9 h 70 fig 3 for daily maximums 27c tempera- LL 60 gg 0 tures were above 26c for 292.999 444.4 h zF 50 in 1995 daily maximum temperatures ranged W 40 1 0 from 180c to 28ooc fig ab2b I could not W 280c uj 30 relocate the thermograph in big jacks creek a the maximum temperature was measured with 20 a maximum registering thermometer the 10 median daily maximum temperature was 0 C 01

fig 2 fleFiefrequencypiequency distribution of daily maximum water 0 temperatures at lowellower distributional limits of redband 1719917 199 C DF 2323259259 C trout in 4 southwesternsouthwestein idaho streamssti earnsearms a 29 june 31 august 1994 b 29 june 2 august 1995 c 29 june 31 IN 2022920 229 OCC 262902629.026 29029.0 OCC august 1996 number of days sampled is shown above the baibalbar toifoifolfor each stream shoofly creekcleekgleek was monitored at the same site in 1994 95 575 7 km upstream of lower distributional limit in 1995 big jacks creekcleekgleek was not monitored inm 1995 due to equipment failure 1999 STREAM temperatures AND REDBAND TROUT 141

TABLE 3 water temperatures diningduringduning the 24 h period maximum temperatures weiewelewere observed at the lower distributional limits of redband trout in 4 southwestern idaho streamssti earnsearms 1994 1996 temperature ICC

1994 1995 1996 stream max minmm aagavg max minmm aagavg max minmm aagavg castle cleekcreek 270 170 225 265 185 210 270 190 225 little jacks creek 290 179 230 250 185 210 260 192 221 big jacks creekgreek 279 185 213 280980 290 190 235 shoofly creekgreek 267 155 210 222251511 155 190 255 165 201

Mmci&uieclcasuredalured with a and t i imixnnumniaxinnno registering thermometer lueannemmem ind minimum temperperaturesituics not nicanicasuredmcisuilclsured onitoredmonitoiedmonitoredMonitored atit saine5 tineline site asis 1994 andind57kmupstieimoflowcidstiilutiiioftioutml99557 1 upstreaniupstreani afloweroflower disbibution of trouttront in 1995

225c and the 95th percentile of daily maxi- discussion mum temperatures for the 3 streams monitored with thennographsthermothermographsthermographygraphs was 260c maximum water pools at the lower ends of the 4 streams temperatures for individual streams ranged were shallow and did not thermally stratify from 225c to 280c table 3 water tem- the continuous distribution of trout at the peraturesperatures fluctuated 7 8cac during the 24 h lower end of the drainages indicated thermal period maximum temperatures were observed refugia from seeps were not present addition- in 1995 little jacks and shoofly creeks tem- ally in 1994 I1 observed redband trout actively peraturesperatures were monitored at the same sites in foraging at a temperature of 262c26 2cac in big jacks 1994 95 with greater stream flows in 1995 creek I1 day before the stream reached its table 1 maximum water temperatures were highest daily maximum temperature of 279c27 9cac about 4c40cac lower for the 2 streams fig 2 these observations indicate that trout were table 2 because of poor stream channel con- tolerating the temperatures measured at the ditions wide and shallow with no pools trout lower limits of their distributions and not could not take advantage of the higher stream moving in and out of thermal irefugia flows and move farther downstream in little this is the first study to systematically doc- jacks creek in 1995 96 ument that redband trout in streams other daily maximum temperatures in 1996 ranged than those inm the owaheeowyhee river basin tolerate from 185c to 290c between 29 june and extreme temperature fluctuations maximum 31 august fig 2cac the median daily maxi- temperatures tolerated by redband trout in mum temperature was 240c and the 95th this study 29c290c were slightly higher than percentile of daily maximum temperatures for behnke 1992 observed for redband trout in all 4 streams was 280c maximum water chino creek in nevada 283c28 3cac maximum temperatures for individual streams ranged water temperatures from this study were mea- from 255c to 290c table 3 water tem- sured in streams with flowing water in con- peraperaturestures fluctuated 7 10cloc during the 24 h trast to behnke s 1992 observation made at a period maximum temperatures were observed pool remaining after stream flows stopped in 1996 the maximum temperature of 255c behnke 1992 states that tolerance of high observed in shoofly creek in 1996 was more temperatures shown by redband trout popula- representative of the maximum water temper- tions evolved through natural selection in atures experienced by trout at the lower end streams of hot and regions over thousands of of their distribution in 1995 temperatures years redband trout inhabiting tributary were monitored in shoofly creek at the public streams to the snake river in southwestern land boundary 575.7 km upstream from the lower idaho also demonstrated tolerance of low dis- end of the distribution of trout maximum solved oxygen concenticoncentrationsconcentinconcenti ationsactions iglg161.61 6 404.04 0 mg L 1 water temperatures observed at the lower dis- during periods of low stream flows vinson tributiontribution of trout in big jacks and castle and levesque 1994 creeks were similar for all 3 yr table 3 rainbow and cutthroat trout oncorhynchus channel conditions were adequate in these 2 clarki typically experience stiessstress when water drainages to allow trout to move farther down- temperatures ilseiiserise above 22c220c with gradual stream during increased flows in 1995 96 increases in temperature 1 gocgog per day loss 142 GREAT BASIN naturalist volume 59

laboratory study 30 1 1 1 1 1 times from another kaya 1978 indicated that 3 rainbow trout stocks 28 acclimated to 17c would tolerate a constant temperature of for 858.5ss ist15715.7 h with a 50 00 of26c26c W 26 mortality rate and would tolerate 29c29cc for ry only 040.4 h 24 the temperature range over which redband trout continue to feed and gain weight 22 functional feeding temperatures may be a more important adaptation than an increase in W 20 the upper lethal temperature that is tolerated behnke 1992 redband trout from a desert 18 basin catlow valley in eastern oregon had

1 growth efficiencies at 16 optimum temperatures 0 4 8 12 16 20 24 19c190c while growth rates of other oncorhyn- HOUR chus mykiss stocks decrease with increasing LITTLE JACKS SHOOFLY temperatures above 16c behnke 1992 higher functional feeding temperatures would BIG JACKS CASTLE allow redband trout to maintain a competitive advantage other fish such as fig 3 typical 24 h temperature cycle at the lower dis over species red tnbutionaltributional limit of redband trout in big jacks 24 july side shiners richardsoniusRichardsonius balbalteatusteatus which 1994 castle 5 august 1994 shoofly 24 july 1994 and are common in warmer low elevation seg- little jacks 13 august 1996 creeks in southwestern ments of streams in the snake river basin idaho during years of greater stream flows redband trout were distributed farther downstream than during drought years reoccupying at about of equilibrium and death occur 3 to 6 km sections of castle shoofly and big 28 29c lee and rinne 1980 behnke 1992 jacks creeks that had been totally dewatered I1 estimates of upper lethal temperatures for or 2 yr previously however the reoccupation trout differ depending on the laboratory method of newly available sections of stream compli- used behnke 1992 studies using long dura cates determinations of whether or not maxi- tion exposures have estimated the upper lethal mum water temperatures limit distribution of temperature for rainbow trout to be 26c260c redband trout during years of greater snow bidgood and berst 1969 charlon et al 1970 packs when surface flows continue farther down hokanson et al 1977 jobling 1981 redband a drainage trout may not be present because trout inm the streams examined in this study tol- they have not yet recolonized the newly erated greater daily fluctuations in tempera- watered segment in general trout were dis- ture than used by lee and rinne 1980 to tritributed downstream to reaches with maxi- determine critical thermal maximum tempera- mum temperatures of 27027.0 290c provided tures of 5 trout species rainbow apache channel conditions allowed trout to move oncorhynchus filaegilae apache gila oncorhyn- downstream chus filaegilae filaegilae and brown trout seimoselmosalmo poor channel conditions and subsiding of trutta tolerated fluctuations from only 21c210cgic to surface flows into streamstreambedbed gravel limited 27c when daily temperatures fluctuated goc trout distribution in big jacks creek in 1994 ovelover a 24 hb period lee and rinne 1980 and little jacks creek during 1994 1996 redband trout in this study tolerated tem- stream flows at the lower end of big jacks and peraturesperatures above 26c260c for durations of up to little jacks creeks were 00030.003 m3ma s 1 red- 444.44 4 h when the upper incipient lethal tem- band trout were essentially distributed the pelaperaturepesatureperapel ture of rainbow trout 256c25 6 was ex- length of surface flows in these 2 streams in ceeded daily for 3 h under a fluctuating tem- 1994 similarly trout in shoofly creek in peratureperature regime ac4c around a daily mean of 1995 96 were distributed down to a stream 22c220c the daily mortality rate was 428 diversion which was the lower limit of avail- hokanson et al 1977 median resistance able habitat however redband trout were not 199911999 STREAM temperatures AND REDBAND TROUT 143 distributed the length of surface flows in castle acknowledgments creek in 1994 1996 in 1994 castle creek flowed at about 00160.016 m3ma s 1 for another 434.3 J nelson and D kearns assisted with field- km below the last pool inhabited by trout work I1 thank M mccoy for preparing the fig- water temperatures may have been a factor ures and reviewing the manuscript R behnke influencing the lower limit of the distribution B rieman and S sather blair provided com- of redband trout in castle creek maximum ments on earlier drafts of the manuscript water temperatures of 29c may have also limited redband trout distribution in big jacks literature CITED creek in 1996 as surface flows continued BEHNKEBEHNKL 1992 native trout of western north amer- sg5.6 RJ westein 5656 km downstream of the lower distribution ica american fisheries society monograph 6 275 of trout PP during years of greater flows and conse- BIDGOOD BYB F AND AHA H BERST 1969 lethal tempera- great quently lower stream temperatures trout in tures for lakes rainbow trout journal of the fisheries research board of canada 26 456 459 jacks able 26456 little creek were not to move far- CHARLON N B BARBIER AND L BONNET 1970 thermal ther downstream because of poor watershed resistance of rainbow trout salmo gairdgmrdnengairdnerinerinerlnefi richard- conditions A highly braided D stream chan- son to abrupt temperature variationsvanations annales dhychy nel stopped trout recolonization of the lower drobiologie 1731 73 89 HOKANSON KEEK E F CFC F KLEINER AND TW tiiorslund drainage probably because the shallow riffle 1977 effects of constant temperatures and diel tem- habitat did not provide resting or hiding cover peraperatureture fluctuations on specific growth and mortal- and also was quickly heated by solar radiation ity rates and yield of juvenileofjuvenile rainbow trout salmo this indicates that stream restoration on the gairdnergairdnengairdgairdnerinerinerlnen journal of the fisheries research board of canada 34 639 lower ends of drainages 34639 648 in southwestern idaho JOBLING M 1981 temperature tolerance and final preferpreter has the potential to increase the range and eidumendum rapidlapid methods foiforholbolhorbor the assessment of opti- numbers of redband trout populations mum growth temperatures journal of fish biology trout populations may also be reconnected 1943919 439 455 by KAYA CMC M 1978 thermal resistance of rainbow trout improving habitat conditions at the lower fromflom a permanently heated stream and of two hatch- end of drainages so that fish can move be- ery strains progressive fish culturist 4013840 138 142 tween some drainages such as big and little KJELSTROM LC MAJ STONE AND W A HARENBERG jacks creeks given current land uses and irri- 1995 statistical summaries of stream flow data for gation diversions trout populations probably selected gaging stations in idaho and adjacent states through september 1990 volume 1 gaging stations cannot be interconnected via the snake river with 10 or more yeaiyealyears s of record USU S geological suisulsur in 1995 96 during typical stream flows for vey water resources investigations report 94406994 4069 big and little jacks and castle creeks red- 534 appp LEE RMR M AND J N RINNE 1980 critical thermal max- band trout were distributed only down to JN imalma of five trout species in the southwesternsouthwestein united 860 900 in elevation the confluence of these states transactions of the american fisheriesFispishenes soci- streams with the snake river was at about 750 ety 109632109 632 635 in elevation the lower elevation of trout dis- ROSGEN DL 1994 A classification of natural rivers catena 22 169 199 tributiontribution was not significantly related to drain- 22169 US geological SURVEY 1977 national handbook of age basin area during low flows in 1994 how- recommended methods for water data acquisition ever during normal flows in 1995 96 trout USU S government printing office washington DC were distributed to lower elevations as drainage VINSON M AND S LEVESQUE 1994 redband trout re- great basin area increased probably because larger sponse to hypoxiahypoxis in a natural environment basin naturalist 5415054 iso150 155 basins provided greater stream flows and lower ZIPPIN C 1958 the removal method of population esti- stream temperatures to lower elevations mation journal ofwildlifeofwildlife management 228222 82 90

received 25 february 1998 accepted 11 august 1998 creatgleatgreat basin naturalistNatmalist 592 019991999 appp 144 150

EFFECT OF SALINITY ON SEED germination OF triglochin MARITIMA UNDER VARIOUS temperature REGIMES

M almalalmaiajmal khanl2khan12 and irwin A ungarl3Ungarungar1313

alislalielawstraciAWSTRACiRA 1 triglochin inarmarimartinarithnamaritiinatiinaithna L allowarrow glassgrass an herbaceous perennial in the family juncaginaceae is widely dis- tributedtiitiltritrl in inland and coastal salt marshes of northnoi th america triglochin mantimamaktimamaritimamarimafitima seeds from a population growinggi owing in a salt maishmarshmalsh at faust utah weiewelewere germinated at 4 temperature regimes 12 h night12 h day 5 15c 5 25c25oc 10 20c and 15 25c25q and 5 salinitiessalinities 0 100 200 300 400 and 500 mol m 3 nacl to determine optimal conditions for germination and level of salt tolerance ungerminated seeds weiewelewere returned to distilled water after 20 d to determine whether seeds could recover fromflom salinity treatments maximum germination occurred in distilled water and increasesinci eases in nacl concenticoncentrationconeonconcentincenti atlon progressively inhibited seed germinationgelgei ruination no seeds germinated at concentrations higher than 400 mol in 3 nacl A temperature regime of low night acaq5q5c and high day 25q25c temperature yielded maximum germination all othelother temperature regimes significantly inhibited seed germination relative to this optimum recovery of germination was highest at 5 25c and lowest at 5 15c recovery of seed germination when seeds weiewere transferred to distilled watelwater fromfi om salt solutions was highest at 5 25c250c 72 for seeds exposed to the 500 mol m 3 nacl pretreatment and sig- nificantlynificantly reducedi at otherothel temperature regimes the recovery germination response indicates a synergistic inhibitory interactioniteimtei actionaelion effect on germination when seeds were exposed to high salinitiessalinities at suboptimal thermopenodstbermoperiods

kenkeifkey words triglochinnighiglochin mantimamaktimamaritimamanmarlmari tima balhalbaihalophyteophite recovery seed gennigenninationgerminationnation thermoperiod utah

tnglochtntriglochintfiglochin maritinamaritimamantzmamatimarimarftima L juncagmaceaejuneaginaceaejuncaginaceae com- french population of T maritinamarimaritimatima had a pri- monly known as arrow glassgrass is a clonal peren- mary morpho physiological dormancy and nial that can form regular clumps up to 2 m that seeds of T maritinamaritimamaritima were more dormant across and 60 cm high davy and bishop 1991 than T papalustrislustris probably because of the more common in saline habitats particularly coastal resistant sclerenchyma tissue in the pericarp marshesmaimal shesashes on rocky shores in temperate subarc- of the former the 2 basic types of dormancy tic andlindtind arctic regions it also extends southward that seeds develop are due either to some to the subtropics davy and bishop 1991 morphological or biochemical characteristics triglochin dantimamantimatnaritima is distributed in inland of the diaspore that produce a primary dor- and coastal brackish and freshwater marshes mancy fruit or seed or to an environmental and bogs of north america sheltlersheatlerSheltler and skog factor that induces seeds into a secondary dor- 1978 ungar 1974 surveyed a triglochin manmatimatl mancy bewley and black 1982 binet 1961a tima community located at park county col- 1961b reported that T maritinamarimaritimatima had a sec- orado and reported that T mantimamaktimamaritimamarimafitima grew in ondary dormancy induced by darkness which almost pure stands in a wetter area with soil in nature is probably triggered by the burial of salinity ranging from 050.5os0 5 to 1 85 170 mol seeds in the soil m 3 triglochin maritinamaritimamanttmamarimarlmafitima was also found grow- germination responses of seeds of T mar ing in salt marshes at the fish springs research itimaidima populations from north america have site juab county utah in an eleochariseicoeihoEleochans not been previously investigated and one of meadow community where salinity averaged the goals of this investigation was to deter- 050.50os 5 total salts bolen 1964 mine if responses to environmental variables germination of halophytes is affected by differ from populations studied from europe temperature and soil salinity content and and related species from africa naidoo and seeds are characterized by varying types and naicker 1992 studied the effect of light tem- degrees of dormancy binet 1965 1968 ungar peraperatureture and salinity on the germination of T 1991 binet 1959 reported that seeds from a balbosabulbosa and T striatastriana populations from south

I1 department of environmentaltaital irtncld plant biology ohio uniuniversityrsityrosity athathens OH 45701297945701 2979 USA 2 permanent addaddressei department of botany university of karachiofkarachi karachi 75270 pakistan 3 corresponding author

144 199911999 germination OF trltriglochinTRICLOCHIN MARITIMA 145

africa and determined that both species have determines if a population will survive to a light requirement for germination and reproductive maturity each species has very achieve highelhigher germination at a warmer tem- specific germination requirements and its re- peratureperature regime 20 30c germination was sponse to stress varies fornhornbornfrom that of other highest in distilled water and decreased signif- species for this reason it is important to de- icantly with an increase in salinity up to 500 termine the range of tolerancetoleibolei anceanee to salinity and mol m 3 transfer of ungerminated salt temperature regime effects on germination treated seeds to distilled water stimulated ger- the effects of salinity and temperature regime miminationnation more in T striatastnatastrianasanata than in T balbosabulbosa on germination and recovery responses of T the interaction between salinity and tempera- mantimamaktimamaritimamarimafitimafima were studied to determine their in-in ture on germination has been the subject of dividual effects and any interaction between investigation khan and ungar 1984 gutter- these factors on seed germination we also de- man 1986 khan and weber 1986 khan et al terterminedmined if salinity and temperature regime 1987 badger and ungar 1989 khan 1991 interact in thentheutheir effects on recovery germination khan and rizvi 1994 because it plays a signif- of seeds initially exposed to saline conditions icant role in determining the timing of germination however no data are available con- MATERIALS AND METHODS cerning the effect of the interaction between different temperature regimes and salinity on we collected triglochin mantimamaktimamaritimamarimarltima L seeds seed germination of north american popula- duringdm ing august 1995 from a salt marsh situated tions of T dantimamaritinamaritimamantimamarimafitima these 2 environmental 30 mi south of the great salt lake at faust factors play a significant role in determining utah seeds were separated from the inflores- whether plants can successfully establish in cence and brought to ohio university where saline habitats because they interact in deter- they were stored at 4cac preliminary tests mining if seeds germinate or remain dormant indicated the seeds were viable and germina- in the seed bank ungar 1995 one of the pur- tion experiments were initiated in september poses of this investigation was to determine 1995 inm 50 x 9 mm celmangelmangeiman no 7232 tight how the germination response of T matimarimatlmaktimamantimamaritimatima fitting plastic petri dishes with 5 ml of test to temperature and salinity may affect its estab- solution each dish containing 25 seeds that lishmentlishment in salt marsh habitats were surface sterilized with the fungicide recovery germination of seeds in fresh- phygonphagon was placed in a 10 cm diameter plas- water after they were exposed to saline condi- tic fetnpetripetn dish as an added precaution against tions has been investigated ungar 1962 1978 water loss by evaporation four petri dishes barbour 1970 parham 1970 macke and ungar containing 25 seeds each weiewelewere used as repli- 1971 seneca and cooper 1971 woodell 1985 cates for each salinity and temperature treat- keiffer and ungar 1995 to determine if seeds ment seeds were considered to be germinated can remain viable after being exposed to with the emergence of the radicle hyphyperhypersalinehypersahnesahneersaline conditions but no similar data are to determine the effect of temperature on available foifolfor T mantimamaktimamaritimamarimarlmafitima seeds the ability of germination we used regimes of 5 15c seeds to germinate after exposure to hyper 5 25c 10 20c and 15 25c we used a saline conditions plays a significant role min the 24 h cycle where the higher temperature 15 establishment of halophyte populations seeds 20 or 25c coincided with the 12 h light of glycophytes cannot germinate after expo- period sylvania cool white fluorescent lamps 2 1 sure to salt stress while halophytes show a 25 kimollimol m s 400 750 nm and the lower range of responses from partial to complete temperature 5 10 or 15c coincided with germination recovery when salinity stress is the 12 h dark period seeds were germinated alleviated woodell 1985 ungar 1991 in distilled water 100 200 300 400 and 500 this study was initiated to obtain a better mol m 3 nacl solutions in each of the temper- understanding of germination requirements of ature regimes and germination was recorded seeds of a population of T maktimamantimamaritimamatimaritima from the every other day for 20 d after 20 d we trans- great salt lake region of utah initial estab- ferred ungerminated seeds from the nacl lishmentlishment of species in salt marsh habitats is treatments to distilled water and a tempera- related to germination response of seeds to ture regime ofofaof55 25c to determine the recov- salinity and temperaturetemperatuietule regime and usually ery germination which was also recorded at 146 GREAT BASIN naturalist volume 59

TABLLFABLL 1 results of a 2 way ANOVA of final percent germination of triglochin maritimamaritimemantwnamarimarltima in different salinity and tem peipelperaperaturepesatureatulturee ti eatmentseattreatmentsments sum of mean significance souiceofvaiiationsource of variation squares df square F offof F temperature 103273 3 34424 724 00001 salinity 177293 5 35459 746 00001 temperaturetempel atul exX salinity 77027 15 5135 108 00001

2 d intervals for 20 d of was rate germination 100 estimated by using a modified timson index of 102010 20 C germination velocity TI lctY gt where G is OM 152515 25 oc 5255 25 C the number of seeds germinating at 2 d inter- 80 f1 EMfa 5155 15 C vals and t is the total germination period khan and ungar 1984 the maximum value possible using this index with our data was 50 60 iei c 100020 and the higher the value the more rapid the rate of germination germination data were transformed arcsine 40 before statistical analysis and data were analyzed with a 2 way ANOVA using SPSS for windows release 616.1gi6 1 SPSS inc 1994 20

RESULTS

different temperature regimes salinity 0 loo100 200 300 400 500 3 and their interaction significantly P 0 0001000010.0001 naci mol M affected the final percent germination of T per dantimamaritinamantimamaritimamarimarltima seeds table 1 germination of T fig I1 percentpel cent germination of triglochin mantimamaktimamaritimamarimarlmafitima seeds in 0 100 200 300 400 and 500 mol m 3 nacl at temper- maktimamantimamaritimamatimarimatltima was highest in distilled water and at ature regimes ofaof5of 5 15c15oc 5 25c25oc 10 20c and 15 25c a regime with low night ac5c50c and high day 25c temperatures fig 1 maximum germination percentages were achieved in 12 d in all treatments germination of seeds decreased temperature regimes of 10 20c and 15 25c with increases in salinity few seeds germi- the rate of germination was similar in the con- nated at salt concentrations higher than 300 trols but the interaction caused by the addi- mol m 3 nacl fig 1 variation in tempera- tion of nacl to the medium adversely affected ture regime significantly affected seed germi- the germination rate at 10 20cgocgog tables 2 3 nation under both saline and nonsalinenon saline condi- after 20 d of nacl treatment seeds were tions in fact there was less than 10 germi- transferred to distilled water where there was nation at the 5 15c temperature regime in less than 25 recovery germination in the the control and all salinity treatments fig 1 10 20cgoc temperature regime at all nacl con- at other temperature regimes there was a sig- centrationscentrations fig 2 at 15 25c and 5 25c nificantnificant inhibitory interaction between tem- the final germination percentages increased to peraperatureture and salinity on final germination per- more than 50 at 500 mol m 3 nacl recov- cencentagestages table 1 fig 1 ery at the highest salinity concentration 500 different temperature regimes salinity mol m 3 was lower than that in the 400 mol and their interaction significantly P 0 0001000010.0001 m 3 nacl treatment which did not differ sig- affected the rate of germination of T maktimamantimamaritimamatimarimatltima nificnificantlyantly from the control at 5 25c fig 2 seeds as determined from the timson index of ungerminated seeds from the 5 15c temper- germination velocity tables 2 3 the rate of ature regime were transferred to 5 25c after germination calculated using a modified tim- 20 d and germination increased but germina- son index of germination velocity was lowest tion was significantly lower than for those seeds in 5 15c and highest in 5 25c table 2 at that initially germinated at 5 25c fig 2 199919991 germination OF TRItriglochinCLOCHIN MARITIMA 147

TABLE 2 index of germination velocity using a modified timson index khan and ungar 1984 to estimate rate of ger- mimination of triglochin mantimamaktimamaritimamarimarltimutima

Temtemperatureperatiirelre i egideegimeregime IC01C nacl mol m 3 10 20 15 25 5 25 5 15 0 15712315715.7 23 17817 8 343 4 24184124 1 333.33 3 070.70707070.7 100 424.2422323 12412.412 4 242 4 16416 4 080 8 0 200 270827 08 661766 17 961296 12 040304 030.3 300 1205121.2 050.5os 5012505.0so 121.2 4504454.5 040.4 oioi0101010.1oi 010.1oi 400 0 1707171.7 070.7 050.5050202 0 500 0 0 0 0

TABLE 3 results ofafofa2of a 2 way ANOVAAN OVA using data from the timson index of germination velocity to estimate rate of ger- mimination of triglochin marimaritinamaritimatima at different salinitiessalinities and temperatures sum of mean significance source of variation squares df square F off temperature 11184 3 3728 512 00001 salinity 24345 5 4869 669 00001 temperature X salinity 10169 15 678 93 00001

recovery germination percentages increased the dark parham 1970 our results indicate with an increase in salinity concentration fig that seeds of this utah population were not 3 at a temperature regime of 10 20c a dormant and that germination was inhibited maximum of 20 recovery germination was by high salinitiessalinities and low day temperatures obtained in 500 mol m 3 nacl but seeds we determined that triglochin maritinamarimaritimatima seeds treated with 400 mol m 3 nacl at 5 25c had had their highest germination percentages in 72 recovery fig 3 distilled water and a progressive decline in germination with increases in salinity similar discussion results were found in populations from europe binet 1960 1965 pigott 1969 letschertlotschertLotschert 1970 triglochin maritinamarimaritimatima germination is most our results agree with those of binet 1965 probably regulated through variation in soil who determined that germination in saline salinity and temperature regime under natural media of seeds from a french population of T conditions when soil salinity is beyond the maritinamarimaritimatima was greatly facilitated by alternating levels at which seeds can germinate seeds may temperature regimes of 5 25c which can sub- die or remain dormant in the soil seed bank stitute substantially for the light requirement seed germination can then take place at a likewise germination of the related species later time in the growing season or in another triglochin bulbosabalbosa and T striatastriana was also high- year after salt stress has been alleviated ungar est in nonsalinenon saline controls and decreased sig- 1995 bolen 1964 and ungar 1974 reported nificantlynificantly with an increase in salinity up to 500 that T maritinamaritimamaritima was found growing in com- mol m 3 naidoo and naicker 1992 higher munitiesmunities with moderate salinity 050.5 101.0io total day temperatures were more stimulating for salts 85 170 mol m 3 we determined that germination compared to lower thermoperi seeds from the utah population required low ods in all of these species of triglochin simi- soil salinity and a temperature regime with lar promotive effects of high daytime tempera- low night socacsog5c50c and high day temperatures tures on germination also were found in other 25c to promote maximum germination perennial halophytes such as cressa creticecretica binet 1959 reported that freshly collected T khan 1991 atriplex griffithiigriffitgriffithivhii khan and rizvi maritinamarimaritimatima seeds enclosed by the pericarp had 1994 salicornia pacifica var utahensis khan an innate dormancy and poor germination in and weber 1986 halopyrumHalopyrum mucromucronatumnatum 148 GREAT BASIN naturalist volume 59

loo100 90 80 102010 2020cOC 152515 2525cOC 0 70 60 50 50 S 40 3 E 30 0 molmoim01 Mm3ma 0 20 100 molmoi Mm3ma 0 3 10 u i 200 mol M pen b 3 prnsen 300 mol Mm3ma 0 arn r 3 ma 100 400 molmoim01 Mm3 500 mol mm3ma3 90 5c 25c OCop 80 525 5155 15 OC 5 25c250c 5 70 15c5 c 60 1 1 1 1 0 50bu T T T T T T T T ro W S 40 al E 30 0 20 10 I1 0

T I1 I1 I1 I1 I1 I1 I1 I1 I1 I1 I1 I1 I1 I1 I1 I1 I1 I1 1 1 20 22 24 26 28 30 32 34 36 38 40 20 22 24 26 28 30 32 34 36 38 40 days days

Ffigpigig 2 peicentpercentpeipel cent geigelgerminationmmcition oftnglochintriglochintrigdochinTrigbochindochin mantimamaktimamaritimamarimarlmafitima seeds after being transferred from 0 100 200 300 400 and 500 mol inm 3 nacl at temperature regimes ofaof5of 5 15c150c 5 25c 10 20c and 15 25c

noor and khan 1995 and chrysothamnus 500 mol Mrrr33 nacl was temperature depen- nauseosus khan et al 1987 dent keiffer and ungar 1995 exposed seeds seeds of triglochin maktimamantimamaritimamatimarimatltima from the utah of 5 halophytes atnplexatriplex prostrateprostrataprostrata hordeum population when transferred to distilled water jujubatumbatum salicorniaSahcorma europaeaeuropaea spergulariaSpergulana after a 20god d treatment at various salinity con- mannamarina and suaeda calceoliformis to salinity centrationscentrations responded differentially under treatments for 2 yr and determined their re- different temperature regimes there was lit- covery responses when transferred to distilled tle lecoveryrecovery 20 with the 10 20cgoc tempera- water they used the woodell 1985 classifica- ture regime in nonsalinenon saline controls but at tion system and placed atriplex prostrateprostrataprostrata seeds 5 25c seeds incubated previously at 400 mol in type I1 recovery inhibited by high salinity m 3 nacl had about 72 recovery it seems hordeum jubatum and Spergspergulariaulana mannamarinamavina in that recovery germination of T mantimamaktimamaritimamarimarlmafitima is type 2 recovery equal to original controls and temperature dependent binet 1961b deter- salicorniaSahcorma europaeaeuropaea and suaeda calceoliformiscalceohformis mined that seeds of T maktimamantimamaritimamatimarimatltima from a french in type 3 salt stimulated recovery greater coastal population had a stratification and light than controls our data from the 500 mol m 3 lequnementrequirement and when immersed in seawater nacl treatment indicate that T dantimamaritinamantimamaritimamarimarlmafitimafima at YC for 60 80 d they were capable of ger- recovery germination could be classified in minaminatingting subsequently in freshwater upon type 1 10 20cgocgog or type 2 15 25c depend- transfer to 25c in the light or dark seeds ing on the temperature regime used in the hornflomfrombormbomm the utah population did not require recovery germination experiment seeds ex- stratification and were not dormant woodell posed to 5 15c in all salinity treatments had 1985 included T maktimamantimamaritimamatimaritimafima in the group of low recovery germination percentages coastal species whose subsequent germination triglochin dantimamaritinamantimamaritimamarimarlmafitima seeds had maximum is stimulated by exposuieexposure to high salinity al- germination at a 5 25c temperature regime though the germination percentages he re- at all nacl concentrations tested few seeds corded were low our results indicate the germinated at the 5 15c in nonsalinenon saline con- iccoveryrecovery germination response of T dantimamaritinamantimamaritimamarimarlmafitima trols inability to germinate at low day tem- seeds in distilled water after 20 d exposure to peraperatureture in the laboratory indicates that a 1999 germination OF triglochinTRICLOCHIN MARITIMA 149

loo100 S a 80 OC 0 102010 20 OC 152515 25 60

40 2.2 M E 20

0 loo100 5255 25 OC 0 2 4 6 8 10 12 14 16 18 20 80 0 60 3 0 mol M loo100 mo 3 .2 40 200 mo m 2 3 M v 300 mol M 3 a 400 mol M E 20 500 mol m 4 0 0

0 2 4 6 8 10 12 14 16 18 20 days

fig 3 percent lecoveryrecovery germination in freshwater of ungerminated triglochin maritinamaritnaritimamaritimatima seeds initially exposed to 0 100 200 300 400 and 500 mol m 3 nacl at temperature regimes ofofaof55 25c 10 20c and 15 25c

threshold of higher day temperatures is neces- germination and recovery of seeds of T mar sary to stimulate germination under field con- itimaidima from hypersalinehypersaline conditions seeds were ditditionsions A combination of reduced salinity and not dormant but did have specific temperature high daytime temperatures stimulates germi- requirements for maximum germination nation and determines the sites along salinity gradients where germination and establish- acknowledgments of T soil ment maritinamaritimamaritima can occur because MA khan would like to thank CIES wash- salinity stress usually increases during the ington for a fulbright scholar research grant summer months salinity conditions early in department of environmental and plant biol- the growing season at the germination stage ogy ohio university for provision of facilities determine whether halophytes will be able to and the university of karachi for granting a successfully establish at a site ungar 1995 sabbatical leave we also thank dr wilford M recovery germination responses were also hess and dr darrell J weber for their help in dependent on temperature ranging from 0 arranging a field trip to collect the seeds from recovery at 5 15c to 72 at 5 25c seeds faust utah and Ms mehar noor and Ms of triglochin maritinamaritimamaritima will germinate when soil Bilquebilqueeses gul for separating and cleaning them salinity is low and the temperature regime is appropriate under natural conditions or they literature CITED will remain dormant in the seed bank until BADGER KSK S AND IAI1 A UNGAR 1989 the effects of salin- soil salinity is reduced and an appropriate ity and tempeltemperatureatnreatnee on the germinationgen nination of the inland temperature regime occurs our laboratory halophyte hordeum jujubatumjubatnmbatum canadian joinjournalnalnai of investigations with seeds from this utah popu- botany 67142067 1420 1425 BARBOUR 1970 germination and eailyeallyearlycarly of lation more precisely MG glowthgrowth define the temperature the strandsti and plant cakile dantimamantimainaritimainaritima bulletin oftheodtheof the tor- and salinity conditions necessary for maximum leyrey botanical club 971397 13 22 150 GREAT BASIN naturalist volume 59

BEWLEYBEWLLY JDJ D AND M BLACK 1982 physiology and bio- KHAN MAM A N SANKHLA DJD J WEBER AND EDE D chemistry of seeds springer verlag berlin 375 appp mcarthur 1987 seed germination characteristics binelBINLIBINET P 1959 doldormancesdoimancesDoiDor maneesmances pripriyairepnmaireprimairemairemalre et secondairesecon daire des of chrysothamnus nauseosus sspasp viriviridulusdulus Astereastereaeae serencessemencessemences de triglochin mantimaritimummantimummaritimum L action du froid asteraceae great basin naturalist 4722047 220 226 et de la lumiere bulletin de la societe linneenneLinneenne LOTSCHERT W 1970 keimingkeimungKeimung transpiration wasser und de normandie caen 9eae seriesserles 1013110 131 142 loneaufnhameloneaufohame bei glycophyten und halophytenHalop hyten 1960 rapportsRaprapeportsports entree 1 eau de meimer et la germina- oecologia Planplantarumplantariumtarum 52875 287 300 tion des serencessemensemencesces de triglochin maritimaritimummum L bul- MACKE A AND IAI1 A UNGAR 1971 the effect of salinity letin de la societe linneenneLinneenne de normandie caen on germination and early growth of Puccipuccinelliapuccinelhanellia nut loeioe10e senesseriesserles 11171 117 132 talhanatallianatashanataltaigallianalianallanailana canadian journal of botany 4951549.515 520 1961a action d une brusque modification de NAIDOO G AND K NAICKER 1992 seed germination in piesplespressionsionslon osmotique et de ph sur laid germination des the coastal halophytes triglochin bulbosabalbosa and triglo- semensentiencesserencessemencesces de triglochin mantimantimummaritimaritimummum L bulletin de la chin striatastriana aquatic botany 422174821742 217 229 societe linneenneLinneenne de noimandieNoinormandienor mandiemandle caengaencaen loeioe series NOOR M AND KHAN MAM A 1995 factors affecting germi- 2116 123 nation of summer and winter seeds of halopyrumHalopyrum 1961b196 ib acquisition de 1 aptitude a germer en mucromucronatumnatum under salt stress pages 51 58 inm MAM A milieu salesaie paipalpar les serencessemencessemen ces de triglochin mantimaritimafitimantz khan and IAI1 A ungar editors biology of salt tolerant mum L bulletin de laid societe linneenneLinne enne de nor- plants department of botany university of karachi mandie caencaen loeioe series 21242 124 128 pakistan 1965 action de divers rythmes thermiques jour PARHAM MRM R 1970 A comparative study of mineral nutri- saliersnaliersndliers suisulsur laid germination de serencessemencessemences de triglochin tion of selected halophytes and glycophytes doct- mantimantimummaritimaritimummum L bulletin de laa societe linneenneLinneenne de oral thesis university of east anglia nolnormandieNoi mandiemandle caen loeioe seriesserles 6996 99 102 PIGOTT CDC D 1969 influence of mineral nutrition on the 1968 dormancesDormances et aptitude a germer en milieu zonation of flowering plants in coastal marshes pages sale chez les halophyteshdlophytes bulletin de la societe de 25 35 in IHI1 H rorisonronson editor ecological aspects of francefianceflance physiologic vegetalevegetdleVegevegetatemege taletaie 1412514 125 132 mineral nutrition in plants symposia of british ecol- BOLENBOLLN EGE G 1964 plant ecology of spring fed salt marshes ogical society 9 blackwell scientific publications ofwesternof western utah ecological monographs 3414334 143 166 oxford DAVY AJA J AND GEG F bisibishopbi&hopioeloeiop 1991 biological flora of the SENECA EDE D AND AWA W COOPER 1971 germination and botishbritish isles no 172 journal of ecology 7953179 531 555 seedling response to temperature daylengthdaylength and GurrcurrGUTILRMANERMAN Y 1986 influences of environmental factors salinity by ammophila breviligulata from michigan on germinationgeigel mi nation and plant establishment in the negev and north carolina botanical gazette 132203132 203 215 highlands of israel pages 441 443 in PJ joss PW SHELTLER SGS G AND LEL E SKOG 1978 A provisional check lynch and OB0 B williams editors rangelands a list of flora of north america revised missouri resource under siege australian academy of sci- botanical garden 199 appp ence canberra SPSS INC 1994 SPSS SPSS 616.16gi 1 for windows update kellierKLIIIERKEIFFER CWC W AND IAI1 A UNGAR 1995 germination re- SPSS inc USA 30 appp sponses of halophyte seeds exposed to prolonged UNGAR IAI1 A 1962 influence of salinity on seed germina- hyphypersalinehypersalmehypersalmesaimeersaline conditions pages 43 50 inm MAM A khan tion in succulent halophytes ecology 4376343 763 764 and IAI1 A ungar editors biology of salt tolerant 1974 population dynamics of inland halophytic plants department of botany university of karachi communities bulletin de la societe Botbotaniquebotamqueanique de pakistan france 121287121 287 292 KHAN MAM A 1991 studies on germination of cressa creticecretica 1978 halophyte seed germination botanical pakistan journal of weed science research 4894 89 98 review 4423344 233 263 KHAN MAM A AND Y RIZVI 1994 effect of salinity temper- 1991 Ecophysiology of vascular halophytes CRC ature and growth regulators on the germination and press boca raton FL 209 appp ealiseailyearlyeally seedling growth of atriplex griffitgnffithligriffithiigriffithivhiihilhll var stock 1995 seed germination and seed bank ecology of siisilsllsitsti canadian journal of botany 724757947572 475 479 halophytes pages 529 544 in J kigel and G galili kilanKHAN MAM A AND ILA1 A UNGAR 1984 the effect of salinity editorsediedltoistols seed development and germination marcel and temperaturetemperatuietule on the germination of polymorphic dekker new york seeds and growth of atriplex triantnangulanstriangularisgularis willd WOODELL SRJS R J 1985 salinity and seed germination pat- american journal of botany 7148171 481 489 terns in coastal plants vegetatiovegetation 6122361 223 229 KHAN MAMAANDDJAND DJ WEBER 1986 factors influencing seed geigerminationmination in salicorniaSahcomia pacifica var utahensis received 10 february 1998 american journal of botany 73116373 1163 1167 accepted 14 july 1998 great basin naturalist 592 199901999 appp 151 159

AN irvingtonianIRVINGTONIAN SPECIES OF brachylagusbbachylacus MAMMALIA lagomorpha FROM PORCUPINE CAVE PARK COUNTY COLORADO

colleen N ramoslramos1ramose

ABSTRACT brachylagus is currently a monotypic genus of uncertain origins and known only from holocene and late rancholabiRanchorancholabreanlabi ean A new species of lepondleporid is described from the early and middle pleistocene irvingtonianIrvingtonian deposits of porcupine cave park county Coloiacoloradodo stratified deposits of the pit and the velvet room 2 localities within porcupine cave have been dated biochronologically and magneticallypaleomagneticallypaleo from the middle irvingtonianIrvingtonian and early to middle irvingtonianIrving tonian respectively brachylagus coloradoensis sp nov is characterized by its conserved p3pa enamel patterns which are intermediate between B idahoensis and hypolagusHypolagus and its size which is slightly larger than that of B idahoensis this suggests a possible ancestral relationship between hypolagusHypo lagus and brachylagus

key words brachylagus irvingtonianIrvingtonian brachylagus idahoensis pygmy rabbit leporidaeLeponfahdaedah lagomorphalagotLagornorpha pleistocene

brachylagus idahoensis pygmy rabbit the brachylagus idahoensis being the smallest sole living species of brachylagus is a small leporid in north america is easily identified rabbit that is restricted to dense stands of by its unique p3pa enamel patterns and its small sagebrush artemisia sppapp in the great basin size even so few paleontological sites include an isolated population also survives in eastern this species in their faunal lists thus it has a washington and is currently the subject of limited fossil record that extends only to the attention by conservationists because of its late rancholabreanrancbolabrean kurtenkurten and anderson highly specific habitat needs as reviewed by 1980 however material from cathedral dobler and dixon 1990 the species is vulner- cave an irvingtonianIrvingtonian site in eastern nevada able to local extirpation as its natural habitat is has yielded isolated teeth easily referable to B modified andor destroyed by agriculture and idahoensis CJ bell personal communication cattle grazing the species has apparently despite its limited geologic presence this existed in the great basin area for much of its genus has been hypothesized to have origi- evolutionary history as only a single paleonto- nated either in the miocene and descended logical site is known from outside its current from the genus alilepusAlilepus hibbard 1963 or range walker 1987 later in the pliocene and again derived from the genus has a long history of taxonomic alilepusAlilepus white 1991b these hypothetical uncertainty but is now generally considered to ancestriesances tries are based upon formation of the p3pa be valid the species now known as B ida enamel pattern found in brachylagus which is hoensis was first described by merriam 1891 completely distinct from other north ameri- but ascribed to the genus lepus miller 1900 can leporidsleporideleporids given the limited fossil record of formally proposed the subgenus brachylagus brachylagus and extensive radiation of the which lyon 1904 elevated to full generic sta- genus Hypohypolaguslagus during the pliocene a rea- tus a usage maintained by nelson 1909 later sonable alternative hypothesis is that brachyabrachy authors placed the species in the genus of lagus arose directly from hypolagusHypolagus in the late sylvilagusSylvilagus grinnell et al 1930 orr 1940 dur- blancaablancan or early irvingtonianIrvingtonian these hypothe- rant 1952 more recent morphological work ses are summarized in figure I1 and evaluated has supported the generic status of brachyla below gus kenner 1965 with recent genetic work the species described here occurs in the reviewed by chapman and ceballos 1990 irvingtonianIrvingtonian deposits of porcupine cave park further substantiating this county colorado the cave contains several

elveienverenvei museum of natural history 2001 Coloiacoloradodo blvd denver CO 80205579880205 5798 and university of Coloracoloradodoatt boulder campus box 334 bouldeibouldenBouldei CO 80309

151 152 GREAT BASIN naturalist volume 59

hypolagusHypolagus atialilepusaftlepusalfAliAftlepus brachylagus

alilepusAlilepus brachylagus

c Q hypolagusHypolagus brachylagus

fig 1 Surnsummarymary of hypotheses concerning evolution of p3pa form seen in brachylagus A brachylagus descending from afilepualilepuhAlilepuhpus postelposterior101 external reentrantleentiant PER fuses with posteriorpostenoi internal reentrant to form separate lobes as seen in bi achylagusachybrachylaguslagus summarized hornfrombormbomm hibbard 1963 B brachylagus descending from alilepusAlilepus which arose from Hypohypolaguslagus during late pliocene 01or eailyearlyeally pleistocene aftelahterafter white 1991b199 ib and C brachylagus descending directly from hypolagusHypolagus PER eventually extends completely across width of the tooth to separate the trigonid fromblom the talonid loomsrooms hornfromhormhomm which fossils have been collected apparently undisturbed these lower levels are moskybarnoskyBa and rasmussen 1988 wood and also yielding mammal species such as hepolahypola balbarnoskyBai nosky 1994 anderson 1995 these loomsrooms gus that are older than those found anywhere and areas lepieleplerepresentsent localities within porcu- else in the cave although other rooms with pine cave and include the pit velvet room fossilized remains exist and have been sam- and mark s sink fossil deposits range in age pled the 3 rooms described above have pro- hornfromhormhomm eailyeallyearly to late irvingtonianIrvingtonian the pit a duced most fossil specimens to date and may small loomroom in the cave was excavated by the collectively span more than one million years calcainegiecarnegieCai neglenegie museum CM in the 1980s and was in the 1980s certain specimens from the pit originallyongmallymaily thought to date between approxi- were identified as brachylagus idahoensistdahoensts by mately 0380.380 38 my to i oughlyhughlyroughly 080.80 8 my barnosky barnosky and rasmussen 1988 while ana- and rasmussen 1988 wood and barnosky lyzing the lepondleporid remains from the velvet 1994 more lecentrecent analyses date the majority room and mark s sink I1 discovered several oftheodtheof the pit aioundaround 850000 ybpybyap p barnosky per- teeth and mandibles similar in size to those of sonal communication paleomagneticPaleomagnetic studies B idahoensisidahoensts but lacking the diagnostic p3pa of the velvet room have found that the upper enamel pattern of this species when I1 reex- 5 levels coneoncorrelateelate to the present normal polar- amined the pit specimens I1 found that they ity while the majority of the lowellower levels were too lacked the B idahoensisidahoensts form of p3pa previ- deposited during the matuyamaMatmaruyamauyama reversed polar- ous morphometncmorphomorphometricmetriemetric work ramos in press pro- ity chron B raynolds and J friedman per- vided a large sample of extant B idahoensis n sonal communication and aiealeare therefore at 85 as well as an understanding of variation least 0780.780 78 million years inm age mark s sink an in dental characters for this species when isolated section of the velvet room contains used for comparison with the porcupine cave biotabioturbatedbiotm bated material in its upper sections fossils these data demonstrated that although while its lower portions aieare stratified and the fossils were similar in several ways to B 1999 irvingtonianIRVINGTONIAN SPECIES OF brachylagus 153

idahoensis distinct differences indicated a systematic paleontology new species of brachylagus class MAMMALIA linnaeus 1758 order lagomorpha brandt 1855 METHODS family LEPORIDAE fischer de waldheim 1817 porcupine cave is located at 2900 in in the genus brachylagus miller 1900 nwi4nw14 ofswl4of sw14 sec 23 tiss r76w in park brachylagus coloradoensis county colorado fossil material was col- new species lected by denver museum of natural history DMNH crews using 064 m2ma grid excavated HOLOTYPE DMNH 33261 incomplete in 2 cm levels for the stratified deposits of the right dentary with p3pa lacking incisor p4pa ml velvet room the upper region of mark s sink m2ma and m3ma coronoid process condyloid pro- is unstratified and the grid system was not cess and angle of mandible fig 2 applied however the lower sections separate into strata that have been carefully excavated utilizing the grid and level system described above material was screened and washed then brought to DMNH for identification and cata- loging material from the pit was excavated using different methodology see barnosky and rasmussen 1988 and much of that material is housed in CM using mitutoyu calipers calibrated to 0020.02 minmm I1 measured the fossil material as well as 85 recent skulls of brachylagus idahoensis for comparison measurements followed previously published conventions bensley 1926 findley et al 1975 white 1987 and are as follows 1 depth of mandible at anterior alveo- lus of p4pa 2 length of alveolar tooth row of mandible 3 length of p3pa 4 width of p3pa 5 width of p4pa 6 width of ml 7 width of m2ma 8 width of m3ma 9 length of diastema 10 length of maxillary alveolar tooth row 11 depth of anterior zygomatic process 12 width of p2pa 13 width of p3pa 14 width of p4pa 15 width of ml 16 width of m2ma 17 width of m3ma Diastemdiastemataata of maxillarmaxillaemaxillae are rarely preserved and were not included in the analysis enamel pattern terminology follows white 1987 fig 3 the data were then subjected to descriptive statistics using microsoft excel in addition a 2 sample t test was performed upon widths and lengths ofofp3p3pa between the 2 species only adult specimens of both the recent and fossil material were used fossil specimens were determined to be adult if the cheek teeth showed no alteration in size and enamel pattern from their base to their occlusal surface sutures were well knit and the bone was not fig 2 DMNH 33261 boloholotypetype for B coloradoensis highly porous specimens were determined to top and mandible of brachylagus idahoensis UCM 5840 shown from labial camera be and excluded both view lucida immature from the analysis if drawing ofp3of p3pa from holotypebolotype labial is to left lingual to the any of the above conditions were not met right top is anterior bottom is posterior 154 GREAT BASIN naturalist volume 59

HYPODIGM pit CM 66408 66409 p3pa than the average B idahoensis although some CM 66431 edentulous right maxilla CM specimens fall within the range of individual 66432 edentulous left maxilla CM 65486 variation for B idahoensis tables 1 2 results left dentary with p4pa m2ma CM 66604 p3pa CM of the 2 sample t test indicate that p3pa of B 65607 p3pa velvet room DMNH 28901 idahoensis and p3pa of B coloradoensis differ in 3325033255338503325533250 33255 p3pa mark s sink DMNH 28902 length f8119fgiagi 19 454.5 P 000010.0001 and width left humerus DMNH 33257 left dentary f8119119 2662 66 p 0001ooi0.001001 these results and with p3pa p4pa DMNH 33260 three isolated the aifferencesdifferences between enamel patterns indi- psp3s DMNH 33272 four isolated psp3s cate that the 2 samples are not drawn from the DMNH 33271 left dentary with p3pa m2ma same population and support the designation DMNH 33270 left dentary with p3pa ml of B coloradoensis as a species separate from TYPE LOCALITY AND AGE middle pleis- B idahoensis the PER ofboflofbB coloradoensis does tocene ealiseailyeallyearly to middle Irvingirvingtonianirvmgtomantonian DMNH not extend completely across the tooth width locality no 1349 porcupine cave 2900 in as in B idahoensis no complete skulls of B park co colorado nwi4nw14 of swi4swibsw14 sec 23 coloradoensis are known from this site how- tiss r76w lat 384345n3804345n long 105 ever portions of dentariesdentaries and edentulous 5r4rw5141w cribblesCribGribgribblergribblesblesbies park 757.57 5 quad maxillarmaxillaemaxillae are present disarticulated postera ETYMOLOGY named for the locality and nial material has been tentatively assigned to following the precedent set in this genus at this species due to the unusually small size present this species is known only from a sin- which allows no other confident identification gle site in colorado within the leporidae several catalogedununcatalogeduncatalogued DIAGNOSIS the enamel pattern of the p3pa humeri ulnae and podial remains have thus is distinct from that of B idahoensis in having been identified to B coloradoensis and are a posteroexternal reentrant PER that extends similar in size to corresponding elements of B between 13 and 23 the width of the tooth idahoensis fig 3 the posterointernal reentrant PIR when present is only a slight indentation discussion dimensions of postpostcranialcranial material strongly resemble those of modern B idahoensistdahoensis but brachylagus coloradoensis sp nov shows are slightly more robust some affinity to the extinct genus hypolagusHypolagus comparisons brachylagus coloradoensis that radiated profusely in the pliocene fig 3 may be distinguished from hypolagusHypolagus species although some psp3s of B coloradoensis display by its smaller size also the PER ofofblofbB colora patterns with clear PIRs as seen in ailalilepusafilepusahiAliAfilepus doensis p3pa extends farther across the tooth none of the upper molars found and ascribed width generally greater than 12 the distance to B coloradoensis display the characteristic fig 3 although B idahoensiszdahoensis is of similar enamel lake found in ailalilepusafilepusahiabiAliAfilepus this lake is also size B coloradoensiscoloradoenszs sp nov is generally absent in hypolagusHypolagus hypolagusHypolagus is easily dis- more robust in comparing the size of p3pa B tinguished by its simpler p3pa enamel patterns coloradoensis is approximately 12 15 larger and larger size

1 C QQc oe DQQ ia A B C D E F

3 fig comparisoncolnCoinpanson ofp3of p3pa enamel patternspatteins for hypolagusHypo lagus B coloradoensis and B idahoensis A is hypolagusHypolagus redrawn from white 1987439 B E DMNH unknown DMNH 33257 DMNH 33255 and DMNH 33260 represent the iangelangerange ofvdiiationof variation seen in specimens of B coloradoensis B predominates slightly in earlier sediments whereas C becomes somewhat moiemolemore plentiful in younger sediments at porcupine cave F is that ofoijaoij3B idahoensis UCM 5840 note also that some specimens ofofblofbB coloradoensis have a slight posterior internal reentrant PIR 199919991 irvingtonianIRVING TONIAN SPECIES OF brachylagusbrachyiagus 155

TABLE 1 dimensions and descriptive statistics of dentary and lower dentition of brachylagusbrachylagns coloradoensis and B idahoensis min parentheses values for B idahoensisiddhoensis are for left measurements only measurement xaf5f s range n alveolar length of toothrow 104810.4810 48 na na 2 919 1 0340.340 34 8248 24 102610.2610 26 85 length of diastema 10910.910log 9 na na I1 90go909.09 0 0430.430 43 8468 46 119211 92 85 depth of dentary at p4pa 8868 86 0300.300 30 7747 74 9569 56 4 7547547.547 54 0420.420 42 6606 60 8268.268 26 85 length ofp3of p3pa 2022 02 0280.280 28 1741 74 2568562.562 56 19 17817810111781.78 oiioli0110.11 1621621.621 62 2282.282 28 85 width of p3pa 1801.801 80 0220.220 22 1401.401 40 2122.122 12 19 iglg161011161.6 olioii011 1421 42 2182.182 18 85 width ofp4of p4pa 2342 34 0120 12 2122 12 2542 54 5 1961.961 96 010 1 1681681.681 68 2222.222 22 85 width of mlm 1 2352 35 0120 12 2242 24 2468462 46 4 1981981.981 98 010.10oi 1 1781781.781 78 2262.262 26 85 width of m2ma 2222.222 22 0160.160olg16 2082.082 08 2402.402 40 3 1941941.941 94 010.10oi 1 1741741.741 74 2202.202 20 85 width of m3ma 1701.701 70 na na 2 1021021.021 02 00900.0909 0820 82 1301 30 85

TABLE 2 dimensions and descriptive statistics of maxilla and upper dentition of brachylagus coloradoensis and B idahoensis in parentheses values for B idahoensis are for left measurements only measurement xT s range n alveolar length of toothrow 100810.0810 08 na na 2 9139139.139 13 03500.3535 8328328.328 32 10loi10110.11 85 depth of zygomatic process 3493 49 na na 2 3493493.493 49 0310.310 31 2882882.882 88 4244 24 85 width of p2pa 2092.092 09 na na 2 1731731.731 73 0140.140 14 1521521.521 52 2362.362 36 85 width ofofp3p3pa 3193 19 na na 2 2962962.962 96 020.2ob0 2 9582582 58 3383 38 85 width ofofp4p4pa 3193.193 19 na na 2 2982 98 0170 17 2522522.522 52 3643.643 64 85 width of mlM 1 3053.053 05 na na 2 2872 87 0180.180 18 2362 36 3263 26 85 width of m2ma 262.62 6 na na 2 26262.62 6 0150 15 2322322.322 32 2962.962 96 85 width of m3ma 1 15 na na 2 10710710181071.07 0180.18 06og060.60 6 1841.841 84 85 156 GREAT BASIN naturalist volume 59

the lower third premolar pap3 is considered of the hypotheses concerning the ancestral somewhat diagnostic among leponislepondsleporidsleporids and has stock of this genus fig 1 it appears just as been used in numerous species descriptions likely that brachylagus arose directly from especially of extinct lepoleponislepondsleporidsrids white 1984 1987 hypolagusHypolagus as from alilepusAlilepus although B col 1991a 1991b hibbard 1963 in B colora oradoensis exhibits psp3s of both hypolagusHypolagus and doensis there is some variation in this charaecharac- alilepusAlilepus form there are no enamel lakes pre- ter which based upon the limited number of sent in any of the upper molars examined specimens available seems to be present re- enamel lakes in upper cheek teeth are absent gardlessgardless of geologic age in short no particu- in hypolagusHypolagus but present in p3pa of alilepusAlilepus lar evolutionary trend is seen in this character and may represent plesiomorphic characters although the simplest form is slightly more retained from paleolaginae from which archae prevalent in older strata and the more derived olaginaeolaginae arose also the time framesfranics hypothe- form seems more prevalent in later strata sized by hibbard 1963 and white 1991a both extremes of the continuum are present in are unsubstantiated by the fossil record as oldest and youngest strata the range ofoftanaofvanavariavarla there is no evidence that brachylagus existed tion in the p3pa enamel pattern of B colora earlier than the very latest blancaablancan to earliest doensis is depicted in figure 3 p2pa irvingtonianIrvingtonian is in the con- genera tains a single anterior reentrant as in B ida such as lepus and sylvilagusSylvilagus are hoensis and hypolagusHypolagus not considered to have originated directly from lagus the size of the mark s sink B coloradoen Hypohypolagus one or more potential inter- mediates have hypothesized sis which is presumably the oldest in the cave been over the Ali plio is somewhat larger than that of either the vel- years including alilepuslepus serengetilagus aliopilo pentapentalaguslagus and others vet room or pit specimens of B coloradoenstscoloradoensis hibbard 1963 white 1991b and fossil evidence fig 4 while retaining more conservative p3pa does not dispute these hypotheses however B coloradoensis enamel patterns this is an intriguing trend appears to indicate a and may indicate a time frame for the clado more direct transition from Hypohypolaguslagus to brachylagmbrachylagus thus the living genic event that gave rise to the genus brachyabrachy pygmy rabbit may be only lagus unfortunately the strata from which the direct descen- dent of an extinct genus and may be more dis- these earliest specimens come are bracketed tantly related to sylvilagusSylvilagus than previously only with upper and lower time limits and thought although halanychmalanychHalanych and robinson there is little refinement in the age estimates in 1997 found some molecular evidence placing of the levels themselves therefore a more brachylagus as sister taxon to Sylvisylvilaguslagus based accurate picture of the rate at which the variavarla varia- upon mitochondrial 12s12 S data other tion narrowed is not possible at this rdnaadna is time aspects of their analysis did not support this however the wide range of size size variation strongly I1 find that the high degree of mor- noted for B specimens of coloradoensis in the phologicalph differences between brachylagus eailyeallyearly of cave Irvingirvmgtomanirvingtoniantonian porcupine fig 4 and sylvilagusSylvilagus as well as the paleontological may indicate that the cladogeneticclado genetic event evidence described above present a strong occurred shortly in geologic terms before the case against a close relationship between them porcupine cave record at least 2 species of see also kenner 1965 green and flinders hypolagusHypolagus as yet unidentified are present in 1980 the older sections of mark s sink with the B given the high degree of habitat specificity coloradoensis material one of these species is exhibited by the extant species of brachylagus very small though still somewhat larger than it would be interesting to determine whether the largest specimens of B coloradoensis the its extinct species was also highly habitat spe- relationship between B coloradoensis and this cific unfortunately paleopaleoecologicalecological evidence small Hypohypolaguslagus is unclear at present A more at porcupine cave is scanty and restricted detailed phylogenetic analysis utilizing several mostly to faunal remains such postcranialpostcranial re- characters is currently in progress and should mains as have been assigned to B coloradoen provide stronger evidence concerning the ori-oriorl sis resemble B idahoensis in almost all details gins of brachylagus and its relationships to this indicates a similar degree of locomotor extinct and extant genera CNC N ramos in adaptation as seen in the extant species braceybrocey preparation lagus coloradoensis was probably also closely 199919991 irvingtonianIRVINGTONIAN SPECIES OF brachylacusbrachylagus 157

252.5

E 2 E w CL 0s 151.5

0CM s 1

050.5

0 0 5 10 15 20 approximate chronological order 161 6 from marks sink 7187 18 from velvet room

fig 4 change in p3pa length over time size data plotted over time for the different rooms in porcupine cave length of p3pa has been used to illustrate the general trend that of wide size range in older sediments then loss of upper size limits and closer approximation to size range of B idahoensis line indicates average p3pa length foifolfor B idahoensis

allied to brush and dense cover and avoided although complete species associations can- open areas although the plant biota compris- not be reconstructed at this time for porcu- ing its habitat is impossible to ascertain at pine cave it is notable that this new species this time however barnosky and rasmussen of lepondleporid was associated with extinct lepoleponislepondsleporidsrids 1988 reported the consistent concurrence of such as azdanolagusaztlanolagus and Hypohypolaguslagus in the old- sage vole lagurus curtatuscurtatus with B colora est sections of porcupine cave yet found with doensis reported as sylvilagusSylvilagus idahoensis extant species such as lepus townsendfowntowntownsendiitownsendutownsendiasenduii and this arvicoline rodent is generally found in sylvilagusSylvilagus audubonnaudubonii in younger sediments at stands of artemisia armstrong 1972 and cur- this site ramos 1998 these associations do rently has a distribution overlapping that of B not necessarily indicate sympatry but do indi- idahoensis both mammal species are absent cate a temporal coexistence sympatry is rare from the wyoming basin as is B idahoensis among species of rabbits and hares and the despite the apparent presence of suitable habi- remains at porcupine cave are the result of tat walker 1987 the presence of an irving predation by raptorsraptores and mammals some of tonian species of brachylagus on the eastern which have large ranges prey remains are side of the continental divide and the persis- thus sampled from a potentially wide radius tence of extant brachylagus in the great basin and do not represent close species associations may indicate that the 2 populations became it is still intriguing to find that this little lep- isolated on opposite sides of the rocky moun- ondorid was able to survive while other lepondleporid tains after dispersing through the wyoming species even genera were going extinct basin where the genus originated is unclear unfortunately the fossil record at porcupine and although I1 find the presence of a small cave truncates several hundred thousand hypolagusHypolagus associated with B coloradoensis in years ago and we cannot know at this time the older strata of mark s sink to be sugges- how long B coloradoensis survived in associa-assoriaassocia tive it certainly is not conclusive tion with modern fauna 158 GREAT BASIN naturalist volume 59

porcupine cave is continuing to be exca- BENSLEY BAB A 1926 practical anatomy of the rabbit P vated and specimens of this new species are biakisblakistonBlakis tonss son & co philadelphia 298 appp CHAPMAN being added to the collections at DMNH in J A AND G CEBALLOS 1990 the cottoncottontailstails pages 95 110 in JAJ A chapman and JECJ E C flux edi- addition temporal limits of localities within tors rabbits hares and pikas status survey and the cave are being clarified which will allow conservation action plan international union for more refined analysis of morphological change conservation of nature and natural resources speciation rate and faunal associations As fur- JUCNIUCN gland switzerland ther analysis of this important irvingtonianirvingtomanIrvingtoniantoman DOBLER FC AND KRK R DIXON 1990 the pygmy rabbit pages 111 115 inOT JA chapman and JECJ E C flux site continues our understanding of this extinct editors rabbits hares and pikas status survey and species will continue to grow hopefully pro- conservation action plan international union for viding new insights into the evolutionary history conservation of nature and natural resources of the single living species of brachylagus IUCN gland switzerland DURRANT SDS D 1952 mammals of utah taxonomy and distribution university of kansas publications muse- acknowledgments um of natural history lawrence 616 1 549 flndleyfendleyFINDLEY JSJ S AHA H HARRIS DED E WILSON AND C JONES david daitch of the university of colorado 1975 mammals of new mexico university of new at boulder provided the excellent drawings for mexico press albuquerque 360 appp GREEN S AND T FLINDERS 1980 brachylagus ida the figures I1 also thank the following individ- JSJ JTJ hoensis mammalian species 1251125 1 4 uals for their help with this work A cheri GRINNELL J J DIXON AND JMJ M LINSDALE 1930 verte- jones and the denver museum of natural brate natural history of a section of northern califor- history dave armstrong of the university of nia through the lassen peak legionregion university of colorado at boulder elaine anderson russ california publications in zoology 35135 1 594 graham kathy honda and kirk johnson of HALANYCH KMK M AND TJ ROBINSON 1997 phylogenetic relationships of cottontailsofcottontailstailstalis sylvilagusSylvilagus lagomorpha the museum of natural cotton denver history jane congruence of 12s rdnaadna and cytogenetic data mol- bock herbert covert carol wessman greg ecular phylogeneticsPhylo genetics and evolution 72947 294 302 carey and jill skarstad of the university of HIBBARD CWC W 1963 the origin of the p3pa pattern ofsylviof sylvi colorado at boulder christopher J bell lagus Caprocaprolaguslagus oryctolagus and lepus journal anthony D barnosky julio friedman CA of Marnmamammalogylogy 44144 1 15 KENNER GHG H 1965 comparative osteology of rabbits of repenningReperming bob raynolds don and jerry the genera brachylagus miller and sylvilagusSylvilagus gray rasmussen lou taylor and all the porcupine unpublished master s thesis university of utah cave volunteers thanks to the following insti- salt lake city 125 appp tutions for the generous use of specimens KURTEN B AND E ANDERSON 1980 pleistocene mam- denver museum of natural history carnegie mals of north america columbia university press new york museum of natural idaho museum of 443 appp history LYON MWM W 1904 classification of the hares and their natural history field museum national mu- allies smithsonian miscellaneous collections seum of natural history university of colo- 451456321451456 321 447 rado museum and cowan vertebrate museum MERRIAMMEBRIAM CHC H 1891 mammals of idaho north ameri- william akerstonagerston anthony D barnosky and can fauna 5755 75 78 MILLER GSG S 1900 A new subgenus for lepus idahoensis john white acted as reviewers and their com- proceedings of the biological society washington ments helped improve the original draft 13157 NELSON EWE W 1909 the rabbits of north america north literature CITED american fauna 29129 1 314 ORROBR RTR T 1940 the rabbits of california occasional papers ANDERSON E 1995 preliminary report on the carnivorescarnearnivores of the california academy of sciences of porcupine cave park county colorado pages 19119igi 1 227 259 282 in K stewart and K seymour editors RAMOS CNC N 1998 evolution and biogeography of north palaeoecologypdlaeoecology and palaeoenvironments of late ceno- american leporidaeLepondae doctoral dissertation univer- zoic mammals tributes to the career of CSC S rufus sity of colorado at boulder 192 appp churcherchuicherChuicherichen university ofaofToftbrontoof brontoorontotoronto press toronto IN PRESS Morphomorphometricmorphometncmetriemetric variation among leponislepondslepoleporidsrids ARMSARMSTRONGtrongTRONc D M 1972 distribution of mammals in col- mammalia lagomorpha of the interior west pro- orado monograph university of kansas museum of ceceedings of the denver museum of natural history natural history 313 1 415 WALKER DND N 1987 late PleistocenepleistoceneholoceneHolocene environ- BARNOSKY ADA D AND DLD L RASMUSSEN 1988 middle pleis- mental changes in wyoming the mammalian record tocene arvicoline rodents and environmental change pages 334 392 in RWR W graham HAH A semken and at 2900 meters elevation at porcupine cave south MAM A graham editors late quaternary mammalian park colorado annals of the carnegie museum 57 biogeography and environments of the great plains 267 292 and prairies illinois state museum springfield 199911999 irvingtonianIRVINGTONIAN SPECIES OF bracmiagusbrachylagus 159

WHITE JA 1984 late cenozoic leporidaeLepondae mammalia 1991b north american leponnaeleporinaeleporidaeLeporLeponinaenae mammalia lagomorpha from the anza borrego desert south- lagomorpha from late miocene cialclarendonianClarenclai endomandoniandonlan to ern california special publication carnegie museum latest pliocene blancaablancanBlancan journal Vertebrateofvertebrateof pale- of natural history 9419 41 57 ontology 116711 67 89 1987 the archaeolaginae mammalia lagomor- WOOD DLD L AND ADA D BARNOSKY 1994 middle pleisto- pha of north america excluding archaeolagus and cene climate change in the colorado rocky moun- panolax journal of vertebrate paleontology 74257 425 tains indicated by fossil mammals from porcupine 450 cave quaternary research 4136641 366 375 1991a A new sylvilagusSylvilagus mammalia lagomorpha from the blancaablancan pliocene and irvingtomanirvingtonianIrving toniantoman received 21 november 19919977 pleistocene of florida journal of vertebrate pale- accepted 22 june 1998 ontology 1124311 243 246 cleatcreatgleatgreat basin naturalist 592 019991999 appp 160 168

A NEW SPECIES AND NEW SYNONYM IN THE GENUS psychoroniaPSYCHORONIA limnephilidaelimnophilidae WITH significant RECORDS FOR caddisfliesCADDISFLIES trichoptera FROM WESTERN NORTH AMERICA

david E ruiterlruiter1

ABSIRACIABSTRACT A new species of caddiscaddisflycaddishlyfly PsychpsychoroniapsychoromaPsychooroniaroma brooksi limnephilidaelimnephihddelimnophilidae is described from new mexico and Psychpsychoroniap&ychoromaoronia brevibrevipennispennis banks 1904 is designated as a junior synonym of PsychpsychoroniapsychoromaPsychooroniatomaroma costalis banks 1901 additional distnbutionaldistributional records and notes foifolfor 51 caddisflycaddishlycaddis fly species are also presented

key words trichoptera caddisliescaddiscaddisfliesitesfliesglieslies limnephilidaelimnephihdaelimnophilidae psychoroniaPsychoronia new species biogeography

the discovery of a new species psychoroofofpsychoro based on examination of the lectotype and niama in new mexico has led to a review of the paratype males in the type series of P costalis other 2 species in the genus and a resultant MCZ 11676 the single female holotype of synonymy additional work at small isolated P brevibrevipennispennis MCZ 11657 and numerous habitats in the andaridarld west will undoubtedly result series of P costalis specimens from colorado in the discovery of additional new species of P brevibrevipennispennis as wiggins 1975 suggested is trichoptera these isolated distributions also a new junior synonym of costalis the holo- emphasize the need to protect such habitats type female of P brevibrevipennispennis is small about 8 whenever possible mm from head to apex of abdomen when two species previously have been placed in compared to female specimens of P costalis the genus PsychpsychoroniapsychoromaPsychooroniafomaroma P brevipenbrevibrevipennisbrevipenmspennisms banks from colorado up to 14 mm total length 1904 and FP costalis banks 1901 wiggnswiggmswiggins however terminalia of the P brevibrevipennispennis holo- 1975 provided rationale for maintaining the type are well within the variability seen in P genus PsychpsychoroniapsychoromaPsychooroniaroma and figured the adults of P costalis females banks 1904 indication that costalis wiggnswiggmswiggins 1975 also noted the similar- the wing membrane of P brevibrevipennispennis does not ity of the 2 species and illustrated the larvae of have hairs is in error the membrane of the P costalis wiggins 1977 PsychpsychoroniapsychoromaPsychooroniafomafomoroma brevi wing is clothed with strong upright hairs as I1 pennis is known only from the female holotype indicated for P costalis ruiter 1995 the wing during preparation for this paper I1 attempted membrane of the new species described below to collect atnearannear the new mexico type locali- also has a few upright hairs although the ma- ties for both P brevipenbrevipennisbrevipenmsbrevipennisms and FP costalis the jority of the hairs on the membrane are fine and top of the las vegas range the type locality of recumbent similar to the genus hesperophylax P costaliscostahs is located west of las vegas new terminology for genital structures follows mexico and just west of beulah the type that of schmid 1980 locality of P brevipenbreulbrevibrevipennisbrevipenmspennisms is also located west of vegas the former las near townsite of beulah psychoroniaPsychoronia brooksi new species 2438 m I1 could not locate populations ofpsyofoppsypsyapsy figs 1 10 choreniachoroniachoronia at or near the beulah townsite based on discussions with residents in the area and the discovery of this new species occurred the condition of existing aquatic habitats it while I1 was looking for P brevibrevipennispennis the appears that many typical P costalis habitats occurrence of psychoroniaPsychoronia in this habitat a headwater springs and very small streams small high velocity stream was totally unex- weiwelwere e altereddestroyedaltered destroyed to create water supplies pected as my previous collections of psychoro for recreational homes being built in the area nia had been only from headwater spring

626016260 south crantgiantglantgrant sheetstreet littleton CO 80121

160igo 199911999 NEW SPECIES IN GENUS psychoroniaPSYCHORONIA 161

sources this collection is another example of triangular in dorsal ventral and lateral views a species occurring on an isolated mountain in in FP costalis the appendages of segment X are the southwestern united states this species very short not extending to apex ofofaofxX is named for bill brooks a comrade with LARVA figs 8 10 most characters typi- numerous interests one of which is occasion- cal of limnephilimlimnephilini wiggins 1977 mandibles ally collecting caddisfliescaddisflies each with cutting edge entire except for sinsin- ADULT wings and body yellowish brown gle subapical tooth in P costalis mandibles forewing membrane patterned with pale areas with numerous apical teeth head and tho- margined with darker brown fig 1 wing racic scsclescieselelentesrites dark nearly black with faint membrane with numerous long upright and muscle scars on head primary setae absent recumbent setae nearly as long as those on from anterior pronotal margin in FP costalis wing veins wings of female extending beyond primary setae are present and equally spaced apex of abdomen as long as wings of male dorsal and ventral gills present on abdominal length from front of head to end of forforewingsewings segments 2 7 most with 3 5 filaments lateral 16 18 mm spurs 1341 3 4 gills present only on segments 2 and 3 most MALE 2 4 with figs tergite VIII an specimens with 2 filaments for each lateral P apical patch of large upright spines in gill abdominal dorsal chloride epithelia absent costalis this patch comprises slender re- ventral chloride epithelia large present on segment IX separated dor- cumbent spines segments 2 7 sally with widest portion slightly above the CASES the larval case is made of sand mid lateral line inferior appendages large grains only slightly tapered from wide ante- directed dorsocaudad segment X with inter- riorbior end to posterior and slightly curved mediate appendages fused into triangular pupal case 17 20 mm made of larger rock caudal structure surrounding anal opening in particles similar to that of P costalis see wig its dorsal extended into view apex narrow ginsamsgms 1977 fig 1046 nearly straight not slightly bifid process in P costalis the apex of tapered several cases have incorporated occa- the intermediate appendages is acute and sional live fingernail clams recurved superior appendages large mesally TYPE SPECIMENS holotype and allotype concave dorsally slightly concave with blunt NEW MEXICO lincoln county north fork rio apices extending caudad nearly to apices of ruidosoRuidoso at entrance to ski apache ski area phallic inferior appendages paparameresparamerosrameres each collected as pupae 24 may 1997 emerged 20 terminated in several long strongly sclerotized june 1997 DED E ruiter deposited museum spines divided completely to base of para- of comparative zoology paratype males same mere the dorsal portion a thick sinuous data emerged 20 22 june 1997 deposited spine curving laterad apically and apex with museum of comparative zoology national minute serrationsserrations along dorsal margin the museum of natural history illinois natural ventral aedaedeagaleagal spines straight clumped at history museum royal ontario museum A their base in P costalis the paramerespaparamerosrameres are paratype male and female same data deposited shorter and the dorsal portion is composed of at NMNH A pupal eluviaexuvia and 4 larvae same short fused spines data deposited at each of the above collec- FEMALE figs 5 7 ventromesal sclero- tions the remainder of the adult 8 males I1 tized spur on sternite VI absent abdominal female pupal and larval material from the sternal setae equal in thickness to tergal setae same collection retained in author s collection tergite IX bardlikebandbandlikelike separated by faint suture DIAGNOSDIAGNOSISis Mmalesmaiesalesaies of P brooksi can be from its very small ventrolateral lobes ventro- separated readily from P costalis by the stiff lateral lobes widely separated ventrally by dark spines on tergite VIII versus the fine broad slightly sclerotized suprasupragenitalgenital plate hairhairlikehairlinelike spines on tergite VIII of P costalis in median lobe of vulval scale clavate with addition phallic parameresparamerosparameres of P brooksi are at broad truncate apex not extending caudad as least twice as long as those of P costalis far as apices of lateral lobes segment X fused females of P brooksi have normal length to tergite IX and with apex tubular its dorsal wings that extend well beyond the abdomen margin slightly cleft its ventral margin con- while wings of all females of P costalis I1 have cave entire appendages of segment X located examined do not extend beyond the ath8th dorsodorsolaterallylaterally extending well beyond its apex abdominal segment and usually do not exceed 162 GREAT BASIN naturalist volume 59

sup appopp oltwit oltwit intidt ix x ix inf s9psap

VMS VIIIS 1xv1 VS appopp

1xv1 x s9psap

parpanpon Vs

VIIIS

v111t

SUP appopp

ix ixtaxt 7

int inf 4 oltwit

figs 1 7 PsychpsychoroniapsychoromaPsychooroniaroma brooksi foiewmgforewing and genitalia 1 right side forewing 2 male genitalia left lateral view 3 male aedeagusacdeagusacdeagus left lateral view 4 male genitalia caudal view 5 female genitalia left lateral view 6 female genitalia ventral view 7 female genitalia dorsaldorsaidoidol salsai view vicitviiitv111t tergite VIII segment VIMVIIIs steritesternitestermte VIII segment ixt1xthixt tergite IX seg ment ixilixvl ventrolateral lobe IX segment X X segment app appendage infmf inferior appendage idtintmt intermediate appendage sapsgp suprasupragenitalsupiagemtalgenital plate sup superior appendage vs vulval scale 199911999 NEW SPECIES IN GENUS psychoroniaPSYCH ORONIA 163

W 10

figs 8 10 PsychpsychoroniapsychoromaPsychooroniafomaroma brooksi larva 8 head and thorax dorsal view 9 mandibles dorsal view 10 ninth abdominal tergite dorsal view the ath5th abdominal segment appendages of REMARKS the occurrence of a macropter- the female P brooksi segment X extend poste- ous female psychoroniaPsychoronia conflicts with the pri- riorly well beyond the apex of segment X mary character banks 1916 used to define while those of P costalis do not reach the apex the genus female short winged wiggins of segment X larvae of the 2 species can be 1977 separated psychoroniaPsychoronia larvae from hes separated based on the setae of the pronotum perophylax based on the lack of lateral gills P costalis has several evenly spaced setae beyond segment 3 in psychoroniaPsychoronia this char- along the anterior margin of the pronotum acter seems to be valid for P brooksi although see wiggins 1977 fig 104610.46 whereas these P brooksi larvae usually have the posterior lat- setae are absent in P brooksi eral gill branched I1 have also seen occasional 164 GREAT BASIN naturalist volume 59

colorado P costalis larvae with branched lat- larvae being present during emergence there- eral gills another character that appears to fore suggesting a univoltine life cycle for P separate PsychpsychoroniapsychoromaPsychooroniafomaromavoma larvae from hesperophy costalis lax is the reduced number of primary setae psyapsy along the anterior pronotal margin of additional RECORDS AND NOTES ON chchornoma choromachoroniaoronia in hesperophylax the primary setae WESTERN NORTH AMERICAN caddisfliesCADDIS FLIES are numerous and not evenly spaced PsychpsychoroniapsychoromaPsychooroniafomaroma brooksi is known only from the the following are additional distributional type collection which consisted of numerous records for western north american caddis larvae and pupae adults were reared from flies special recognition goes to the following pupae by placing the pupae in plastic bags institutionsindividualsinstitutionsinstitutionsindividualsindividuals for providing most of along with a piece of damp moss the bags these specimens colorado state university were transported in a cooler on iceieelee for about a CPCE gillette museum arthropodofarthropodof diversity week and then placed in a home refrigerator BC kondratieffcsukondratieff CSU university of wyoming adults started emerging in the refrigerator RJ lavigne UW purdue university A about a month later larvae were numerous provonsha PU brigham young university throughout all but the highest velocity habi- monte L bean life science museum RW tats of the north fork rio ruidoso this flow baumann BYU illinois natural history sur- ing water habitat along with the scraper vey KA methven INHS california academy mandibles of the larvae is unusual for most of sciences VE lee CAS dean blinn DB membelmemberss of the limnephihmlimnephilini but similar to GZ jacobi GZJ and SR moulton II11 SRM that of hesperophylax the stream had a maxi-maxi mum width of about 252.52 5 m at the time of col- apataniidae lection pupae were located in of in aggregations apataniaapalania shoshone banks 1924 ALASKA katmaigatmai 5 10 along to the lateral margins of the largest peninsula at lights alaknak river alaknak lodge boulders I1 could turn over in the stream just WG downs 22 august 19872m1987 2mam IF DER below the water substrate interface this is often the same type of habitat where hesperthespero apataniaapalania zonella zetterstedt 1840 WYOMING phylanphylax pupae are found this is not usually the albany county meadow creek at glacier lakes case for colorado P costalis pupae which are medicine bow national forest H copeland 29 often lying scattered throughout the vegeta- june 1987 IF west glacier lake medicine bow tion of headwater seepage area habitats national forest BC kondratieff & WB painter although they may be attached to rocks and 21 july 1987 IF glacier lakes T ebert 4 july 1988 CSU carbon county swept from sticks I1 have not seen P costalis in streams af3f herbage near edge of lake marie 20 km W of cent- larger than about 04m040.4 m wide they are most ennial PH arnaud jr I1 august 1973 IF CAS often found within 3 m of the headwater spring source there also seems to be some brachycentridae type of habitat partitioning between P costalis and hesperophylax whereas hesperophylax micrasema aniscaonisca ross 1947 ARIZONA coconino county front passage to roaring springs cave occidentalisoccidentalistalis banks is very common in high occident grand canyon national park D blinn 28 septem- altitude colorado headwater seepage areas in in ber 1994 IM IF DB those seepage areas which contain P costalis H occidentoccidentalisoccidentalistalis is found further downstream min glossosomatidae the outlet stream A single hesperophylax sp larva along with larvae of lepidostomaLepidostoma sp agapetus boulderensisboulderensis milne 1936 NEW MEX- ICO colfax county creek US 64 west of cammar and oligophlebodes sp was collected along cimmar ron M harris 8 august 1990 IF CSU with the P brooksi larvae lim 10lomm larval the presence of mm cases and 20 Glossoglossosomaglossocomasoma velonum ross 1938 YUKON TERRI- mm pupal cases in late may suggests a 2 in yr TORY stewart river at mayo JC abbott & KW life cycle for P brooksi in colorado P costalis stewart 24 june 1996 IF ross river at ganol emerges from late july through early septem- road 212.1 mi N of perry barge JC abbott & KW ber and I1 have not seen evidence of small stewart 25 june 1996 3mam af7f SRM 1999 NEW SPECIES IN GENUS psychoroniaPSYCHORONIA 165

frotoptilaprotoptila coloma ross 1941 WYOMING park hydroptila ajax ross 1938 california river- county fireholeFirehole river near old faithful BC side county colorado river route 95 BCB C kon kondratieff 8 june 1987 ism CSU teton dradratieffdratieff&dragiefftieff & RWR W baumann 20 june 1988 2mam CSU county gibbon river gibbon falls picnic ground this appears to be a significant western extension yellowstone national park RJR J lavigne 13 july of records for this species 1989 3mam UW hydroptila angusta ross 1938 WYOMING crook Goercoergoendaegoeridaeidae county belle fourche river hulett BCB C kon dratieffdradragiefftieff & 15 july 1997 Goergoereillaeilla baubaumanmmanni denning 1971 washing- dratieff&rwRW baumann IM CSU extends the distribution ofthisorthisof this ton spokane county small streams big spring this species further northwest picnic ground on mount spokane DED E ruiter 9 june 1996 IM DER hydroptila arctia ross 1938 NEW MEXICO hydropsychidaeHydropsychidae grant county west fork of gila river at road 15 gila national forest P mccafferty A provonsha ceratopsycheCeratopsyche protis ross 1938 ARIZONA santa & D bloodgood 6 may 1981 IM af5f PU cruz county santa rita lodge madera canyon PA opler 30 july 1991 2mam CSU WYOMING hydroptila conconsimilissimilis morton 1905 NORTH green sublette county river at highway 191 DAKOTA hettinger county cannonball river new KDK D alexander & LAL A stewart 4 august 1995 9mam england route 22 BCB C kondratieff & RWR W bau- af3f SRM mann 15 july 19974m1997 4mam af4f CSU

cheumatopsyche lasta ross 1938 COLORADO hydroptila pecos ross 1941 MONTANA bighorn las animasammas county purgatoirePurgatoire river iron canyon county bighorn river fort smith DED E ruiter 5 rourke ranch PCMS BCB C kondratieff 14 july september 1991 17m DER SOUTH DAKOTA fall 1991 13m IF otero county purgatoirePurgatoire river river county fall river hot springs BC kon- route 109 BC kondratieff 9 april 1992 2mam dratieffdratieff 15 july 1988 2mam IF 5 february 1995 2mam CSU IF CSU

Hydrohydropsychepsyche californicacahformcacalicailfornica banks 1899 NEW MEX- hydroptila salmo ross 1941 WYOMING carbon ICO county at light below catron taylor creek county medicine bow river about 2 mi E of elk wall gila national B C & lake forest BC kondratieff mountain on interstate 80 RJR J lavigne 23 august R durfee 24 july 1994 22m 15f dona ana 1982 5mam UW county radium springs HEH E evans 13 may 1989 9mam county grande percha state sierra rio dam Leucoleucotrichialeucotnchiatrichia pictpictipesipes banks 1911 SOUTH area B C kondratieff 17 july 1989 recreation BC DAKOTA fall river county fall river hot springs gom CSU af8f BCB C kondratieff 15 july 1988 lomIOM af4f 5 february 19956m1995 6mam IF CSU Hydrohydropsychepsyche occidentoccidentahsoccidentalisoccidentalistalisallsails banks 1900 NEVADA washoe county truckee river verdi fish hatch- ochrotnchiaochrotrichiaOchrotrichia alsea denning & blickle 1972 ery RW baumann 10 may 1983 IM BYU SOUTH california butte county diamond timber 252.52 5 DAKOTA bennett county 10 mi E of martin mi la mi E of hwybwy 52 and rd 150g PA opler & E creek national wildlife refuge PA opler 26 may bucknerbuekBuckneinel 9 july 19935m1993 5mam CSU 1990 IM IF fall river county hot springs BCB C kondratieff 15 july 1988 CSU lim ochrotnchiaochrotrichiaOchrotrichia logana ross 1941 ALASKA katmaigatmai peninsula at lights alaknak river alaknak lodge smicndeasmictidea atella mclachlan 1871 fasciatellafascifascianella califor- WG downs 22 august 1987 IM af2f DER this nia riverside county colorado river route 95 represents a major northwestern distribution exten BCB C kondratieff & RWR W baumann 20 june 1988 sionslon 6mam IF colorado river mayflower park blythe BCB C kondratieff & RWR W baumann 20 june 1988 ochrotnchiaochrotrichiaOchrotrichia stylitastylata ross 1938 NEW MEXICO 6mam 12f CSU grant county west fork of gila river at rd 15 gila provonsha hydroptilidaehydrophilidae national forest P mccafferty A & D bloodgood 6 may 1981 IM PU Agraylea salteseasaltesea ross 1938 WYOMING park county virginia cascades yellowstone national Ochroochrotrichiaochrotnchiatrichia tartarsalismarsalissalis hagen 1861 SOUTHSOUJTH park HRH R lawson 19 july 1989 3mam af2f CSU DAKOTA fall river county fall river hot springs 166 GREAT BASIN naturalist volume 59

BCB C kondratieff & RWR W baumann 5 february appear to be the southwesternmost records for this 19953m1995 3mam af4f CSU species

orthotrichiaOrthotrichia aegerfasciella chambers 1873 oecetisdecetis inconspicua walker 1852 NEW MEX- COLORADO larimer county black light trap mail ICO san miguel county pecos river near creek PA opler 18 august 1988 IF 10 august monastery lake GZCX jacobi 5 october 1980 IM 1989 2mam CSU this appears to be the western- CSU most collection for this species triaenodes tardusbardus milne 1934 california butte county 10 mi ESE ofchicoof chicochleo Centcentervilleerville PA opler oxyethira hallidapalhdapallida banks 1904 COLORADO 11 august 19932m1993 2mam IF CSU baca county east carrizo creek carrizo creek picnic ground BCB C kondratieff 16 july 1992 IM limnephilidaelimnophilidae larimerlarmier county light trap mail creek fort collins asynarchus circopa ross & merkley 1952 PA opler 29 august 1989 1mam black light trap circola WYOMING big horn county meadowlark lake shields pond off shields road BCB C kondratieff 5 bighorn national forest sanderson 24 july 1991 24m 15f black light trap shields pond MW august 1954 IM INHS fremont county golden fort collins BCB C kondratieff & R durfee 5 july lakes middle fork bull rees 6 1991 22m 30f montezuma county toten reser- lake creek D august 1996 2mam CSU teton county lewis voir east of cortez BCB C kondratieff 2 may 1992 lake yellowstone national park lavigne 31 july lomIOM af4f CSU RJ 1990 IM UW Zumzumatnchiazumatrichiaatrichia notosanonosa ross 1944 NEW MEXICO asynarchus montanus banks 1907 herrmann grant county west folkfoikfork of gila river at rd 15 et al 1986 did not report this from gila national forest P mccafferty A provonsha species colorado doubting the record of schmid 1955 1 & D bloodgood 6 may 19814m1981 4mam af2f PU I have now seen specimens from the west side of rocky lepidostomatidae mountain national park COLORADO grand county 121.2 mi S of boksubowsu baker trailhead rocky moun- lepidostomaleptdostomaLepidostoma amomumapomumapornum denning 1949 ARIZONA tain national park PA opler 23 july 1994 1mam coconino county front passage to roaring springs harbison picnic area rocky mountain national cave grand canyon national park D blinn 28 park PA opler & E buckner 29 july 1995 IM september 1994 IM1mam DB COLORADO larimer green mountain employee area rocky mountain county upper beaver meadows picnic area rocky national park PA opler 30 august 1997 IM park mountain national PA opler 12 july 1990 CSU OREGON clatsop county black light trap am 2m CSU laramie river at honholtzHonhongoltzholtz lakes astoria K goeden I1 september 1969 IM CAS access DED E ruiter 15 july 1988 9mam af3f DER asynarchus mutatismutatus hagen 1861 UTAH du- Lepidoleptdostomalepidostomastoma cascacascadensedense milne 1936 NEW chesne county white rocks river below chepetachapeta MEXICO taos county red river at zwergel dam lake ashley national forest RC mower 20 july G Z jacobi am GZ 29 july 1980 1m GZJ 1984 IM summit county china meadows uinta mountains RW baumann & BJ sargent 14 july leptocendaeleptoceridae 1986 2mam BYU these collections represent a sig- ceracleaCeraclea annuliannulicomisannuhcomiscomis stephens 1836 WYOMING nificantnificant southern distributional extension for this carbongalbon county big creek 3 mi above confluence species it seems that the uinta mountain area of with north platte river D rees 27 june 1988 3mam utah contains many interesting distributional records af3f CSU park county slough creek camp- see discussion under hydatophylax hesperus ground yellowstone national park RJ lavigne 2 august 19902m1990 2mam af2f UW chyrandaChyranda cencentraliscentralestralis banks 1900 NEVADA elko county stream above angel lake ruby moun- decetisoecetis cinerascens hagen 1861 COLORADO tains RW baumann 3 august 1990 IM IF BYU baca county picture canyon BCB C kondratieff 15 june 1994 1mam CSU clistoroniaClistoronia flavicollisflavicollis banks 1900 washington chelan county minotaur creek 10 mi W of decetisoecetis mobilisimimmobilisimmobihs hagen 1861 COLORADO natcheewenatcheeWe lake site 6 in uncut timber north saguache county at lights russell lakes state and south ofofrdoardrd 2728 JR wood 14 july 1976 1mam1 M wildlife area R durfee 6 july 1994 3mam 17 july DER this appears to be the most southern 1994 IM af3f 8 august 1994 2mam af8f CSU these record for this species 199919991 NEW SPECIES IN GENUS psychoroniaPSYCHORONIA 167

glyphopsycheclyphopsyche irroratairrorateirrorata fabricius washington polycentropus crassicorniscrassicormscrassicornis walker 1852 MON- kittatusKittatus county teanawayTeanaway river hwybwy 10 4 mi S of TANA rosebud county coiColcolstnpcolstripcolstrupstrip quicktrapquicktrap TF r2ra cle elum RW baumann & SD smith 6 may q33 leetham 20 june 1975 IM DER 1982 IF BYU psychomyiaPsychomyia flaridaflavida hagen 1861 ARIZONA hydatophylax hesperus banks 1914 UTAH coconino county oak creek along USU S hwybwy 89a wasatch county bryantsabryants fork creek strawberry north of sedona SRS R moulton 12 june 1998 2mam reservoir M whiting S wells & L liu 14 june af2f SRM 1989 1mam BYU this record is based on a 1989 collection subsequent to that collection the entire rhyacophilidae drainage including the headwater springs was rhyacophila narvae navas 1926 WYOMING park roterotenonednoned in an attempt to restore a nonnativenormative county virginia cascades yellowstone national sport fishery in a downstream reservoir the bryantsabryants park HRH R lawson 19 july 1989 IM IF CSU fork creek locality was revisited in 1995 and 1996 rhyacophila kellisapellisa ross 1938 NEW MEXICO and no caddiscaddisfliesflies were located in an effort to cre taos county red river at zwergel dam GZG Z ate maintain a nonnative sport fishery it is likely jacobi 29 july 1980 IM CZJGZJ that isolated native aquatic species within this drainage such as hydatophylax hespersushespersus have been uenoidaeUeno idae eliminated Neophyneophylaxlax nckenricceririckeri milne 1935 COLORADO jack- son county black light trap north platte river at limnephilus moestusmodestus banks 1908 NEW MEXICO ginger quill ranch WG downs I1 september santa county medio at santa fe creek fe ranch 1990 IF DER this appears to be a major south- ski alexander & stewart 5 july lodge KD LA eastern distributional extension for this species 1995 IM SRM ohgophlebodesofigophlebodes minutesminutus banks 1897 YUKON limnephilus sansoni banks 1918 ruiter 1995 TERRITORY little rancheriaRanchena creek at alaska high- indicated that the colorado record of dodds and way JCJ C abbott & KWK W stewart 26 june 1996 IM hisaw 1925 was questionable and did not include SRM limnephilus sansoni from colorado I1 have now seen specimens from grand county colorado on acknowledgments the opposite western side of the continental divide from the locality reported by dodds and in addition to the individuals and institu- hisaw grand county green mountain employee tions mentioned above who supplied much of area park rocky mountain national PAEA opler 30 the material that this paper is based on I1 august 1997 6mam af2f CSU would like to thank both john morse and an anonymous reviewer for providing very help- psychoglypha prita milne 1935 record for the ful comments and for this paper I1 ross corrections psychoglypha onniaeonnive 1938 from teton county also thank the museum of comparative zool- wyoming ruiter and lavigne 1985 is an error ogy for the loan of the type specimens of P these specimens are psychoglypha prita milne brevipennisbrempenmsbreulbrevipennis and P costalis C lynn bjork drew 1935 teton county taggert creek 7000 feet HE the larval figures evans 6 october 1983 25m CSU

CITED psychoglypha schuhlschubl denning 1970 WYOMING literature sublette county 25 lead creek DE ruiter octo- BANKS N 1901 some am insects of the hudsonianHudsoman zone of ber 1995 3m DER this appears to be only the new mexico neuropteroid insects psyche 92869 286 287 and2nd collection reported for this species type local- 1904 neuropteroid insects from new mexico ity is in nevada transactions of the american entomological society 309730 97 110 polycentropodidae 1916 A classification of ouioulour limnephilidlimnophilid caddice hiesflies canadian entomologist 4811748 117 122 cornelluscyrnellusCyrnellus frafraternusternus banks 1905 COLORADO DODDS GSG S AND FS HISAW 1925 ecological studies on bent county caddoacaddoan creek at road CC BC aquatic insects IV altitudinal range and zonation of kondratieff 14 july 1992 IF black light trap john maymayfliesflies stonestonefliesflies and caddisfliescaddis flies in the colorado rockies ecology 63806 380 390 martin reservoir BC kondratieff 16 july 1992 prowers HERRMANN SJS J DED E RUITER AND RJR J LAVIGNE 1986 3mam county arkansas river US 50 lamar distribution and records of colorado trichoptera BC kondratieff 23 august 1996 IM CSU southwestern naturalist 3142131 421457457 168 GREAT BASIN naturalist volume 59 runnerullerRuflErunnellRUIILRii DE 1995 the adult limnephilus leach trltri- partie 7 agricultureAgnculture canada publication 1692 296 chopterachop terateya limnephilidaelimnophilidae of the new world bul- PP letin of the ohio biological survey new series WIGGINS GBG B 1975 contribution to the systematics of iliiii11l1ilil1111111 1 200 the caddisflycaddiscaddishlyfly family limnephilidaelimnephihdaelimnophilidae trichoptera II11 rullerrulRUIILRRUIteitteltTEti DE aniAND RJ lavigneLAVICNE 1985 distribution of canadian entomologist 107325107 325 336 wyoming trichoptera university of wyoming agri- 1977 larvae of the north american caddiscaddisflycaddishlyfly culturaleuitcultuialulalulai experiment station publication sm47 102 genera trichoptera university of toronto press PP 401 appp S hmilIIMID F 1955 contribution a 1 etude des limnophili- dae trichopteraa mitteilungen der schweizerischenschweizenschenSchweizerischen received 20 march 1998 entomologischen Gesellgesellschaftgesellsehaftschaft 28128 1 245 accepted 26 june 1998 1980 genera des tnchopterestrichopt6res du canada et des ttatsthatsetats adjacentsadjacents les insectinsectesinsectedes et arachnitesarachnidesarach nides du canada great basin naturalist 592 199901999 appp 169 174 EMERGENCE PATTERNS OF LARGE stonefliesSTONE FLIES plecoptera pteronarcys calineuriaCALIN EURIA hesperoperla IN A MONTANA RIVER

andiewandrew L sheidonsheiSheldonsheldonlsheldon1sheldonaSheldonldoni1

ABSTRACT Emeremergentgents of pteronarcys californicacalifornica calineunacalineuriaCaliCalineurlaeurianeuna calicailcalifornicacaliformcacaliforfornicamcamoa and hesperoperla pacifica were captured with leplicatedreplicated emergence traps at 3 sites along the banks of rock creekgreek montana pteronarcys emergence in early june was short 88 of individuals in 6 d and attained latesrates of 5 m 1 d 1 shoreline distance and cumulative densities up to 19 m 1 yr 1 caltcalineunacalineuriaCaliCalinentiaentlaeurianennaneuna emerged synchronously 89 in 9 d 2 wk later at densities up to 7 m 1 d 1 and 40 mnrr11 yr 1 the less numerous hesperoperla 5 individual m 1 yr 1 had a longer emergence period partially coinciding with calineunacalineuriacalzcaizCaliCalineurlaeurianeuna median males of pteronarcys and caltcalineunacalineuriaCaliCalineuriahuriahurlaneuna emerged 2 3 d before median females protandrypiotplotandry was not sig- nificantnificant in hesperopeihesperoperla la sex ratios were female biased in pteronarcys and hesperoperla but male biased in caltcalineunacalineuriaCaliCalineurlaeurianeuna size trends through emergence were very weak although females showed a greater tendency toward larger size early in emergence

key words plecoptera phenology emergence traps sex ratio size

emergence the transition from the aquatic 1995 but aside from investigations of drum- growth phase to the reproductive and disper- ming behavior stewart and laketonmaketon 1991 sal activities of terrestrialaerialterrestrial aerial adulthood is a and calineuriaCalineuria flight behavior poulton and critical event for aquatic insects macan 1958 stewart 1988 little has been written about corbet 1964 zwick 1990 for the biologist adults since muttkowski 1929 described collection of emeremergentsgents provides a census of emergence of pteronarcys and hesperoperla in readily identified individuals at a defined point the yellowstone river in the life history thus emergence data are emergence of stonestonefliesflies has been described essential for life history studies and are poten- from sweepnetsweepnet catches sheldon and jewett tially useful for monitoring stattnerStatzstatznerner and resh 1967 that confound adult longevity with 1993 synthesized a large set of emergence emergence timing counts of exuviae haro et studies in an analysis of species richness and al 1994 alexander and stewart 1996 and other properties of stream communities this captures in traps of various designs kerst and paper however emphasizes population biology anderson 1974 zwick 1977 kuusela 1984 in the western united states emergenciesemergences of ernst and stewart 1985 bagge and hynynen the stonestonefliesstoncfliesflies pteronarcys californicacalifornica newport 1995 giberson and garnett 1996 I1 used pteronarcyidae calineuriaCalineuria californicacalifornica banks catches from emergence traps designed for and hesperoperla pacifica banks perlidae this project to describe phenology abun- are notable for the numbers and visibility of dance and size and sex composition of adult the large colorful diurnally active adults to pteronarcys calineuriaCalineuria and hesperoperla in avoid confusion involving the californicacalifornica epi- rock creek montana thet I1 refer to these species by their generic names throughout this paper As salmonfliessalmonflies METHODS pteronarcys and golden stonestonefliesflies caline uria hesperoperlahesperoper1a these insects are impor- rock creek a noted trout stream originates tant in angling lore and literature leiser and in the anaconda range of western montana boyle 1982 the nymphal biology of these and flows north 110 km to its confluence with taxa is moderately well known stewart and the clark fork river 4643n46043n IISWW stark 1988 freilich 1991 dewalt and stewart three sampling stations were established valeyvalleyvaileymaley

I1 division of biological sciences university of montana missoula mt59812MT 59812

169 170 GREAT BASIN naturalist volume 59

10 cm diameter

fig 1 emergence trap used in this study arrow indicates direction ofwaterof water flow along the shoreline

of the moon stream km 404.04 0 1095 m elevation anchored the entire trap an informational sign ranch creek 20020.020900goo 0 km 1207 m bitterroot flat was placed by each trap traps were tended 38038.038 0 km 1305 m daily and moved up or down to accommodate the traps used fig 1 were designed to changing water levels Stonestonefliesflies were col- intercept emerging stonestonefliesflies as they left the lected from the cylinder and also from the water cost was kept to a minimum since I1 used apron and outer surface of the funnel since numerous replicates at each site and planned especially on cold mornings teneteneralsgeneralsrals and pre to sample at more sites I1 also anticipated sub- ecdysial nymphs sometimes failed to pass stantial trap loss from changing water levels through the funnel before the traps were and human interference traps were con- tended nymphs were included in numbers strucstruestructedted of plastic window screen PVC pipe and sex ratios but not in size data and duct tape each trap consisted of an apron at each site 20 traps were placed at 152 m leading upward to the PVC pipe which con- soft50 ft intervals along one bank since the tained a screen funnel past the PVC collar opposite shore was inaccessible at spring dis- insects were captured in a stapled screen charge identification sex determination and cylinder closed by a heavy rubber band which size measurement head width including eyes allowed easy access apron funnel and cylin- with dissecting microscope and ocular micro- der were attached to the PVC pipe with duct meter were done in the laboratory data pre- tape total cost was 200 per trap more sented here are aggregated by site a statistical refined models with apron and cylinder treatment of betweenofbetween trap variability and pre- attached by hose clamps and the funnel with cision of this and other emergence studies is epoxy resin weiewelewere nearly twice as costly how- in preparation ever duct tape was completely satisfactory preliminary fieldwork in 1989 using less except when the entire trap was submerged retentive traps without funnels began 5 june for hours midway in the pteronarcys emergence and in use the edge of the apron was placed at ended 13 june in 1990 I1 tended traps daily 29 or just below the water s edge and weighted may 30 june this is a period of very heavy with gravel and small stones if needed a use by anglers so I1 chose not to install record- large stone or two held the PVC pipe and a ing thermometers although temperature data stone at the rubber band end of the cylinder would have been very useful 1999 STONEFLY EMERGENCE 171

statistical methods follow sokal and rohlf 8 1981 and zar 1984 calineuriaCalineuriaeurla

RESULTS 6 pteronarcys

emergence of these species occurs during E peak spring runoff from snowbeltsnowsnowmeltmelt because 0 4 high discharge covers rocks exposed at other E seasons and limits emergence to the shoreline z at the study sites the entire emergent popula- 2 tion was vulnerable to the traps however hesperoperla high and variable discharge washed 1 away many goor A b 0 0 q 0 traps on the gently sloping shore at lower- 0 0 0.0 the 0 7 most valley of the moon site in both years 28 30 1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 thus the quantitative data from that site can- may june not be used one trap at ranch creek and fig 2 emergence number m 1 d 1 at ranch creek the 3 over season at bitterroot flat were lost 1990 length is shoreline length double these number to to flow and tampering analyses are based on estimate number per stream length data from traps persisting to the end of emergence for a particular species ie n 19 for pteronarcys and n 17 for calineuriaCalineuria and ranchbanch cr 1990 hesperoperla at bitterroot flat 5 x bitterrootbifterroot flat 1990 1990 few pteronarcys in very adults were 13 ranchbanch cr 1989 present at ranch creek when traps were 4 installed 28 may snow and low water temper- E ature inhibited emergence over the next few 3 E days and again on 5 2 june fig however Z3 2- pteronarcys emergence was extremely syn- chronouschronous with 88 of adults emerging over 6 consecutive days the first hesperoperla adults emerged with pteronarcys but the 29 31 1 2 3 6 7 8 10 11 12 majority were contemporaneous with the abun- may june dant calineuriaCalineuria pteronarcys and calineuriaCalineuria peaks were separated by a week with little fig 3 emergence of pteronarcys at 2 sites in 1990 and 1989 data 6 activity although calineuriaCalineuria emergence was at ranch creek in no before june length measure is shoreline length protracted relative to pteronarcys 89 emerged during 9 consecutive days emergence timing varied within and be- no exuviae were found at bitterroot flat in tween years fig 3 pteronarcys emergence 1989 anglers and I1 observed massive emer- in 1990 at the upper 2 sites was in phase and gence all along the lower 80 km of rock creek controlled by cold weather in late may and on on 5 june 5 june not shown in figure 3 are 3 excep- densities expressed as emergentsemergents per tiotionallynally late pteronarcys emeremergentsgents at bitter- meter of shoreline estimated densities per root flat males on 16 18 june female 24 meter of stream length will be twice these val- june emergence at ranch creek in 1989 was ues appear high although few data exist for slightly later than in 1990 anglers reported comparison at ranch creek pteronarcys and the first heavy emergence on 5 june spatial calineuriaCalineuria attained peak emergenciesemergences of 5 7 variation in timing was not adequately sam- nr1mnry 1 d 1 cumulatives for ranch creek were pled with complete data from 2 sites only pteronarcys 19619.6 nr 1 yliyilyriylyr 1 calineuriaCalineuria 40040.0 mmr1mra1 anglers rely on a steady upstream progression yr 1 and hesperoperla 535.3 wrim 1 yr 1 incom- of the hatch and this is readily apparent in plete data for 1989 fig 3 yielded 13913.9 some years in 1990 numerous adults were pteronarcys m 1 yllyliyiyr 1 of similar magnitude to present on 28 may at the lower site valley 1990 catches at bitterroot flat were consid- of the moon a few were at ranch creek and erably less pteronarcys 606.0go m 1 yliyriyr 1 calineutiacalineuriaCalineurlaeuriaeuTia 172 GREAT BASIN naturalist volume 59

767.67 6 m 1 yr 1 and hesperoperlahesperoperiaHesper operia 7777.77 m 1 yr 1 TABLE 1 sex composition by species and locations an interesting perspective is obtained by cal- species location male female cuculating that I1 adult of the combined species pteronarcyspteronareys ranch 93 121 emerged in 1.5lsis1 5 cm of shoreline in 15 at ranch creekci eek bitterroot 25 34 and in 4744.77 cm at bittenbitterrootoot flat determination of sex ratios and sexual phe- caimcalmeunacalineuriaCalinCalmeuriaeurlaeufiahunaeuna ranch 175 120 bitterroot 37 37 nology of emeremergentsgents is a necessary first step toward understanding operational sex ratios hesperoperiahesperoperlaHesperoperia ranch 16 29 and mating tactics stewart 1994 sex ratios bitterroot 24 49 table 1 were female biased in pteronarcys at both sites although not significantly so xax22 test ratios at the 2 sites are not different P 0990.990ogg99 nor do the combined data depart signifi- divided at medians of size and time A significantly from aallali11 sex ratio P 0100.10olo0 10 sex ratios cant yax2xay2 indicates that size and emergence time of hesperoperlahesperoperiaHesperoperia were female biased signifi- are not independent and the phi coefficient cantly so for the larger collection not different a parametricnonparametricnon correlation coefficient with P 0990.990ogg99 at the 2 sites and significantly P range 1 to 1 measures the strength and 00050.0050 005oos biased in the combined data calineunacalineuriaCaliCalineuriaeurlaneuna direction of the interaction size time interac- in contrast was strongly male biased at ranch tions table 2 were weak and only 3 of 11 creek not different P 0100.10olo0 10 at the 2 sites tests were significant seven of 11 phi clefficoeffi and significantly P 00050.0050 005oos male biased in elenis were negative ie later individuals were combination these results suggest very differ- smaller including all the significant cases ent mating systems especially when compar- effects on females were stronger than males in ing the 2 perlidae all species and locations the collective result operational sex ratios also depend on sex indicates little or no size change through the specific emergence phenology and on the relatively short emergence periods of these functional life span especially of males which species if fitness attributes such as fecundity probably can mate more than once longevity and mating success are size dependent qual- was not measured in this study differential ity of potential mates varies little over time emergence by sex table 2 illustrates the pro- this conclusion may depend in part on the tantandrousdrous pattern seen in many insects median conservative contingency test used blackburn males of pteronarcys and caltcalineunacalineuriaCaliCalineuriaeufianeuna emerged et Aal 1993 showed that in sampling from 2 3 d before median females differences in skewed size distributions size and abundance distributions of emergence times of the sexes were confounded to avoid this effect I1 used a were tested with kolmogorov smirnov tests parametricnonparametricnon test applying the same test to 2 tailed contrasts involving pteronarcys and comparable data sheldon 1972 on male caltcalineutiacalineunaCaliCalineuTianeuna were significant except for pteronar sawalaskwala curvatecurvata hanson perlodidae yields cys at bitterroot flat where sample size was phi 0480.48 P 00050.005 a much stronger size small and where late emeremergentsgents mentioned time interaction than observed in rock creek above included males the pattern for hes stonestonefliesflies although conservative the tests can petpereeroperoperlaperoperiapetopero perlaperiaoperia was quite different median emer- detect temporal size patterns where they exist gence dates were very similar especially in natural history observations on post emer view of the extended emergence of this species gents include the following adults are readily fig 2 and distributions of emergence times apparent on bushes where mating occurs by sex were not significantly different oviposition flights occurring as air tempera- attributes other than sex may vary through tures rise appear to be directed upstream the emergence period A tendency for early and many flying adults of pteronarcys were erneiernelemergentsemeremmel gents to be larger than later ones has observed several kilometers upstream from been reported in stonestonefliesflies khoo 1964 shel- reaches where from exuvial density and num- don 1972 and other insects examination of ber of adults on bushes significant emergence size vs time plots suggested that such patterns had occurred supporting muttkowski s 1929 were weak or absent in these species formal observations on predation I1 saw numerous tests and metrics were based on 2 X 2 tables birds especially flocks of 30 40 western 199911999 STONEFLY EMERGENCE 173

TABLE 2 emergence timing and size trends by sex M F and location median dates alediedleare all in june 1990 and proba- bilitiesbilities are fromblom kolmogorov smirnov contrasts of temporal distributions ofofemeigenceemergence of the 2 sexes Conecorrelationslations phi are negative if later emeemergersrgers are smallersmailersmalleilerier probabilities flomfrom x2xa2 median date size time M size time F species site M F P phi P phi P pteronarcys ranch 3 5 0001 002 0750 75 005 050 bitterrootBittenoot 4 6 005 004 0750 75 026 010 cabcahneunacalineuriaCahCalineurlaeuriaeuylaneuna ranch 17 20 00010.0010 oolooi001 017 0025 030 0005 bitterroot 15 18 00010.0010 ooi001 017 075 013 0250 25 hesperoperla ranch 10 11 0400 40 014 0250250.250 25 bitterroot 13 13 090 oil011 0500500.500 50 032 005

tanagersTamanagersnagers piranga ludoludovicianaviciana foraging in used here may have failed to detect lealleaireal phe- riparian vegetation and hawking at flying nomena however blackburn et al 1993 adults particularly pteronarcys have shown that skewed size distributions and temporally varying collection sizes as in most discussion emergence data can produce statistically sig- nificantnificant artifacts with parametric tests the simple emergence traps worked well at repeatability of observations is an impor- the 2 sites where stream banks were steep tant issue emergence biology of pteronarcys level did enough that a moderate rise in water in rock creek is very similar to the same not submerge the entire trap daily mainte- species in colorado dewalt and stewart to accommodate fluctuat- nance was necessary 1995 the differences in numbers and species ing discharge and to yield temporally precise proportions at ranch creek and bitterroot emergence data aggregated over longer data flat may indicate a longitudinal trend but I1 periods can be very useful masteller 1983 attribute them to differences in poolrunrifflepoolpooi run riffle kuusela 1984 but miss the details of short proportions and other physical factors phe- synchronous emergences emergencies nology and sex ratios were similar at the 2 sites although common elements appear each of haro et al 1994 detected substantial varia-variavarla the 3 species had unique elements in its emer- tion of and sex ratio of a perlodidperlodid stonstoneflyefly gence phenology pteronarcys and calineuriaCalineuria size among sites and years although intraintraspecificallyspecifically synchronous were emergence a single event a life history temporally well separated whereas the less is in emphasis in this is on timing identity abundant hesperoperla had a longer less syn- in paper and sex of emergentsgents more generallyallyaily chrchronousonous emergence overlapping the other size emer geneigenel of the two synchrony has been suggested as a tactic emeremergentsgents are the product aquatic life to the adult for satiating predators sweeney and vannote phase of the history and input 1982 perhaps hesperoperla benefits from population emergence data could be a sensi- emerging with abundant calineuriaCalineuria tive indicator of nymphal population biology patterns of size and sex ratio also vary be- and environmental conditions between years tween species A general tendency toward and locations haro et al 1994 of equal inter- protandrous emergence in insects is supported est the numbers and quality sex size of emer in pteronarcys and calineuriaCalineuria species with gents must influence mating behavior and re- short synchronous emergence periods the productive success detailed studies of stonstoneflyefly selective advantage fagerstrom and wiklund reproduction alexander and stewart 1996 1982 of protandry apparently is reduced in would be enhanced by a firm comparative the less common hesperoperla with its pro- base of emergence data such data are rela- tracted emergence size time interactions were tively easy to obtain and for pteronarcys and weak although females showed a greater ten- the perlissperlids of rock creek provide informa- dency than males for early emeremergentsgents to be tion at the time of high discharge when the larger the conservative parametricnonparametricnon statistics stream itself is unworkable 174 GREAT BASIN naturalist volume 59

acknowledgments LEISER E AND RHR H BOYLE 1982 Stonestonefhesstonefliesflies for the angler alfred E knopf inc new york 174 appp MACAN 1958 causes this research was supported by a grant frombrom TT and effects of short emergence periods in insects verhandlungen der Internationintemationaleninternationalenalenaien the rock creek advisory council and the vereinigung fur theoretheoretischetische und angewandte lim montana department of state lands cologienologie 1384513 845 849 MASTELLER ECE 1983 emergence phenology of plecop- literature CITED tera from sixmile creek erie county pennsylvania USA aquatic insects 515 1 8 muttkowski RAR A 1929 the ecology of trout streams in ALEXANDER KDD AND KWW STEWART 1996 description yellowstone national park roosevelt wild annals and theoretical considerations of mate finding and life 21512 151 240 other adult behaviors in a colorado population of bisi POULTON AND W STEWART 1988 aspects flight ciaclaasseniaclaassemaClaClaasasseniasema sabulosesabulosasabulosa plecoptera perlidae annals of BC KW of behavior in cabcahneunacalineuriaCahCalinneuna cahcabcalicallcailpornica plecoptera perli- the entomological society of america 892908999089 290 296 in euriaeufia californicacahfornicafornica dae from a rocky mountain lake outlet system bacceBAGGEBACGL P AND J HYNYNEN 1995 plecopteran communi- entomological news 9912599 125 133 ties and annual emergence inm five forest streams and SHELDON A L 1972 comparative ecology ofarcynopteryx two lake outlet streams of central finland entomoantomo AL and plecoptera in a california stream archivarchev lodicalogica fennicafenmcabennica 6996 99 108 ciuradiura in fur hydrobiologie 6952169 521 546 BLACKBURN TMM KJ castonGASIONGASTON ANDNEAND NE STORK 1993 SHELDON ALA L AND SCS G JEWETT JR 1967 stoneflyStonefly emer- temporal dynamics of body size of beetles on oaks a gence in a sierra nevada stream pan pacific ento- cautionary tale ecological entomology 1839918 399 401 in momologistlogist 43143 1 8 corbelCORBEICORBET PS 1964 Tempotemporalialralrai patternspatteins of emeigenceemergence in SOKAL RRR R AND ROHLF 1981 biometry WH aquatic insects canadian entomologist 9626496 264 279 ejFJ rohlerohlk free- man and company san francisco 858 appp DEWAFTDEWALT RER E AND KWW STEWART 1995 life histories of STATZNER B AND VH RESH 1993 multiple and year stonestonefliesflies plecoptera min the rio conejos of south- site analyses of stream insect emergence a test of eco- ern Coloiacoloradodo great basin naturalist 55155 1 18 logical theory oecologia 966596 65 79 ERNST MRM R AND KWW STEWART 1985 emergence pat- STEWART W 1994 considerations mate terns and an assessment of collecting methods for KW theoretical of finding and other adult behaviors of plecoptera adult stonefliesstoneflies plecoptera in an ozark foothills aquatic insects 169516 95 104 stream canadian journal of zoology 63296263 2962 2968 STEWART KWW AND M MAKETON 1991 structures used fagerstrom T AND C WIKLUND 1982 why do males by nearctic stonestonefliesflies plecoptera for drumming emerge before females protandry as a mating strat- and their relationship to behavioral pattern diversity egy in male and female butterflies oecologia 52 13 33 164 166 aquatic insects 1333 53 STEWART KW AND BP STARK 1988 nymphs of FREILICH J E 1991 movement patterns and ecology of north JE stonstoneflyefly genera plecoptera entomologi- pteronarcys nymphs plecoptera observations of american cal society of america thomas say foundation 12 marked individuals in a rocky mountain stream 1 460 freshwater biology 2537925 379 394 SWEENEY BWW AND RLR L VANNOTE 1982 population syn- albersonGIBFRSON D J AND HLH L carnettgarneriGARNETIGARNETT 1996 species compo- DJ chrony in maymayfliesflies a predator hypothesis sition distribution and summer emergence phenol- in satiation evolution 36 810 821 ogy of stonestonefliesflies plecoptera from catamaran brook 36810sioslo ZAR J H 1984 Biostatistical analysis prentice hall inc new brunswickBlunswickswiek canadian journal of zoology 74 JH englewood 718 1260 1267 cliffs NJ appp ZWICK P 1977 pleopteren emergentemergenz zweier lunzer HARO RJR J K EDLEY AND MJM J WILEY 1994 body size and size bache 1972 1974 archivarchev fur hydrobiologie 80 sex ratio in emergentemeigent stonstoneflyefly nymphs Isogenisogenoidesoides 458 505 ohvaceusolivaceusolivaolivaceousceus perlodidae variation between cohorts 1990 emergence maturation and upstream and populations canadian journal of zoology 72 ovi- flights from 1371 1375 position of plecoptera the breitenbach with notes on the adult phase as a possible control kerseKLRSF CDD AND NHN H ANDERSON 1974 emergence pat- of stream insect populations hydrobiologia 194 terns of plecoptera in a stream in olegonoregon USA in in 207 223 freshwater biology 42054 205 212 KHOO SGS 1964 studies on the biology of capniaacapniacapma bifrontbifrons received 9 october 1997 newman and notes on the diapause in the nymphs accepted 14 july 1998 of this species Gegelwassergewasserwasser und abwasserabwdsserAbwasser 3435233435 23 30 kuuselaKUUSLLA K 1984 emergence of plecoptera in two lotic habitats in the oulankasulanka national pariparkpaikpalk northeastern finland annales de Limnlimnologielimnologicologie 206320 63 68 gleatgreat basin naturalist 592 019991999 appp 175 181

competition AND NICHE partitioning AMONG PSEudoroegneriapseudoroegneria SPICATA HEDYSARUM BOREALE AND CENTAUREA MACULOSA

james S jacobsijacobbijacobs1 and roger L sbeleylsheley1

ABSTRACT maximizing desired plant diversity has been suggested as a means of minimizing non indigenous plant invasion on rangeland by maximizing niche occupation competition between 2 desired indigenous species pseudoroegfseudoroeg nerianerlanena spicatespicata pursh love bluebunchbluebunch wheatwheatgrassgrass and hedysarum boreale nutt vaivar boreale northernnortheithel n sweetsweetvetchvetch and a non indigenous invader centaurea macumaculosemaculosalosa lam spotted knapweed was quantified using growthglowth of isolated individuals and 2 three species addition seriesserles experiments seeding densities of P spicatespicata remained constant at 0 200 400 and 800 seeds m 2 in both experiments H boreale and C macumaculosemaculosalosa seeding densities were 0 200 400 and 800 2 seeds m 2 respectivelyi in the ist experiment and 0 400 800 and 1600 seeds m2ma respectively in the and2nd experiment densities weiewelewere facfactoriallytonally arranged pots were placed in an environmental chambelchamber 12c 12 h day length 200 armol photons m 2 s 1 spectralspeetspectiallallai light in a randomized complete block design after 90 d the growth rate ofeof P spicatespicata 99192192 1 mg d 1 shoot growth was greater than that of the 2 forbs iglg161.61 6 and 555.55ss5 mg d 1 for H boreale and C macumaculosemaculosalosa respectively and growth rates of the 2 forbs were similar to one another curvilinear i egressionregression indicated that intraspecific competition was moiemotemore important in determining shoot weight than intelinterspecificspecific competition in addition the 2 forbs competed more directly with each other than with P spicatespicata competition coefficient ratios 1421421.421 42 and 1531 53 for P spicatespicata with H boreale and C macumaculosamaculoselosa respectively and 1031.031 03 for H boreale with C macumaculosamaculoselosa indicated substantial partitioningpaipal titionmg of lesourcesresources between P spicatespicata and each of the forbs little or no resource partitioning occurred between forbs this study suggests that increasing desired plant diversity may minimize weed invasion by increasing niche occupation

kelfkeifkey words plant diversity three species addition series competitive relationships resource preemption niche parti- tiotioningning spotted knapweed centaurea macumaculosamaculoselosa pseudoroegnenapseudoroegneria spispicatespicatacata hedysarum boreale

the ecological integrity and environmental it has been hypothesized that longtermlong term quality of grassland and sagebrushsagebrushgrasslandgrassland sustainable management of ecosystems under ecosystems of western north america are invasion by exotic species must focus on pro- being degraded by non indigenous invasive moting or establishing desirable species with plants these ecosystems once dominated by plant traits that maximize niche occupation species such as artemisia tridentata nutt big larson et al 1994 sheley et al 1996 many sagebrush pseudoroegneria spicataspicate blue aggressive exotic invaders appear to be able to bunch wheatwheatgrassgrass festuca abrellascarrellascscabrella torr occupy most niches in plant communities they rough fescue and festuca idahoensis elmer invade for example sheley and larson 1996 idaho fescue now contain extensive areas suggested that centaurea diffusediffusa lam diffuse dominated by non indigenous weeds such as knapweed monomonotypicallytypically dominates range- centaurea solstitiasolstitialislis L yellow starstarthistlethistle land by maximizing niche occupation through maddox 1983 bromus tectoriumtectorumtectorum L cheatcheatgrassgrass developing a hierarchy of age classes within mack 1981 euphorbia esula L leafy spurge the population in native communities estab- lajeunesse et al 1997 and centaurea macu lishmentlishment of desirable species having diverse losa lam spotted knapweed sheley et al above and belowgroundbelowground growth forms may 1998 it is widely accepted that invasion of minimize interspecific competition maximize rangeland by these species reduces resource vertical and temporal structure and enhance values including species diversity forage pro- their resource capture pyke and archer 1991 duction wildlife habitat and ecosystem func- tilman 1996 maximizing vertical and temporal tion lacey et al 1989 olson and lacey 1990 resource capture by increasing the diversity of thompson 1996 desirable species may limit exotic invasion

idepartmentepclitment of plant soil and environmental sciences montana state university bozeman MT 59717312059717 3120

175 176 GREAT BASIN naturalist volume 59 through resource preemption sheley and equilibrate to pot capacity ten seeds were larson 1995 broadcast on the soil surface and covered with ouiour objective was to develop an initial less than 2 mm of dry soil the soil surface understanding of the effect of combining was kept moist for the ist wk of the study by desirable plant species with contrasting above fog misting and then covering with clear plas- and belowgroundbelowground growth forms on their inter- tic no additional watering took place actions with C macumaculosemaculosalosa because C macu one week after emergence plants were losa is a taprooted forb we hypothesized that thinned to I1 plant per pot each species had 6 the addition of a desirable taprooted forb in a harvest dates at 14 d intervals beginning 14 d grass population would increase interspecific after thinning there were 4 replications at competition by maximizing niche occupation each harvest date pots were arranged in an and reduce C macumaculosemaculosalosa growth we compared environmental chamber 12c 12 h day length growth rates and competitive interactions 200 emoltmolumol photons mnr2nry2 s 1 spectral light in a among P spispicatespicatacata hedysarum boreale nutt var randomized complete block design boreale northern sweetsweetvetchvetch and C macu harvesting involved manually washing soil losa using growth analysis and addition seriesserles from the roots roots were cut from shoots methods root length was scanned using a codaircomair root length scanner codaircomair corp melbourne MATERIALSMAFERIALS AND METHODS australia and leaf area was measured using a licor 3100 area meter with conveyer belt LI- we used 2 indigenous species H boreale COR inc lincoln NE roots and shoots and P spispicatespicatacata as desired species C macumaculosemaculosalosa were then dried to a constant weight at 60cgoc a non indigenous perennial invasive weed for 48 h and weighed root length root weight currently dominates about 242.42 4 million ha of leaf area shoot weight and total weight of iso- rangeland and open forest throughout western lated individuals were incorporated into linear north america sheley et al 1998 habitat regression equations as dependent variables types dominated by P spicatespicata are commonly with time as the independent variable invaded by C macumaculosamaculoselosa H boreale a deep differences of growth rates between 2 taprooted legume found throughout the species were calculated using the variance northwest hitchcock et al 1977 occurs on ratio determined by the extra sums of squares sagebrush slopes and plains and in open grass- procedure for comparing slopes ratkowsky lands isley 1955 we chose H boreale because 1983 milliken and milliken mackinnon 1998 its taproot appears to allow this species to the equation used was occupy a more similar belowbelowgroundground niche to that of C macumaculosamaculoselosa than to a fibrous rooted F f RSS rssdfcrssjdf dfsjrsssdqdfrssdf grass therefore H boreale should compete more directly with C macumaculosamaculoselosa than with P riscrsscrssg and dfdfpdap are residual sums of squares and spicatespicata degrees of freedom respectively for regres- seeds of C macumaculosamaculoselosa were collected from sions run using pooled data of 2 species RSSSrssg gallatin county montana in august 1995 P and df are residual sums of squares and spicatespicata goldar variety and H boreale seeds degrees of freedom respectively for regres- were purchased from granite seed co lehi sions run using the separate data for the 2 utah in september 1995 species A variance ratio larger than the critical 0.05005 df dfdfdenominator value individual growth of fa 005 dfatopdfnumeratoramornumeratoratop denominator rejects the null hypothesisator that the slopes are isolated plants the same isolated individuals of each were species competition experiments gown in polyvinylpoly vmylvinylamyl chloride PVC pots to compare intrinsic growth rates of the species Monomonoculturescultures and mixtures of P spicataspicatespicata H pots were 15 cm in diameter and 100 cm deep boreale and C macumaculosamaculoselosa were arranged to and were filled with a pasteurizerpasteurized sifted 10 provide 2 addition series experiments with mm sievesleve soil mixture of 23 farland silt loam total stand densities ranging from 25 to 1650 fine slitssilty mixed typic argiboroll and 13 sand plants m 2 spitters 1983 radosevich 1987 soil was saturated with water and allowed to the P spicatahspicataH borealeboteborealeCC macumaculosemaculosalosa seeding 199911999 PLANT competition AND NICHE partitioning 177

densities in experiment I1 were factoriallyfaefactonally of intrainterspecificintra interspecific competition between the arranged combinations of 0 100 400 and 800 subscripted species negative BWB bvby and seeds m 2 seed production of C macumaculosamaculoselosa on bkK are the efficiencies of resource utilization infested rangeland in montana jacobs and by the populations and were estimated to be 1 sheley 1998 washington and idaho shirman firbank and watkinson 1985 the model 1981 falls within this range recommended uses the data to estimate CWV dwiD Ccyyy dy seed rate for P spicatespicata ranges from 277 to 484 ckcy and dy and provides a 95 confidence 2 seeds m holzworth and lacey 1991 and for interval around its mean if covcwvcyyy dakdwkdyyj cawcvwcyyy H boreale is 125 seeds m 2 granite seed co dkdy cy and dy 1 intraspecific competi- 1996 in experiment 2 seeding densities of H tion had a greater effect on plant weight if boreale and C macumaculosamaculoselosa were doubled 0 200 carvcvrv Dawkdwk CGVW ivkdakdvk ckwcaw andd div 1 inter- 800 1600 seeds Mrrr22 because of low establish- specific competition had a greater effect on ment of these species in experiment 1 seed- plant weight ing densities of FP spicatespicata weiewelewere the same as partitioning of resources between P spicatespicata experiment 1 as were potting conditions and H boreale P spicatespicata and C macumaculosamaculoselosa pots soil establishment conditions and and H boreale and C macumaculosamaculoselosa was calculated growth chamber conditions for the competi- using the competition coefficients from non- tion experiments were the same as described linear regressions in the equation iccicxcx1 CC for the growth of isolated individuals with the spitters 1983 connolly 1986 joliffe 1988 exception that seeds were broadcast on the deviations from unity indicate increased soil surface and manually arranged until a uniuni- resource partitioning niche separation form distribution was achieved there was no hand thinning of established plants RESULTS AND discussion ninety days after emergence plants in each in growth plants pot were clipped at the soil surface separated individual of isolated by species dried to constant weight at 60cgoc growth rates measured from root and shoot for 48 h and weighed shoot weights for the 3 weight root length and leaf area were greatelgreater species were used to determine competitive for P spicatespicata than C macumaculosamaculoselosa and H boreale interactions using nonlinear procedures proc tables 1 2 root growth measured by weight nimniin SAS institute 1991 regression equa- was greater foiroirolfor P spicatespicata 69569569.569 5 mg d 1 com- tions were of the following forms watkinsonwatkmson pared to H boreale 21212.12 1 mg d 1 and C macu 1984 losa 64646.46 4 mg d 1 based on shoot weight FP spicatespicata growth rate was 92192.192 1 mg d 1 compared P spicatespicata w11W to 161.6iglg1 6 mg d 1 for H boreale and 555.55 5 mg d 1 for wmjlwmwl AN CN dnkdenkdknk bw C macumaculosamaculoselosa root length and leaf area growth rate was 8 for P H H boreale Wwy times greater spicatespicata than boreale and C macumaculosemaculosalosa H boreale AN bv root walwml CN dnkdenkdknk length and leaf area growth was greater than C macumaculosamaculoselosa wkW that of C macumaculosamaculoselosa there was no difference wmkllwmijl akikaknk CNcknwconw DNdkvnv bk between H boreale and C macumaculosamaculoselosa in root weight however C macumaculosamaculoselosa shoot weight wyWWI Wwy and wkW are the mean shoot weights increased faster than that of H boreale per plant ofot P spispicatespicatacata H boreale and C mac P spicatespicata rate of soil depth penetration ulosaalosau10sa respectively wmamywmy wmvemywmy and amkwmkwm are 10 1 cm d 1 was greater than that of C macu the mean weights per plant grown in isolation losa 71717.17 1 cm d 1 but not that of H boreale and were derived for each species from the 80808.08 0 cm d 1 tables 1 2 rates of soil depth individual growth of isolated plants away Aay penetration were not different between C and akA represent the area required by a plant macumaculosamaculoselosa and H boreale to achieve wm and in this experiment they soil moisture is depleted first from the indicate area of the pot nwny N and nkN are upper soil horizon in semiarid ecosystems the densities of P spispicatespicatacata H boreale and C daubenmire 1970 reichenbergerReichenberger and pyke macumaculosamaculoselosa respectively cy awkdwkD C Ddakdvk 1990 found that survival of P spicatespicata seed- c1cagly and divD are the per plant equivalents of lings increased with greater depth of root w v and k and can be interpreted as the ratio competition with big sagebrush rapid growth 178 GREAT BASIN naturalist volume 59

FABLITABLE 1 crowthgrowth rates peipelper day harvest date is the independent variable for root length leaf area root weight shoot weight and soil penetration dependent variables of isolated individual pseudoroegnenapseudoroegneria spicatespicata PSSP centaurea inaculosafnaculosa CEMA and hedysarum boreale HEBO determineddetel mined from linear regression equations root leaf root shoot soil length area weight weight penetration species dmdmmmmdd 1 r2ra cm2 r2ra mg d 1 r2ra lugingmg d 1 r2ra cm d 1 r2ra PSSP 564 053 136 062 695 058 921 058 101 090 lieboHEBOIIEBO 73 048 41 019 21 072 16 074 80 076 CEMA 69 078 10 042 64 065 55 014 71 042

1 TABUTAISLI 2 variancevanance latiosratios using the extra sums of squares procedure comparing slopes growth rate d of the regres- 1 1 sionssions between 2 species in table critical f054243F 05 42 43 16565 for i ejectingrejecting the null hypothesis that the slopes are the same species root leaf root shoot soil compared length aleaarea weight weight penetration psspcemalPSSP cemalcema1 192 88 248 300 555 PSSPIIEBOPSSPHEBO 164 89 91 161 103 CEMAHEBOCEMAMEBO 176 32 12 70 053

spiesspicssp icslesiesersLKS nainesn niitulit s listed intabletintablein I1 ililcbililc 1

TABU I1 3 competition coefficients upper and lower confidence intervals P 005oos0 05 and coefficient of determination values rar2 fornhornfrombomm curvilinear legiessionregression equations using species harvest densities to predietpredictpi edletedict shoot weight of pseudo roegneriatoegroegnenaroegnerlanerianena spicatahpiuitaspicate PSSP hedysarum boiealeboreateboreave HEBO and centaurea macumaculosemaculosalosa CEMA lower upper dependent competition competition confidence confidence species species coefficient interval interval r2ra PSSP lieboIIEBOHEBO cvC 50 15 86 052 CEMADCEMAgema D 39 04 74 HEBO PSSPCPSSP cawcvw 71 42 99 039 CEMA daldvlD 30 06 53 CEMA PSSPCPSSP cawckw 60 66 84 033 HEBO dij 29 10 49

rate is believed to give a plant an advantage spicataspicate under field conditions may be due in over its neighbors because it is able to pre- part to its longer duration of growth jacobs empt their resource use harper 1980 under and sheley 1998 rather than its intrinsic the conditions of our study and when emer- growth rate gence and duration of growth are equal times competition series the rapid growth rate and soil depth penetration of P spicatafpicataspicate give it an advantage over C intraspecific interference was more impor- macumaculosamaculoselosa and H boreale of capturing both tant than interspecific interference for the above and belowbelowgroundground resources root length prediction of shoot weight for the 3 species and leaf area results suggest that H boreale table 3 the influence of P spicataspicate density may have a slight advantage over C macumaculosamaculoselosa on its own shoot weight was 5 times greater the rapid relative growth rate of B fectotecto than H boreale density and 4 times greater rum has been used to explain its dominance than C macumaculosamaculoselosa density table 3 fig 1 on P spicatespicata range harris 1967 sheley and however the confidence interval for C macu laisonlarson 1994 found the growth of C solstiholsti losa density predicting P spicataspicate weight in- tialisbialis another member of the knapweed group cludes 1 suggesting that C macumaculosemaculosalosa density of plants was 7 times greater than that of B is as important in predicting P spicataspicate weight tectoriumtectorumtectorum the success of C macumaculosemaculosalosa over P as P spicatespicata density these results suggest that 199919991 PLANT competition AND NICHE partitioning 179

tV 13 C

1- 0 F

1uau sp

2 fig 1 response surfaces using P spicatespicataspicata H boreale and C macumaculosemaculosalosa density plants rn to predict P spicataspicate shoot weight mg plant 1 with the equation wwW 0541 00182n 39nk39n 5nj 1 the light shaded surface has an H boreale density Nny ofofooto0 the dark shaded surface Nny 500 plants m 2 and the unshaded surface N 1500 plants M 2 including H boreale with P spicataspicate in a plant niche partitioning between these forbs and community will have less impact on P spicatespicata the grass was evident from competition coeffi- than on C macumaculosamaculoselosa cient ratios the coefficient ratio of C macu competition coefficients using plant den- losahlosaiosabosah H boreale was 1031.03 close to unity show- sity to predict H boreale shoot weight show ing little or no niche partitioning ratios test- H boreale density was 7 times more important ing P spicatespicata with C macumaculosamaculoselosa or H boreale than P spicatespicata density and 3 times more were 1421.42 or 1531.53 respectively indicating important than C macumaculosemaculosalosa table 3 fig 2 greater niche partitioning between grass and the confidence interval for the coefficient forbs than between the forbs predicting the importance of C macumaculosemaculosalosa den- competition in semiarid plant communities sity on H boreale shoot weight includes 1 is an important determinant of community suggesting that C macumaculosemaculosalosa is as important as structure fowler 1986 our results support H boreale density for predicting H boreale the theory that combining desirable plant weight we conclude that H boreale competes species with contrasting above and below more directly with C macumaculosamaculoselosa than with P ground growth forms will maximize niche spicatespicataspicata occupation and may increase resource pre- regressions predicting C macumaculosamaculoselosa shoot emption pyke and archer 1991 sheley et al weight showed C macumaculosamaculoselosa density 6 times 1996 tilman 1996 growth characteristics of more important than P spicatespicata density and 3 H boreale were more similar to those of C times more important than H boreale density macumaculosamaculoselosa than P spicatespicataspicata and H boreale com- table 3 fig 3 only the confidence interval peted more directly with C macumaculosamaculoselosa than for the coefficient predicting H boreale density with P spicataspicatespicata we expect H boreale to coexist effect on C macumaculosemaculosalosa shoot weight included 1 with P spicataspicate because our results suggest that suggesting that H boreale density may have these 2 species effectively partition resources the same impact on C macumaculosamaculoselosa shoot weight and minimize competitive exclusion as C macumaculosemaculosalosa density these results show the hypothesis that greater species diversity that the 2 forbs have a greater impact on each reduces community invisibilityinvasibility by increasing other s shoot weights than on P spicatespicata shoot niche occupation and facilitating more com- weight and suggest that there is some niche plete use of limiting resources is being pro- overlap between the 2 species posed as a sustainable weed management 180 GREAT BASIN naturalist volume 59

110

ppp dsa000esaooo 110

e oe ih 6 yo

fig 2 response surfaces using P spicatespicataspicata H boreale and C macumaculosemaculosalosa density plants m 2 to predict H boreale shoot weight lugingjugmg plant 1 with the equation W 0051 00182n TIN 3noano3nan 1 the light shaded surface has a C macumaculosemaculosainaculosalosaiosa density nkN of 0 the dark shaded surface nkN 500 plants m 2 and the unshaded surface nkN 1500 plants in 2

0 00100 01

00oo 10ae10.10 100 lo10

dararrerr110wer11o nd

fig 3 response surfaces using P spicataspicatespicata H boreale and C macumaculosemaculosalosa density plants m 2 to predict C macumaculosamaculoselosa shoot weight mg plant 1 with the equation wiwk oii011 00182ni00182nk00182 NiNk 39nw39n 5n15nyany 1 the light shaded surface has an H boreale density Nny of 0 the dark shaded surface Nny 500 plants rn 2 and the unshaded surface Nny 1500 plants rn 2 strategy on rangeland mcnaughton 1993 niche occupation and may be more effective in robinson et al 1995 sheley and larson 1995 minimizing invasion of taprooted weeds than tilman 1997 our study supports this hypoth- the grasses alone together they maximize esis and suggests that maintaining taprooted resource capture and preempt resource use by forbs along with caespitose grasses increases weeds sheley et al 1996 1999 PLANT competition AND NICHE partitioning 181

acknowledgments MACK RNR N 1981 invasion of bromus tectoriumtectorumtectorum L into western noinorthth america an ecological chionchroniclechronehron leleicleleie agro ecosystems 71457 145 165 material is based upon work supported this MADDOXMADUOX DMD M AND A MAYFIELD 1985 yellow starthistlestaistalstar thistle by the cooperative state research educa- infestations aiealeare on the inciincreaseease california agricul- tion and extension service USU S department ture 391039 10 12 of agriculture under agreement no 95 mcnaughton SJ 1993 blobiodiveiBio diversitysity and function of 37315240937315 2409 NRI competitive grants program grazingglazing ecosystems pages 361 383 in EDE D schuleschulze and HAH A mooney editors Biodiversity and ecosys- USDAUS DA it was published with approval of the tem function springer verlag berlin directors montana agricultural experiment PYKEpykedaandsDA AND S ARCHER 1991 plant plant interactionsiteimtei actions station as journal no J 5174 affecting plant establishment and persistence on re vegetated langelandrangeland journal of range management 4455044 550 557 CITED literature radosevich SRS R 1987 methods to study intelinteractionsactions among crops and weeds weed technology ligo11901 190igo 198 CONNOLLY 1986 on difficulties with replacement J reichenbergerrhlchenbcrgerREIC HENBERGER G AND DAD A PYKEPYKL 1990 impact of seriesserles methodology in mixture experiments journal early lootroot competition on fitness components of fourhourbourmoul of ecology 23 125 137 applied 23125 semiarid species oecologia 8515685 156 166 R 1970 steppe vegetation of washington daubenmire ROBINSON GR JF QUINNQUTNN AND ML stanlonSTANIONSTANTON 1995 washington agricultural experiment station techni- invasibilty of experimental habitat islands in a cali- cal bulletin 62 131 appp fornia winter annual grassland ecology 7678676 786 794 FIRBANK AND WATKINSON the analysis LGL G ani ARA R 1985 on SAS INSTITUTE INC 1991 SASSTAT user s guide release of plants of competition within two species mixtures 6036 03 edition SAS institute inc carygary NC 1028 appp journal of appliedofapplied ecology 2250322 503 517 SHELEY RLR L AND LLL L LARSON 1994 observation com- FOWLER of in plant com- N 1986 the role competition in paiapalaparaparativetive life history of cheatcheatgiasscheatgrassgrass and yellow staistalstar munimunitiesties in andaridarld and semiarid regions annual review thistle journal of range management 4745047 450 456 of ecologicalofecological systems 178917 89 110 1995 intelinterferenceference between cheatcheatgiasscheatgrassgrass and yel- GRANIFEGRANITE SEED granite seed UT 72 granire co 1996 lehi appp low starthistlestaistalstar thistle at 3 soil depths journal of range HARPER JLJ L 1980 population biology ofplantsof plants academic management 4839248 392 397 press london 892 appp 1996 emergence date effects on resource parti- HARRIS some GAG A 1967 competitive relationships be- tioning between diffuse knapweed centaureacentatireacentanrea dif agropyron and bromus tectorumtec tween spicaspicatumtum tectoriumtorum fusa seedlings journal of range management 49 ecological monographsM 378937 89 111 241 244 HITCHCOCK C L A CRONQUIST M OWNBEY AND J W CL JW SHELEY R L J S JACOBS AND MLM L carpinelli 1998 pacific RL JS THOMPSON 1977 vascularVascu latlai plants of the distribution biology and management ofot diffuse press northwest part 3 university of washington centaurea dlfdifdibfusagusa and spotted knapweed centau- seattle 614 appp rea macumaculosemaculosalosa weed technology 1235312 353 362 HOLZWORTH AND LACEY 1991 selection L J species cri- SHIRMAN R 1981 seed production and spring seedling teria for seeding dryland pastures in montana mon- establishment of diffuse and spotted knapweed tana state university extension bulletin ebig 12 appp journal of range management 344534 45 47 ISLEY D 1955 of the north central the leguminosae SNEDECOR GW AND WG COCHRAN 1980 statistical united states II11 hedysareae iowa state college methods iowa state university press ames 507 journal of science 303330 33 118 PP S AND R SHELEY 1998 observation life his- JACOBS JSJ RLL SPITTERSRS C J 1983 an alternative approach to the analy- range manage- smittespitte CJ tory of spotted knapweed journal of of mixed experiments 1 estimation of ggs sis cropping ment 5166551 665 673 competition effects netherlandsnethel lands journal of agricul- 1988 effects of JOLIFFE PA evaluating the competitive tural selencescienceS 313 1 1 11 on plant performance journal of interference theo- THOMPSON MJ 1996 winter foragingtoiloifol aging lesresponseresponse of elk to retical biology 130447130 447 459 spotted knapweed removal northwest science 70 LACEY JRJ R AND BEB E OLSON 1991 environmental and 10 19 pages 5 16 economic impacts of noxious weeds in TILMAN D 1996 resources competition and the dynam- LLEF james JOJ 0 evans MHM H ralphs and RDR D child acs of pages 51 75 michael press ics plant communities in editors noxious range weeds westviewWestview boul- crawleygrawleyCi awley editor plant ecology blackwell scientific der CO boston LACEY JRJ R CBC B MARLOW AND JRJ R LANE 1989 influence 1997 community mvasibihtyinvisibilityinvasibility recruitment limita- of spotted knapweed centaurea macumaculosemaculosalosa on suisulsur tion and grassland bioblodiversity ecology 788178 81 92 face yield technology runoff and sediment weed WATKINSON ARA R 1984 yield density relationships the 36273 627 630 influence of resource availability on growth and self S AL 1997 spurge biology ecol- lajeunesse ET leafy thinning in populations of vulpicvulpia fasciculatefasciculatafasciculata annals ogy and management montana state university of botany 5346953 469 482 extension bulletin eb134 25 appp LARSON SIIELEYSHELEY M MCINNIS 1994 LLL L RLR L AND MLL mclnnis received I1 december 1997 vegetation management and weed invasion in sus- accepted 8 may 1998 taining rangeland ecosystems USDASAREUSDA SARE symposium lagrande OR gleatgreat basin naturalist 592 199901999 appp 182187182 187

A NEW SPECIES OF PLECIA FROM THE GREEN RIVER FORMATION AND NEW combinations OF FOSSIL bibionidae DIPTERA

scott J Fitzfitzgeraldlfitzgeraldsfitzgeiald1geraldl

ABSTRACABSTRACT 1 pihota akerionanaakerionana n sp is described from the green river formation colorado and diagnosed with P minuminutolaminutulatula rice P myersimyersii peterson and P rhodopterina cockerell phecie interintermediaintermedialmedia scudder the genotype of myce tophaetustophaetnstophaetus and pienaplena creedencreedensissis james aiealeare transferred to the genus benPenpenthetnapenthetriathetna and hesperinushespennusHespHespe rinusfinusennus immutabilisiinmutcibilis melander is transferredti anshel i ed to aleciaplecia

Kkeikeyy words diptera bibionidae fossil aleciaplecia new species green river formation

ovelover 20 fossil species min 5 genera of bibio- genus aleciaplecia wiedemann nidae have been described from the shales of florissant colorado oligocene while only aleciaplecia wiedemann 1828 72 type species hirtha 4 species all belonging to the genus aleciaplecia fulvifulvicolliscollis fabricius 1805 by subsequent desig- have been described from the green rivelriver nation of bianchardblanchardblanehard 1840 576 formation coloichloi adowyomingutahcoloradowyomingutah eocene although there are few bibionid species known aleciaplecia akerionanaakerionana fitzgerald from the green river formation they are new species among the moiemolemore abundant diptera as was figs 1 3 found in a survey of several sites in the parachute creek member of the green river for- HOLOTYPE Y USA COLORADO rio mation codington 1993 in which bibionidae blanco county 4 mi W rio blanco bob ham- represented 22 of diptera specimens one mon USNM 498201 green 1 gleencreen river site site flF l wyoming is partic- ETYMOLOGY the specific epithet is de- ularly rich in bibionid specimens with 80 96 rived from the greek akeriosakerros lifeless and nan of all insects representing aleciaplecia pealeipealed scud- dwarf der scudder 1890 grande 1984 discussion aleciaplecia akerionanaakerionana was col- this study describes an additional species parachute green lected from the upper creek mem- ofofpleciaaleciaplecia from the river formation colo- ber of the green river formation lake uinta and of rado reassesses generic assignments locality U 2 of grande 1984 and is esti- several other fossil bibionids described from mated to be 45 47 million yr old dayvault et colorado al 1995 morphology follows mcalpine 1981 mea- DIAGNOSIS aleciaplecia akerionanaakerionana can be dis- surementssurements were made with an ocular microm- tinguished from most other eter wings were illustrated with the aid of a north american fossil aleciaplecia by its minute size and is most sim- camera lucida the following individuals made green materials available for study philip perkins ilar in size to P rhodopterina cockerell and michael kelley museum of comparative river formation colorado eocene P myersimyersii zoology harvard university MCZC virginia peterson canadian amber cretaceous and P scott entomology and peter robinson pale- minuminutulaminutolatula rice british columbia eocene ontology university of colorado boulder table I1 provides wing measurements for com- UCMC conrad labandeira and mark flo- parison of the 4 smallest based on wing length rence national museum of natural history species of nearctic aleciaplecia aleciaplecia akerionanaakerionana is smithsonianSmithsoman institution USNM specimens most similar to P myersimyersii but can be distin- have been deposited at these institutions guished by the longer and relatively narrower

I1 lputnilntdepartnient oiof entomologylntornology coloradoColo radoiddo stitesittestate university foitroitroltfort collins CO 80523 present address department of Entornoentomologylogy oregon state university cmvallisCmoi villisvallis OR 97331290797331 2907

182 1999 NEW SPECIES OF PLECIA 183

TABLE 1 comparison of wing measurements of the 4 smallest nearctic fossil aleciaplecia species P minuminutulaminutolatula rice P rhodopterina cockerell P akerionanaakerionana n sp and P myersimyersii peterson using methods of rice 1959 and melander 1949 measurements of P minuminutulaminutolatula and P myersimyersii are taken from rice 1959 and peterson 1975 respectively wing measurements in mm minuminutulaininutulaminutolatulatuia rhodopterina akerionanaakerionana myersimyersii

rice 1959195911 WL 535.3 500soo5.00 3473.47 2702.70 WW 22 216 1321.32 1201.20 RL 31 3123.12 2452.45 1921.92 abcaxc 063 0620.62 0360.36 0350.35 1134 101.0iolo 084 0380.38 0350.35 r5ra 18 1641.64 099 0500.50oso SC 0380.38 034 019 0200.20 RF 141.4 1541.54 1401.40 1071.07 rax 09 086 057 0470.47 MC 0350.35 0200.20 0150.15 0070.07 melander 19491949b ABA B 0860.86 0570.57 0470.47 ACA C 192 1321.32 110iioilo1.10 AEA E 154 1401.40 lot1071.07 ALA L 3203.20 2392.39 1871.87 BCBGB CG 0380.38 olt0170.17 022 BJB J 022 0150.15 007 CDC D 0400.40 0340.34 0200.20 CFC F 1361.36 1201.20 0900.90ogo CPC P 0220.22 0230.23 0170.17 DFD F 1041.04 0820.82 065 DHD H 1 0340.34 0190.19 0200900.20ogo EDE D 0840.84 0380.38 030 EFE F 164 0990.99ogg 0820.82 MJM J 044 067 050 MKM K 076 0880.88 075 N 0 is 070 057 0520.52 PRP R 0180.18 0230.23 0200.20 akeykey towingto wing measurements of rice 1959 WL wing length WW wing width at point of rm crossveincrossvein rice does not indicate at what point this ineasiiierneasujv ment is taken RL length ofiadiilof radial sectersector abcaxc distance between rm crossveinciosiivemcros svein and costa r4r314ra length ofr230 RZ flomfrom folkforkmoik to tip re length ofrirofairof 11415 tiornfbiorn forkfolkmoik to tip SC height of submarginal cell at tip ofofrcofrr23 RF distance between origin of radial seetotsector and its fork rax distance between origin ofofiaofiadialradialdial sector and rm croscrossveinciossvemsvein MC height of cell sc malmarginalmaiginal cell at point ofrmofomof rm crossveincrossvein bforbaorforfoi key to wing measurementsmeasuimeaski ementscements see melanderMelan derdei 1949 01or petetsonpetersonPeteisonetson 1975 measurementMeasiiiliilil enienttenient not available

wing table 1 younger age and ratio of the GENERAL description compression fos- rm crossveincrossvein to the pedicel of mg 115ils11.5 in sil on pentagonal piece of light gray shale P akeriakerionanaonana and 13 in P myersimyersii however approximately 5 cm x 5 cm and 202.02 0 555.55 5 mm due to the difference in age locality and type thick dorsoventrally preserved with wings of preservation P akerionanaakerionana more likely and legs outstretched fig 1 reverse not would be confused with P rhodopterina present which can be distinguished by the longer rel- FEMALEFLMALE head dark brown left antenna atively wider wing table 1 fig 4 and r23kg3kgb with at least basal segments difficult to distin- straight fig 4 rather than evenly curved guish 9 round compact segments apical fla figs 2 3 gellomeregellomere small nipple like anteromedial the 4 aleciaplecia species known from the green region of head with 2 faint segmented light river formation are based on wing length brown structures likely the labial palpi tho- significantly larger and thus easily distin- rax and scutellum light brown legs apex of guished from P akerionanaakerionana P dejecta scudder right mid femur tibia and 5 tarsomerestarsomeres apex wing 757.5ts mm long X 303.0 mm wide P pealeipealed of right fore femur tibia and 4 tarsomerestarsomeres loo10010.0 mm x 353.5 mm P rhodopterina 500soo5005.00 mm apex of left mid femur and tibia and apex of x 2162.16 mm and P woodruffwoodtufftrufft cockerell 70707.0 left fore femur and tibia brown with dense mm X 2752.75 mm short dalidaikdalkdark brown appressed hair apices of 184 GREAT BASIN naturalist volume 59

awe isknse F alt

figsFigsdigs 1 2 pleciaplecidplecin akerionanaakerionana photograph of holotype 1 habitus 2 wing 199919991 NEW SPECIES OF PLECIA 185 fore tibiae more robust than that of mid tibiae right fore and mid tibiae with short slender anteroanteroapicalapical spur right mid basitarsus 0400.400 40 mm long 5 times as long as wide tarsomeretarsomere 3 two 0170.170 17 mm long tarsomerestarsomeres three through five 0500.500 50 mm long right fore basitarsusbasitarsus 0540.540 54 mm long 6756.756 75 times as long as wide wings anteroanterobasalbasal portion of left wing and right wing minus most of anal lobe light brown fumose wing tip slightly pointed without anteroanteroapicalapical emargination anterior veins especially costa subcosta radius radial sector and dark blownbrown posteriorpostenor r23rg3rge r45 4 veins lighter pterostigmapteiostigma absent pedicel of m1212 15isls1.51 K times as long as rm crossveincroseroscrossvemsvein r45 262.62 6 times as long as r23rgsggs 123 relatively short and vertical evenly rounded figs 2 3 see table 1 for wing measurements abdomen dark brown posterior end tergittermitestergiteses 8 on con- cealed under rock layer posterior edge of each tergite with a thin darker brown line marking 5 the division between tergittermitestergiteses abdomen robust 3 5 of holotype 3 plecia somewhat swollen in appearance figs interprctationinterpretationinterintel pi etationelation wing of bolotype alecia akeriakerionanaakerionaiwonana 4 aleciaplecia rhodopterina 5 aleciaplecia hninutabilisiminutulnlis MALE unknown baibar 1 mmramnam aleciaplecia immutabilisimmutabihs melanderMelan deidel fig 5 not clear on the characters that distinguish these 2 genera the shape of the antenna and HespHespe immutabihsimmutabilis melanderMclanderiander 1949 20 holo hespennushesperinusennusrinus the shorter r23kg31 oblique in relation to 141514 1 5 type 112554 USNM USA COLORADO flor- indicate this species belongs to the genus aleciaplecia issant lacoe collection aleciaplecia immutabilisimmutabihs melander new combination genus penthetria deigenmeigen

discussion the family hespenmdaehesperinidae or benPenpenthetnapenthetriathetna deigenmeigen 1803 264 type species benthebentke subfamily hespenninaehesperininaeHespenninae of the bibionidae triafunebristria fufunebrisnebris deigenmeigen 1804 by subsequent mono depending upon current views of classifica- typy in deigenmeigen 1804 104 tion treated here as hesperinidaehespenmdae is differeddifferendiffeidiffee en mycetophaetus scudderScuddeidel 1892 20 type species mycetophaetus intermedius scudder 1892 by tiatedteated in part from the bibionidae by the synonym antennal flagellomeresflagello meres slender elongate and monomonotypemonotypytypy new longer than the thorax hardy 1981 antenna penthetria creedencreedensissis james rohdendorfRohdendorf 1991 this differs greatly from the antenna of typical bibionidae which have aleciaplecia creedencreedensissis james 1938 114 holotype female relatively sholtshort stout antenna with compact 4523 MCZC USA COLORADO slopes of flagellomeresflagello meres examination of the holotype of willow creek nealnear creede H immutabilisimmutabihfi is not particularly helpful as penthetria creedencreedensissis james new combination the antennae are not visible although based on wing venation raara3r23 oblique in relation to discusdiscussionS lonION based on wing venation in r4545 thene specimen clearly belongs to hesper the illustration of P creedencreedensissis provided by inus or aleciaplecia fig 4 however the photo- james 1938 115 fig 2 and examination of graph of paratype 112588 USNM of H paratype 19117 UCMQUCMC this species belongs immutabilis provided by melander 1949 to the genus penthetria penthetria can be dif- reveals that the antenna are of the bibionid ferentiated from aleciaplecia by ra3r23raa longer than and not the hesperiidhesperhespermidhesperinidhespermidinid type melander 1949 and nearly parallel to r4sr45 also treats species of aleciaplecia described by there seems to haversbeen some confusion cockerell as hesperinushespennusHespHespe rinusfinusennus and apparently was among early workers in distinguishing aleciaplecia 186 GREAT BASIN naturalist volume 59 hornfromhormhomm benPenpenthetnapenthetriathetna as cockerell 1911 discussing de castlenauCastlenau editor histonHis toiree natuinamatuiturelleelleelieeile des animaux the genus aleciaplecia from florissant Coloiacoloradodo artiartlarticulesarticulusarticul6sculeseules annelides ciustacescrustac6s arachnitesaiachnidesarachnidesarachnides myrimylimyllmyrl apodes et insects tome troisiemetroitroisi6mesiemesleme dumelldumenldum6ril parispans is still found states that the genus is in the same 672 appp eglonregioni ireferringelenefen ing to benPenpenthetriapenthetnathetna heteroptera say CARPENTERCARPLNTER FM 1992volume1992 volume 4 supelsuperclassclass hexapoda extant nearcticneal etie aleciaplecia are found only in semi- pages 279 655 in treatise on invertebrate paleontol- tropicaltiopical parts of the southeastern united states ogy paitpaltpart R arthropoda 4 volume 221 geological furthermore historic examined society of america boulder CO and university of specimens kansas lawrence as aleciaplecia UCMQUCMC identified by james het- cockerell TDA 1911 fossil insects from florissant eroptera also represent benPenrenpenthetnapenthetriathetna heteroptera colorado bulletin of the american museum of natural history 307130 71 82 benPenpenthetnapenthetriathetna interintermediaintermedialmedia scudder CODINGTONCODING TON LA 1993 climatic implications of terrestrial arthropods flomfrom the parachute creek member middle eocene green river formation gaigatgatfieldgarfieldfield county micetophaetusmycetophaetus intermedius scudder 1892 20 syn colorado unpublished mastelmaster s thesis fort hays types 3494 3463 reverse of one another state university fort hays KS 76 appp COIORADOCOLORADO MCZCMCZCUSAUSA florissant DAYVAULT RDR D LAL A CODINGIONCODINGTON D KOHLS WD HAWESHAWCS plecia alecia interintennediaintermediaintermedialmedia scudderScuddeidel as treated by evenhuis AND PM oteOYE 1995 fossil insects and spiders fromhrombrom 1994 three locations in the green river formation of the Penpenthetnapenthetriathetna interintermediaintermedialmedia scudderScuddeidel new combination Picepiceanceance creek basin colorado pages 97 115 in the green river formation in Picepiceanceance creek and scudder 1892 erected the eastern uinta basins field trip 1995 grand junction discussion geologic grand CO genus mycetophaetus society junction monotypic mycetophilid evenhuisEVLNHUIS NLN L 1994 catalog of the fossil flies of the world to include the species intermedius although insecta diptera backhuusbaekbackhuysbacklmysBackhuys publishers leiden finding formal synonymy of the genus has FABRICIUS JCJ C 1805 systemasystems antliatorumantliatorum secundussecundumsecundsecundurnumurn been nebulous various authors have treated ordines genera species adiecta synonymissynonymsynonymsis locis mucetophaetusmycetophaetus as an extinct genus of hes observationobseivationibusobservationibusibus descriptiondescriptionibusdescnptiombusibus C reichard Brunsvigbrunsvigaeae idae Rohd 1974 perinpeiimdaeperinidae rohdendorfendorf mycetophilidae GRANDE L 1984 paleontology of the green river forma- gaigalcarpenterCai penter 1992 and a junior synonym of tion with a review of the fish fauna 2ndand edition aleciaplecia bibionidae evenhuis 1994 examina- geological survey of wyoming laramieLaiamieramie bulletin tion of the synsyntypestypes of P intermedius and the 63163 1 333 HARDYuy pages illustration provided by scudder 1892 plate IlAi DE 1981 bibionidae 217 222 in JFJX mcalpine BYB V peterson GEG E shewell HJH J teskey 11 5 clearly this s II fig places speciespecleseeles in the JRJ R vockeroth and DMD M wood manual of nearctic bibionid genus benPenpenthetnapenthetriathetna which is differenti- diptera volume 1 agriculture canada monograph ated flomfrom aleciaplecia by Rs longer than and 27 neailynearlyneally parallel to r45 and from hesperinidaehespenmdae JAMESJAMLS MT 1938 family bibionidae pages 114 116 in by the short compact antennal flagellomeresflagello meres FM carpenter TE snyder CP alexander MT james and FM hull fossil insects from the creede formationfoimationhoiholFoi mationmatlon colorado part 1 introduction neurop- acknowledgmentacknowledgments S tera isoptera and diptera psyche 4510545 losios105 119 acalmcalmcalpinepinePINL JFJY 1981 morphology and terminology I1 sincerely thank bonsboris C Kondratickondiatieffkondraticffkondratieffff col- adults chapter 2 pages 9 63 in JYJ F mcalpine BXB V peterson orado state university toitolfolfor his generous sup- Peteison GE shewell HJ teskey JR vockeroth and DMD M wood manual of nearctic diptera vol- of port this searchresearchle and a critical review of ume 1 agricultureAgnagoculture canada monograph 27 the manuscript I1 thank neal evenhuis MEIGENMLICLN JW 1803 bersuchversuchveisuch einer neuen gattungs bishop museum foifolforror trying to help determine eintheilung deidelder europaeuropaischeneuropdischenischen zweiflugligenzweiflilgligen insekansek the nomenclatural status of mycetophaetus ten mag insektenkd 2259 81 and whitney cranshawCi coloradodo state 1804 klassifikazion und Beschreibung der eulocuroeuro anshaw Coloia uni- papaischenpdischcnpaaschenischen zweiflugligenzweifliagligen insekten dipteraDip teiatela linn ver foifolfor of I1 versityveivel sitssity use photography equipment eisterelstererster band abt I1 xxvnxxvii appp 1 152 11 vi appp also thank bob hammon fruita research 153 314 reichardrelReichaid braunschweig centelcentergentel colorado foifolfor making material avail- MELANDERmeimelMLI anderANDLR ALA L 1949 A report on some miocene diptera able foifolfor study and those listed with respective fromblom florissant Coloiacoloradodo american museum novi for loan of type material tates 140714071 1 63 institutions PETERSON BXBV 1975 A new cretaceous bibionid from canadian amberambel dipteraDipteiatela bibionidae canadian literature CITED entomologist 107711107 tii711 715 roiidendorrroiidlndore BB 1974 the historicalhistonealoncal development of BIANIIAKIBLANCHARD E 1840 iiistonchistoireHis toire naturelle des insectinsectcsinsectesinsectedcses diptera translatedlianiianilanslated flomfrom the russian by JEJ E moore oithoptciesorthopt6res ncvioptucsn6vropt6res h6mipt6rcshcmipteics hymenopteieshym6nopt6res and I1 theile university of albeltaalberta presspi ess edmon- lcpidoptlics16pidopt6res ctet diptcicsdipt6res in FLEL dcde laporte cointecorntecomtc ton english translation of rohdendorfRohd endorf 1964119641964.1 1999 NEW SPECIES OF PLECIA 187

1991 order diptera two winged insects pages Coloiacoloradodo and utah bulletin of the united states 444 502 in BBB B rohdendoifrohdendorfRohdendorf editor fundamentals geological survey 93193 1 34 of paleontologpaleontologyy volume 9 arthropoda tracheata wiedemannwllwil dl MANN CRWC R W 1828 ausseieuiopaischcaussercuropdische zweiflugeligczweiflftgclige cheilcheliceiatacheliceratachelicerateChelicerata smithsonianSmith soman bianesblaneslibrariesLi and national sci- insekten als foitsetzungfortsetzung des meigenschen weikeswelkes ence foundation washington DC xxxi 894 appp erster theilthelltheli schulz hammhamin xxxii 608 appp english tiantrantranslationslation of Rohdrohdendoifrohdendorfendorf 196211962 SCUDDERSCUODCR SHS H 1890 the tertiary insects ofnorthof northnorte amer- received 3 april 1998 ica united states geological survey of the territo- accepted 22 june 1998 nesrepoitl3ries report 1311 734 1892 some insects of special interest hornfrombormbomm florissant Coloiacoloradodo and othelother points in the teittelttertiariesianeslanes of gleatgreat basin naturalist 592 019991999 appp 188 192 assortative MATING IN SOLDIER BEETLES cantharidae chauliognathus TEST OF THE MATE CHOICE hypothesis

ruth Beinstein 1 and stephen BernBernsteinsteinlsteini1

ABSIHACABSTRACT 1 soldier beetles of 2 species chauliognathus basahsbasalisbabalis and C deceptus weiewelewere examined to test the crespi hypothesis that positive assortative matinginciting by size is caused by mate choice specifically we tested the piepredictiondiction that if mate choice involves choosing the largest mate available then mating individuals will be larger than nonmatingnonmating individuals foulfour samples weiewere taken at different times during the mating season fromflom each of 2 sites each sample consisted ofmatingof mating pairspans nonmatingnonmating males and nomnatingnonmatingnominatingnonmating females some of the samples contained beetles of both species others contained beetles of a single species foifor each gender elytron lengths of mating individuals weiewelewere compared with elationelytionelytron lengths of nonnianonniatingnonmatingnonmatingting individuals we found no effect ofmatingof mating status mating vs nonmatingnonmating on elytron lengths in samples that exhibited assortative mating which occurs where 2 species coexist surprisingly we found a consistent effect ofinatingofinaof matingting status on elytron lengths in samples that did not exhibit assortative mating which occurs where only 1 species exists ouioulour results do not support the mate choice hypothesis instead mate choice and assortative mating appear to be alternativealtelaitel native mating patternspatteins in which mate choice occurs where a single species exists and assortative mating occurs where 2 species coexist

keykeikhixhi words mate choice assortativematingassortative mating soldier beetles chauliognathus deceptus chauliognathus basalisbabalis

positive assortative mating by size occurs a preference for larger males crespi 1989 when the body sizes of mating pairs aieare more when male choice occurs in soldier beetles it similar than if they mated at landomrandom this is most likely because larger females carry mating pattern has been observed in soldier more eggs ridley 1983 when female choice beetles chauliognathuschauhognathus as leportedreported by mason occurs the apparent preference may be due to 1972 mccauley and wade 1978 mclammclainmelam the superior ability of larger males in over- 1982 1984 1985 and bernstein and bern- coming the higher resistance to mating of stein 1998 the ultimate cause of positive larger females mccauley and wade 1978 assortative mating by size may be sexual selec- mccauley 1981 A prediction of the mate tion in which differencesdiffeidiffee ancesences inm reproductiverepi eductiveoductive suc- choice hypothesis is that the mean size of mat- cess caused by competition over mates are ing individuals will be larger than the mean i elatedrelated to body size anderssonAndeissonelsson 1994 alter- size of nonmatingnonmating individuals arnqvistarnquistArnqvist et al natively positive assortative mating by size 1996 most data on arthropods do in fact may be an artifact of environmental factors show this mating pattern either for females or such as tempoitempostemporalal or spatial covariancecoval lancelanee of body for both sexes crespi 1989 size among mating males and females in the study described herein we exam- many hypotheses have been offered to ex- ined the mate choice hypothesis in 2 species plain assortative mating in arthropods crespi of soldier beetles chauliognathuschautiognathus deceptus le 1989 one of these is the mate choice hypothe- conte and C basalisbasaltsbabalis fender these beetles sis based on sexual selection inm which indi- mate conspicuously occur in large populations viduals choose larger mates because they ben- and remain coupled for many hours in a pre- efit reproductively and aiealearcare differently capable liminary study with marked pairs we found ofot exercising choice darwin 1871 ridley 1983 that 68 of mated pairs remained together for male choice involves a large male mating ad- more than 5 h and 34 for more than 17 h in vantage in male male competition foiroirolfor females a previous study bernstein and bernstein 1998 combined with a preference foiforholbol larger females we found positive assortative mating by body female choice involves factorsfactois that mcleasemeleaseincrease size in some populations but not in others largelailalial ge femalelemale pairingpan mg probability combined with arnqvistarnquistArnqvist et al 1996 lists for the mate choice

1qartnntdcpiitnifiitolof viavivinvihiimionirnntinwohd population andincloiainisinilorganisinicOrgani sinic biology uiiiversityuinvisituluUinvisit ofot coloradocoloichloi ado Boulbouldeiliouldeibouldendeidel CO 80309

188 1999 MATE CHOICE IN SOLDIER BEETLES 189

hypothesis the following predictions 1 the babosaitssalissaltssaits an unexpected result since the 2 form of assortative mating will be true 2 species cannot be distinguished in the field mating males will be larger than nonmatingnonmating males were identified to species by the shape males and 3 mating females will be larger of the copulatory organ using the key pro- than nonmatingnonmating females the first prediction vided min fender 1964 females were identi- has been tested bernstein and bernstein 1998 fied by species specific correlations between and found true the correlation between body length of the posteriorpostenoienol elytron spot and length sizes of mates is true linear rather than appar- of the elytron bernstein and bernstein 1998 ent in which the variance in male size changes the right elytron of each beetle was severed with increasing female size here we test the from the body and its maximum length was remaining 2 predictions specifically that mat- measured to the nearest 00010.0010 ooi001 mm using a ing individuals are larger than nonmatingnonmating indi- binocular microscope with an eyepiece viduals in the populations that exhibit assorta- micrometer tive mating but not in the populations that do the effects of sampling time and mating not exhibit assortative mating status mating versus nonmatingnonmating on body size were analyzed by 2 factor analyses of variance METHODS sampling time represents both progression of the mating season and variations in assortative we collected mating and nonmatingnonmating beetles mating as only the first 2 samples at the from 2 sites one on the plains and the other in canyon site exhibited this mating pattern a canyon within 30 kmkin of boulder colorado bernstein and bernstein 1998 all samples the plains site is a meadow elevation 1760 in were reduced in size by random elimination near eldorado springs where beetles were to the smallest sample in order to meet the feeding and mating on sunflower helianthus recommended equal sample sizes for the 2 annausannuus L blossoms the canyon site is a factor ANOVA we began with 24 groups a roadside in south st vrain canyon elevation male group and a female group from each of 1830 in near lyons where beetles were feed- the 4 samples of C deceptus at the canyon ing and mating on blossoms of rabbitbrushrabbitbrush site from each of the 4 samples of C basahsbasalisbabalis at chrysothamnus nauseosus pall britt each the canyon site and from each of the 4 sam- sampling site encompassed an area of less ples of C basahsbasalisbabalis at the plains site five of the than 050.5os ha at each site 4 samples were taken 24 groups were eliminated from analysis 4 at lwi1 wk intervals during the approximately because they were too small the male group month long mating season all beetles in each and female group from samples 3 and 4 of C sample were collected on a single morning basahsbasalisbabalis at the canyon site and I1 because it was between 0900 h and 1030 h at this time of so much smaller than the other samples sam- day beetles are too cool for rapid locomotion ple 4 of C basalisbasahsbabalis females at the plains site and so are likely to have been coupled since at that we preferred to eliminate it rather than least the previous evening the sluggish condi- greatly reduce the other samples where sig- tion also prevents sampling bias as any beetle nificantnificant effects of sampling time were found regardless of size or mating status is easily the means of mating and of nonmatingnonmating indi- captured by sliding it from a blossom into a viduals were compared using the newman collecting vial whenever possible each sample keuls multiple comparison test zar 1996 with consisted of 40 mating pairs 40 nonmatingnonmating the level of significance set at 0050.050 05 males and 40 nonmatingnonmating females samples of in presenting a summary of our results we mating pairs are the same ones reported in an compare the pattern of assortative mating with earlier publication in which patterns of assor- the pattern of size differences between mating tative mating were described bernstein and and nonmatingnonmating individuals among the samples bernstein 1998 part of this comparison involved testing for beetles were frozen within a few hours after differences among the pearson product corre- capture and then preserved in 70 alcohol lation coefficients r our measure of assorta- plains samples consisted entirely of chauliog tive mating for these tests we followed the nathusbathus babasalisbabalissalis canyon samples consisted of 2 statistical procedures described by zar 1996 species chauliognathus deceptus and C with the level of significance set at 0050.05oos0 05 190 GREAT BASIN naturalist volume 59

TABLITABLE 1 two factor ANOVA results iolfoifor males effects of sampling time and mating status mating versus nonmatingnonmating on elytron length NS P 005oos0 05 NA not applicable species effect of effect of interaction newman keuls test site sampling time mating status effect P 0050 05

C deceptusdeceptns f321626 42704.2704 270 f1216F 216 1605016.05016 050 f3216faf3 g 49814.9814 981 mating x x2xax x3xaxy x4xax CIcanyonTyon P ooi0010 01 P 0 0005000050.0005 P 0 0025000250.0025 nonmatingNon mating xxxaxxT 5f2 jckjc3 t4tax 1 P C bawlf f136fi 3 03800 380 f136fy 1264264 f136i 36 01570 157 NA canyon NS NS NS

C hdsalvbasalisbabalis f3240F 240 02390.2390 239 f1240FI 240 3447634.47634 476 faf3f3240240 02210.221221 NA plains NS P 0 000500005 NS

IAISLITABLE 2 two lactolfactor ANOVA results foiroirolfor females effects of sampling time and mating status mating versus nonmatingnonmating on elytron length NS P 0050.050oos05 NA not applicable species effect of effect of interaction newman keuls test site sampling time mating status effect P 0050.05oos0 05

C deceptufdeceptus F 47524.7524 752 F 30.82730 827 8.4748 474 af5f 5f2 5f3 x fy13176 f171 30827 f171faf3 8474 mating xxxaxx t4ta canyon P 00050 005oos P 0 000500005 P 0 000500005 nonmatingNon mating xxxaxxT y2ya t3ta x5f4 C 124 NA bawkshawksbamks fif1363 012400.124 fj36F 3 00270270 fif13636 01080 losios108 canyon NS NS NS

C basalishasahsbabalis f2234t 234 00140.0140 014 FIf1234234 1541015.410iai5 410 f2234 05340.534534 NA plains NS P 0 000500005 NS

RESULTS there are no effects of sampling time mating status or interaction on the elytron length of results of the 2 factorfactor analyses of variance either males or females for C basalisbabalis at the aiealeare presentedpiesen ted for males in table I1 and females plains site there is a significant effect of mat- in table 2 each table shows the effect on ing status on elytron length but no effect of mean elytron length of sampling time mating sampling time nor of an interaction ie mat- status and interaction between sampling time ing and nonmatingnonmating individuals are different in and mating status results of the newman size and this difference remains the same keuls test for differences among sampling across samples times are provided for each ANOVA with a results of our studies are summarized in significant difference among samples table 3 where for each sample we give the males and females exhibit the same relative abundance of a congener correlation ANOVA pattern in C deceptus there are sig- coefficient r of elytron lengths of mating nificantnificant effects of sampling time and mating pairs size ratio of mating to nonmatingnonmating males status as well as the interaction between sam- and size ratio of mating to nonmatingnonmating females pling time and mating status on elytron lengths for both species samples 1 and 2 from the an interaction effect indicates that size differ- canyon site have higher correlation coeffi- ences between mating and nonmatingnonmatmgnonmating individ- cients higher percentages of the less abun- uals occur in some samples but not in others dant species and virtually no size differences according to the newman keuls multiple between mating and nonmatingnonmating individuals comparisons tests elytron lengths of mating for C deceptus in samples 3 and 4 from the individuals remain the same across samples canyon site and for all C basalisbabalis samples from whereas in nonmatingnonmating individuals there is a the plains site correlation coefficients are low significant difference between samples 2 and percentages of the less abundant species are 3 iei e nonmatingnonmating individuals are smaller in low and mating and nonnonmatingmating individuals samples 3 and 4 than in samples 1 and 2 for are more different in size than in samples 1 C basahsbasalisbabalis at the canyon site samples I1 and 2 and 2 from the canyon there are significant 1999 MATE CHOICE IN SOLDIER BEETLES 191

TABLE 3 summary of sample values percent of the less abundant species n total number of individuals in the sample strength of assortative mating coefficient r of the correlation between elytron lengths of mates n number of pairs and size ratios of mated and unmated individuals n number of ratios n of groups in the ANOVA

sample 1 sample 2 sample 3 sample 4 C deceptus canyon site less abundant 294 325 106 131 species in sample n 160 n 160 n 160 n 160 r values 0421 0651 0157 0019 differences among n 30 n 28 n 38 n 36 the values P 0050 05 male size ratio 1000 1002 1057 1064 mated unmated n 28 n 28 n 28 n 28 female size ratio 1006 1001 1087 1067 mated unmated n 23 n 23 n 23 n 23

C basalisbasahsbabalis canyon site less abundant 294 325 106 131 species in sample n 160 n 160 n 160 n 160 r values 0533 0529 too few too few differences among n 10 n 12 the values NS

male size ratio 1022 1013 too few too few mated unmated n 10 n 10 female size ratio 1003 1015 too few too few mated unmated n 10 n 10

C basalisbasahsbabalis plains site less abundant 0 0 0 0 species in sample n 160 n 160 n 160 n 124 r values 0042 0095 0168 0088 differences among n 40 n 40 n 40 n 40 the values NS male size ratio 1048 1042 1047 1033 mated unmated n 31 n 31 n 31 n 31

female sizesiwe ratio 1039 1037 1020 too few mated unmated n 40 n 40 n 40

differences xax22 95719.571 among correlation which individuals prefer larger mates A pre- coefficients r of the 4 samples of C deceptus diction of this hypothesis in addition to assor- at the canyon site there is no significant dif- tative mating is that mating individuals are ference z 00110.011 between correlation coeffi- larger than nonnonmatingmating individuals because cients of the 2 samples of C basalisbabalis in the larger individuals mate more often andor canyon nor are there significant differences remain coupled for longer periods of time xax2 03090 309 among the 4 samples of C basalisbabalis than the less desirable smaller individuals in the plains site we tested the specific prediction that mating individuals are larger than nonmatingnonmating individ- discussion uals in populations that exhibit positive assortative mating and found the prediction to be one hypothesis to explain positive assorta- false for 2 species of soldier beetles C basalisbabalis tive mating is the mate choice hypothesis in and C deceptus in populations of these species 192 GREAT BASIN naturalist volume 59 that exhibit assortative mating as measured by advantage of being paired regardless of mate corrcorrrelationsrelations of the elytron lengths of mates size under these conditions in our study the theretheithel e are no differences between elytron lengths advantage of having a large mate may be of mating and nonnonmatingnonmatmgmating individuals in popu- countered by the disadvantage of mating with lations that do not exhibit assortative mating an individual of the wrong species the rela- howeverhowe veivel mating males and females are larger tiontionshipship between inhibition of mate choice and than nonnonmatingnonmatmgmating males and females stimulation of assortative mating however Is mate choice the usual mating pattern in remains unclear soldier beetles where mate choice as defined by larger mating than nonmatingnonnonmatmgmating individuals literature CITED was examined in previous studies all involving C pennsylvanicuspennsylvamcuspennsylva nicus the results were mixed ANDERSSON M 1994 sexual selection princeton univer- press princeton 599 mason 1980 found evidence of female choice sity NJ appp ARNQVIST G L ROWE JJ KRUPA AND A SIH 1996 larger mating than nonmatingnonnonmatmgmating males in 2 of 3 assortative mating by size a meta analysis of mating populations sampled in northern new york patterns in water striders evolutionary ecology 10 but no evidence of male choice mclain 1982 265 284 found both male and female choice in all 6 BERNSTEIN R AND S BERNSTEIN 1998 assortative mating by size in two species of chauliognathuschauhognathus henz populations sampled in northern georgia A coleoptera cantharidae southwestern naturalist later study mclain 1985 found no evidence 436243 62 69 of female choice only males were measured CRESPI BJB J 1989 causes of assortative mating in arthro- in a dinehineclineeline of 15 populations in northern georgia pods animal behavior 3898038 980 1000 DARWIN C 1871 descent of and selection in A of our results the man in tentative interpretation is relation to sex john murray london that mate choice is the normal mating pattern FENDER KMK M 1964 the chauhognathimchauliognathini ofamericaof america north in soldier beetles but that presence of a con- of mexico coleoptera cantharidae northwest sci- gener on the same host plant inhibits this nor- ence 3852 64649595 106 mal behavior and triggers assortative mating MASON LGL C 1972 natural insect populations and assortative mating american midland naturalist 8815088 150 157 is what is the evidence that assortative mating 1980 sexual selection and the evolution of pair and mate choice are alternative mating pat- bonding inm soldier beetles evolution 3417434 174 180 terns in soldier beetlesbeetles99 three previous stud- mcgaulMcCAULmcgauleyMGCAULEYmccaulbydebynebYDEeyneDE 1981 application of the kencekenee bryant ies all involving C pennsylvamcuspennsylvanicuspennsylva nicus have eval- model of mating behavior to a natural population of soldier beetles 117 uated both mating patterns min the same popu- american naturalist 117400400 402 MCCAULEY DED E AND MJM J WADE 1978 female choice lations mclain 1982 found both male and and the mating structure of a natural population of female choice but no asssortative mating in 6 the soldier beetle chauliognathus pennsylvanicuspennsylvamcuspennsylva nicus populations he sampled in georgia a result evolution 3277132 771 775 that lends support to our interpretation how- MCLAIN DKD K 1981 interspecific interference competition and mate choice in the soldier beetle challichaulichauh ever in a later study mclain 1985 found nei- ognaognathusthus pennsylvanicuspennsylvamcuspennsylvanicus behavioral ecology and ther mate choice nor assortative mating in a sociobiologySociobiology 9659 65 66 dineclineelinegine of 15 populations in georgia mccauley 1982 density dependent sexual selection and and wade 1978 found both mate choice by positive phenotypic assortative mating in natural males and by females and populations of the soldier beetle chauliognathus assortative mating pennsylvanicuspennsylvamcuspennsylvanicus evolution 36122736 1227 1235 in populations they studied in illinois thus 1984 host plant morphology speciation and the the resultslesults on C pennsylvanicuspennsylvamcuspennsylva nicus do not form a economics of mate choice in the soldier beetle consistent pattern chauliognathus pennsylvanicuspennsylvamcuspennsylvanicus evolutionary the- our results suggest that the soldier beetles ory 7637 63 67 1985 clinal in morphology and assorta- studied exhibit mate choice behavior variation in we except tive mating in the soldier beetle chauliognathus in the presence of a congener mclain 1981 pennsylvamcuspennsylvanicuspennsylva nicus coleoptera cantharidae biological in studies of C pennsylvanicuspennsylvamcuspennsylvanicus also found journal of the linnaean society 2510525 105 117 inhibition of mate choice behavior in females RIDLEY M 1983 the explanation of organic diversity the this comparative method and adaptations for mating by the presence of other species in case clarendon press oxford wasps wasps were less aggressive toward pairs ZARJHZAR JH 1996 statisticalbiostatisticalBio analysis ard3rd edition prentice ofbeetlesofbeetles than toward individuals so that paired hall inc upper saddle river NJ 662 appp females were able to feed more efficiently than impaired females thus the advantage of received 31 march 1998 accepted 29 june 1998 mating with a larger male is countered by the great basin naturalist 592 019991999 appp 193 194

BODY SIZE DYNAMICS OF COUGARS FELIS CONCOLOR IN OREGON

stephan G Kohlmannkohlmann1kohhnann11 and richard L Greenlgreenigreenl2greenlegreen122

key words felis concolor oregon body mass

we investigated body size dynamics of 1076 of the 4 yr old cougars while 54 of the harvested male and female cougars in oregon premolar pairs of older cougars were the same from 1987 through 1997 cougars were checked age or differed by I1 yr weighed and measured by oregon depart- body length BL nose to ottip of tailtalltalitaii and ment of fish and wildlife biologists within 48 body mass BM were significantly correlated h of harvest one or 2 upper prepremolarsmolars were for both sexes males BL 14567145.67145 67 removed and age was determined by cemen- 10731 073bm1073bmBM n 645 F 112371123.71123 7 P 0 0001000010.0001 tum annuli at latsonsmatsons lab milltownMilltown MT r2ra 0630.630 63 females BL 12366123.66123 661 68bm168bm precision of age determination was estimated n 431 F 5964596.4596 4 P 0 0001000010.0001 r2ra 0580.580 58 from 864 blind tests of pairs of prepremolarsmolars for mean body mass of male cougars was on aver- which the identity of the individual cougar age 508 142 gigreatereater than that of was not known to the analyzing laboratory females this difference was statistically sig- personnel ages assigned to each of the 2 pre nificantnificant for all age classes except kittens table molars agreed exactly or varied by I1 yr in 93 1 and supports results from earlier studies

TABLE 1 mean body mass kg standard error and sample size of harvestedhaihar vested male and female cougars in oregon 1987 1997 asterisks indicate significant differences in body mass between the sexes P 0050.05oos0 05 significant differences among seasonal mass within age groups are represented by different supelsuperscriptscript letters tukey test P 00500.0505 jun aug sep nov dec feb mar may total sex age xY sts n T svs n x svs n x s n x sts n male 0 boo20020goo0 444.44 4 6 17117.117 1 3873 8 7 22622.622 6 15181.5181 5 18 15015.015iso 0 1481 4 8 19719.719 7 133913391.3391 3 39 1 37337.337 3 202 0 18 43443.443 4 181818181.8181 8 18 43843 8 181 8 32 44044 0 3083.083 0 8 42242 2 iili111.11 1 76 2 38138.138 1 323.23 2 81 49049 0 134011 3 4011 50650 6 080.80 8 8311 46046.0460249124 9aaa 49149 1 070 7 140 ab 1 3 521236iib521 232.3 6a1gai 56815215681521ab56856.8 15211521alal 57257 2 101.0io1 0 71711 49649.649 6 321113 2 ii11 56156 1 080.80 8 109 4 55355 3 323 2 7 58858.858 8 13181 3 18 59659 6 101.0loio1 0 54 57957.957 9 28112 8 11 58958 9 080.80 8 9090s1 5 55055 0 3343 3 4 59459.459 4 16161.6161 6 16 60360 3 11461 1 46 56856 8 2322.33 2 59759 7 090.90og 9 68681 6 57357 3 606.0go6 0 4 60060.060goo 0 18161 8 16 62262 2 115411541.1541 1 54 53853.853 8 202 0 3 612gig61 2 090.90og 9 77r77 7 54554.554 5 010 1 61461.461 4 3063.063 0 6 65565.565 5 151 5 25 63663 6 010 1 64464 4 131 3 33r33 8 616gig61 6 1621.62lgb1 6 2 73673.673 6 595.95 9 3 63663.663 6 19161 9 16 63663 6 010 1 64864 8 171.71 7 22 9 63163.163 1 8848 8 4 65565 5 262.62 6 4 64264.264 2 3193 1 9 68068 0 262 6 3 64864 8 222.22 2 20 10 71871 8 010 1 71271 2 2032 0 3 66766.766 7 18231 8 23 61461 4 010 1 67267 2 16281.6281 6 28 female 0 14314.314 3 1941.941 9 4 15712111157 gillgili2111 18018 0 17161.7161 7 16 18718.718 7 21112 iiiili111 17217 2 10421 0 42 1 284317284317lb28428.4 31 7alaallb 336336ll21a11 21a 31931 9 11 28th28ab 27727.727 7 141 4 1311311 31331 3 070.70 7 69- 2 31531 5 343.43 433abaab 36136107261107 26a11 36936 9 06 gsa 3253252020 1211211 36036 0 050 5 106 3 31801318 0 1 38138 1 131 3 18 37237 2 090 9 41 34434 4 2422.44 3 37237 2 070.70 7 63 4 38438.438 4 222 2 10 39139 1 070.70 7 34 38938.938 9 191.91lgig 9 6 39039 0 070.70 7 50 5 2912929.11 0 1 40440 4 222 2 7 39939 9 070.70 7 32 31831.831 8 919.19gi 122 39339 3 080.80 8 42 6 4094040.99 0 1 42842 8 131 3 8 39039 0 0800.8os 8 20 35535.535 5 0 1 40040 0 070 7 30 7 35035.035 0 0 1 43543.543 5 252 5 5 41841 sioslo810101.0iolo 11 45045.045 0 0 1 42042 0 101.0io1 0 18 8 4774747.77 020 2 40940.940 9 15101.5101 5 10 48042042 0 151.51 5 12 9 48248 2 010 1 41541.541 51111 11 35935 9 010 1 41641 6 12 13 10 35935 9 737.37 3 2 43643.643 6 050.50 5 2 40940 gli911siisillilg111616 45045 0 0 2 41141 1 101.0loio1 0 22

oregon department of fish and wildlife wildlife population laboratory 7118 NE vandenbergg ave colcorvallisCoi vallis OR 97330 2correspondingoi responding authorauthol

193 194 GREAT BASIN naturalist volume 59

anderson 1983 body mass of subadult cougars dimorphic and speculate that significant weight increased rapidly until 4 yr of age after which changes inm subadultssubadults are a resuitresult of food depri- annual weight gain increments were less than vation during dispersal 1 table 1 body mass of harvested cougars varied literature CITED among seasons table 1 but significant seasonal mass changes occurred only in subadultssubadults ANDERSON AEA E 1983 A critical review of literature on puma felis concolor colorado division of wildlife young cougars experienced significant weight special report 54 losses in spring and summer of their and2nd yr HORNOCKER MGM G 1970 an analysis of mountain lion hornockerHomocker 1970 suggested that young cougars predation upon mule deer and elk in the idaho at the time of dispersal are vulnerable to star- primitive area wildlife monographs 21 and accidental mortalitiesmortali we conclude vation ties received 12 august 1998 that cougars in oregon are highly sexually accepted 15 october 1998 greatgi edteat basin naturalist 592 019991999 appp 195 197

HELMINTHS OF THE LOWLAND BURROWING TREEFROG ptyernohjlaptyernohyla FODIENS HYLIDAE FROM SOUTHERN ARIZONA

stephen R coldGoldGoldbergberglbergi1 charles R bursey2bmsey2 and guillermo Gagalindolgalindosgalmdo1lindol

key words pternohylaiteiptei nohylamohyla fodiens hylidae helininthshelminthshelminthes arizonaai izonatzona

the lowland burrowing treetreefrogfrog pternohyla rhabdiasRhabdias americamericanosamencamisamericanusamenanuscamis baker 1978 physaloptera fodientfodiens boulenger 1882 occurs from pima sp larva only and skjabinopteraskrjabinoptera sp larvae county arizona south into western mexico only number of helminthshelminthes prevalencepievalence mean in mexico it is found from sonora to michoa- intensity range mean abundance and infec- ehneancan at elevations from sea level to about 1490 tion site are given in table 1 selected hel m stebbins 1985 there are no published mintasminths were placed in vials of ethanol and accounts of helminthshelminthes from this species the deposited in the USU S national parasite collec- purpose of this note is to report the helminthshelminthes tion beltsville maryland distoichometra bufo of pternohyla fodientfodiens from southern arizona msnis 87748 aplectana wocanensisitzocanensis 87749 cos and to list the other anuran species in which mocercella haberihabera 87750 Rhrhabdiasrhabdtasabdias americanosamericanusameric anus these helminthshelminthes occur 87751 physaloptera sp 87752 skrjabinoptera forty pternohyla fodientfodiens 9 females 31 sp87753sp 87753 males mean snout vent length 551ssi55.1 mm none of the helminthshelminthes found during this 292.999gg s range 45 70 mm collected 353.5 km E study was unique to pternohylafodienspternohyla fodientfodiens dis of sells pima county arizona 3155n toichometratoichometra bufonisbufomsbunonis has been reported previ- 11155w elevation ca 2300 m during ously from 6 anuransanusans from arizona bufo cog 1958lllwllew1962 were borrowed from the herpetol- natus B micromicroscaphusscaphus B punctpunctatusatus B reti ogy collection of the university of arizona formis B woodhourwoodhoumwoodwoodhousiihouahoum and scaphiopus couchicouchncouchii tucson UAZ numbers 16141 16155 16161 goldberg and bursey 1991a 1991b goldberg 16164 16166 16200 16204 16206 16208 et al 1996b 1996c it also has been reported 16212 16240 16258 16261 16908 16913 from bufo boreas from california B cognatescognatuscognatus 16915 16918 the treetreefrogsfrogs were originally from new mexico and oklahoma B terrestristerrestnstetreterrestris fixed in 10 formalin and preserved in 70 from georgia B woodhousiiwoodhousnwoodhousewoodhousn from nebraskaneblNebi aska isopropanol the body cavity was opened gas- and hyla regilla from oregon dickey 1921 trointestinaltrointestinal tract removed and the esopha- kuntz 1941 douglas 1958 koller and gaudin gus stomach small intestine and large intes- 1977 hardinhardm and janovy 1988 goldberg et al tine were cut open and the contents exam- 1995 because distoichometra bufomsbufonisbunonis has a ined in addition the lungs and bladder were wide distribution in north america its occur- searched for helminthshelminthes nematodes were rence in pternohylapternohylafodiensfodientfodiens is not unexpected identified utilizing the glycerol wet mount it is a new host record procedure cestodescustodesCestodes were stained with hema aplectana wocanensisitzocanensis has been reported toxylintoxylon and mounted in canada balsam for from 7 anuransanusans from arizona bufo alalvanusvarius B identification terminology usage is in accor- cogcognatescognatusnatus B micromicroscaphusscaphus B punctatuspunctatus B dance with bush et al 1997 retiretiformisformis B woodhousiiwoodhousnwoodhousewoodhousn and CastroGastrogastrophrynecastrophrynephryne the helminth fauna of pternohyla fodientfodiens ohvaceaofivacea goldberg and bursey 1991a 1991b consisted of I1 species of cestode distoichome goldberg et al 1996b 1996c 1998 addition- tra bufonisbunonis dickey 1921 and 5 species of ally it has been reported from bufo cogcognatuscognatesnatus nematodes aplectana iftocanensisitzocanensis bravo hol- from new mexico and B marinusmartnus from costa lis 1943 cosmocercellaCosmocercella haberihabera steiner 1924 rica and mexico bravo hollis 1943 brenes

I1 department of biology whittierWhittieitier college whittierwhittlerWhit tieltiei CA 90608 depaitmentdepartment21epartment ofot biology pennsylvania state university Sheshenangoshcnangonango Caincampuspus 147 shenangoShe nango avenue shalonsharon PA 16146

195 196 GREAT BASIN naturalist volume 59

TAB I1 1 bommerom talm 1 helminthshelinintlishelminthes fornhornfrom pternohlylafodienspternohylafodiens N 40 from southeinsousouthernsounheinthein arizona evaleneepiprevalenceevalence mealmeani intensity mean abundance helminth N 6 iangelangerange s site CESTODAcl sioda distoichometradi&totchmnetra bufomsbufonisbunonis 7 8 232 3 232 3 1 5 018081018 081 small intestine nlNEIMAIODAMAIODA aplectana itzocaneiwiftocanensis 110 18 15715 7 36736 7 1 99 2752 75 15615.615 6 intestines Cosinocosinocercellaco&mocercellacercellacercelia habenhahhnhaberl 2587 55 lit1176117 6 1261 2 504 646864 68 10986109.86109 86 intestines rhabdiasRhabdias americanosamericanusamericanus 6 6 60 0150 15 0950.950 95 lung phisalopteraphysaloptera sp larvaelaivae 7 7 181.81 8 050 5 1 2 0180.180 18 055oss0 55 stomach skjabinopteraskrjabmoptera sp loiilarvalcii va 1 1 10 0030.030 03 0160.16olg0 16 stomach

and bravo hollis 1959 caballero deloya 1974 the appearance of larvae from these 2 genera baker 1985 goldberg et al 1995 the known is the result of treetreefrogfrog diet rather than a nor- distribution of aplectana iftocanensisitzocanensis suggests mal part of the parasite s life cycle to our it may be a middle american species that knowledge no case of adult physalopterans in i eachescachesreaches its northern distribution in the south- anuransanusans of north america has been reported western deserts of the united states its any insectivore could be expected to have lar- occurrence in pternohylafodienspternohyla fodientfodiens is a new host vae of these 2 genera record in summary pternohyla fodientfodiens was found cosmocercellaCosmocercella habenhaberl has been reported to harbor 1 species of cestode and 3 species of hornfromhormhomm 2 anuransanusans from arizona hyla alemareniaremarenicoloraremcolorcolor nematodes which also occur in other anuransanusans and H wnghtorumwrightorunt goldbergGoldgoldbeigoldbegbeibel g et al 1996a as of north america pternohylafodienspternohyla fodientfodiens is a new well as H aremareniarenicoloraremcolorcolor from utah H cinerea host record for each of these species exami- from north carolina and H versicolor from nation of samples of P fodientfodiens from the south- virginia and ontario canada stermerstemmersteinersterner 1924 ern part of its range in mexico will be neces- pairyparry and grundmann 1965 campbell 1968 sary before the diversity of parasitic helminthshelminthes bakelbaker and adamson 1977 parry and grund- in this cylidhylid can be known mann 1965 reported I1 of 23 rana pipienspipkens from utah to be infected by cosmocercellaCosmocercella we thank charles H lowe department of habenhaberl the only report of infection of a non ecology and evolutionary biology university cylidhylid species because of its wide distribution of arizona for permission to examine speci- as a parasite of hylidshylins its occurrence in pter mens of pternohylafodienspternohyla fodientfodiens nohylamohyla fodientfodiens is not unexpected it is a new hosthostihostrecordhostiecordecordrecord literature CITED Rhrhabdiasrhahdiasabdias americanusamericanosamericamencanusanus has been reported from 5 anuransanusans from arizona bufo alvanusaldariusvarius B andehsonrcANDERSON RC 1992 nematode parasites vertebratesofofvertebratesvelteveitebrates thentheutheir development and tiantransmissionsmission CAB inter- cognatescognatuscog B microscaphusscaphus B natus micro retiretiformisformis and national wallingford axonoxon UKU K 578 appp B woodhousiiwoodhousnwoodhousewoodhousuusn goldbergGoldgoldbeigoldbegbeibel g and bursey 1991a BAKERBAKLR MRM R 1978 morphology and taxonomy of rhabahab goldberggoldbeigoldbeg g et al 1996b 1996c it has also been dias sppapp nematoda rhabdiasidae fromhrombrom reptiles and reported from B americamericanusamericanosamencanusanus and B wood amphibians fromhiom southern ontario canadian jour- housti from eastern north america and canada nal of zoology 56212756 2127 2141 1985 redescription ofaplectana wocanensisitzocanensis and baker 1978 and B cognatescognatuscognatus from new mexico A incerta nematoda cosmocercidaecosmoceicidae from amphib- goldberg et al 1995 this is the first report ians transactions of the american microscopical of rhabdiasRhabdias americanusamericanosamericamencanusanus in an anuran other than society 104272104 272 277 a bufonid and its occurrence in pternohyla BAKERBAKLR MRM R AND MLM L ADAMSON 1977 the genus cosmo cercellace steinersternedsternerstemmed 1924 nematoda cosmocercoideacosmoceicoidea is icella fodientfodiens is a new host record canadian journal of zoology 55164455 1644 1649 species of physaloptera require insects as BRAVO HOLLIS M 1943 DOS nuevoscuevos nemanematodosncmdtodosnematodestodos parasites an intermediateintel mediate host while species of skraskr de anurosacuros del sur de puebla analesanaces del institutoinstituteinstitutor de jabinoptera require ants as an intermediate Biologia umveisidaduniversidad nacional aut6nomaautonomaAutautonomyonoma de mexico 146914 69 78 host anderson 1992 since only larvae weiewelewere pter BRENEsBRLNLS RRR R AND M BRAVO HOLLIS 1959 helmintosHelmintos de found in this study we assumed that laa republica de costa rica VIII nematoda 2 nohylamohyla fodientfodiens is an unsuitable host and that algunos nemaneraftodosnematodesnematodostudostodos de bufo mannusmarinus mannusmarinus L y 1999 NOTES 197

a1gunas alkunasalgunas consideracionescon&ideraciones sobrebobre los geneiosg6nerosgeneiosa oxyso GOLDBERG SRS R CRC R BURSEY KBK B MALMOS BKB K SULLI- mariummatiumtnatium y aplectana revistadevista de BiBioloologiagiagfa tropicalTiopical 7 VAN AND H CHEAM 1996b helminthshelminthes of the south- 35 55 western toad bufo micromicroscaphusscaphus woodhouse s toad BUSH AOA 0 KDK D LAFFERTY JMJ M LOTZ AND AWA W SHOSTAK bufo woodhousiiwoodhousnwoodhousewoodhousuusn bufonidae and thentheir hybrids from 1997 parasitology meets ecology on its own terms central arizonaanzona great basin naturalist 5636956 369 374 margolis al et revisited journal of parasitology 83 GOLDBERG SRS R CRC R BURSEY BKB K SULLIVAN AND QA 575 583 TRUONG 1996c helminthshelminthes of the sonoran gleengreen CABALLEROCABALLLRO DELOYA J 1974 estudio helmmtologilohelmintol6gico de toad bufo retiformisretiformis bufonidae hornfromhormhomm southernsou thein ari-an- los anianimalesanimatesmales silvestressilvestrassilvestres de la estacion de biologia zona journal of the helminthological society of tiotropicalpical los tuxtlas veracruz nematoda I1 algunos washington 6312063 120 122 nemanematodesnematodosnem1todostodos parasites de bufo horriborrihornbihshorribilisbilis wiegmann GOLDBERG SRS R CRC R BURSEY AND H CHEAM 1998 nema- 1833 anaces anales del institute de BioloBiologiagiagfa universidaduniveisidad todes of the cleatgleatgreat plains narrownainal i ow mouthed toad gas nacional aut6nomaautonomaAutautonomyonoma de mexico 454545 45 50 trophryne olivolivaceaolwaceaacea mieroMicromicrohyhdaemicrohylidaehylidae from southern CAMPBELL RA 1968 A comparative study of the para- arizona journal of the helminthological society of sites of certain salientia from pocahontas state paikparkpalk washington 6510265 102log 104 virginia virginia journal of science 191319 13 20 HARDIN ELE L AND J JANOVY JR 1988 population dynam- DICKEY LBL B 1921 A new amphibian cestode journal of ieslesics of distoichometra bufonishufomsbunonis cestoda nematotacninematotaem parasitology 71297 129 136 idae in bufo woodhousiiwooclhousii journal of parasitologyPaiaparasitology 74 DOUGLAS LTL T 1958 the taxonomy of nematotaenndnematotaeniid ces 360 365 todes journal of parasitology 4426144 261 273 KOLLER RLR L AND AJA J GAUDIN 1977 an analysis of hel- GOLDBERG SRS R AND CRC R BURSEY 1991a helminthshelminthes of minth infections in bufo boreas amphibia bufonidae toads three bufo alalvanusdarlusdariusvarius bufo cognatiiscognatescognatuscognatus bufonidae and hyla regilla amphibiaAmpbibia hylidae in sousouthernthein and scaphiopus couchiscouchii pelobatidae from southernsouthein california southwestern naturalist 2150321 503 509 arizona journal of the helminthological society of KUNTZ RER E 1941 the metazoan parasitespaiapalasitessltes of some okla- washington 5814258 142 146 homa anura proceedings of the oklahoma academy 1991b helminthshelminthes of the red spotted toad bufo of science 213321 33 34 ptinctatuspunctpunctatusatus anura bufonidae from sousouthernthein ari-an- PARRY JEJ E AND AWA W GRUNIMANNGRUNDMANN 1965 species compo- zona journal of the helminthological society of sition and distribution of the paiaparaparasitessitessltes of some com- washington 5826758 267 269 mon amphibians of iron and washington counties GOLDBERG SRS R CRC R BURSEY AND I1 RAMOS 1995 the utah Procprocedingsproceedingsedings of the utah academy of aiartsts and component parasitepaiapatasite community of three sympatric sciences 4227142 271 279 toad species bufo cognatescognatuscognatus bufo debilisdebihsdabilis bufonidae STEBBINS RCR C 1985 A heidheldfield guide to western reptiles and and speaaspea multiplicatamultiphcatamultiplicate pelobatidae from new mex- amphibians houghton mifflin company boston icoleoieo journal of the helminthological society of wash- 336 appp ington 625762 57 61 STEINER G 1924 some nemas flomfrom the alimentary tract GOLDBERG SRS R CRC R BURSEY EWAE WA GERGUS BKB K SUL- of the carolina tree foghogbogfrog hyla carolinensis pennant LIVAN AND QA TRUONG 1996a helminthshelminthes from journal of parasitology 11 1 32 three treetreefrogsfrogs hyla areniarenicolorcolor hyla wrighwrightorumtorum and pseudacnspseudacris tntrisesenatariatafiataflata hylidae flomfrom arizonaanzona received 16 march 1998 journal of parasitology 8283382 833 835 accepted 11 may 1998 gleatgreat basin naturalist 592 199901999 appp 198 200

HELMINTHS OF THE MADREAN ALLIGATOR LIZARD ELGARIA KINGII SAURIA ANGUIDAE FROM ARIZONA

stephen R GoldgoldberglGoldberggoldbergegoldberg1berglbergi1 charlescharies R bursey2 and hay chearlcheamlcheam1

key words elgariaelgaiidalgaria kingnkingiikingbi anguidae helininthshelminthshelminthes arizona

the madreanmadiean alligator lizard elgariaalgaria kingiikingbi harwood 1930 travassos 1931 spauligodon gray 1838 occurs from the southern edge of goldbergigoldgoldbergebergi bursey and mcallister 1996 physa the central plateau of arizona southward in loptera sp larvae and skrjabinoptera sp lar- the sierra madre of mexico to jalisco mexico vae prevalence mean intensity range and it frequents chaparral oak woodland and mean abundance are given in table 1 elganaelgariaalgaria pine fir forests and occurs from 760 to 2070 m kingiikingnkingbi is a new host record for each helminth stebbins 1985 there are no accounts of species helminthshelminthes from this species the purpose of none of the helminthshelminthes found in this study our paper is twofold to provide the first was unique to elganaalgariaelgaria kingiikingbi gravid individu- report of helminthshelminthes from E kingiikingbi collected in als of the following 3 species were found arizona and to furnish a list of parasites known oochoristicaoochonstica eumecis was originally described from the genus algariaelgariaElgaria from the skink eumeceseumenes fasciatusfasciatus from texas we borrowed 31 E kingiikingbi from arizona harwood 1932 and has been reported from from the herpetology collection of the univer- ctenosaura pectinatepectinatapectinata from mexico flores sity of arizona UAZ tucson mean snout barroeta et al 1958 E kingiikingbi is the ard3rd host vent length 68 mm 828.2 s range 48 80 mm record cosmocercoides dariadatlavariavanabihsvariabilisbilis is known the lizards were originally preserved in 10 from a variety of amphibians and reptiles from formalin and stored in 70 propanolisoisopropanol spec- north america baker 1987 E kingiiktngnkingbi repre- imens examined are listed by county of collec- sents the 26th host record spauligodon gold tion in appendix 1 the body cavity was opened bergi was 01 igmallyoriginally described from the ground and the gastrointestinal tract excised by cut- snake sonora semisemiannulataannulata from central texas ting across the esophagus and rectum the by bursey and mcallister 1996 E ktngnkingiikingbi is esophagus stomach and small and large in- the and2nd host record testineste were slit longitudinally and examined three species of helminthshelminthes were represented separately under a dissecting microscope the by immature forms tetrathyndiatetrathyridia of mesocesmesones body cavity and liver surface were also exam- toidesboides sp occur commonly in the coleomic ined each helminth was removed to a tempo- cavities of lizards and snakes which are con- rary glycerol mount for examination nema- sidered to be paratparatenicparatonicparatemcparateenicenleeniemc hosts bolette 1997 todes were identified from these temporary adults of physaloptera and skrjabinoptera are mounts cestodescustodesCestodes were stained in hematoxylinhernatoxylinhaematoxylin frequently seen gastric parasites of lizards and mounted in canada balsam for identifica- baker 1987 because only ard3rd stage larvae of tion voucher specimens were deposited in these 2 genera were found their importance the US national parasite collection appen- to the helminth load of E kingiikingbi cannot be dix 2 terminology usage is in accordance assessed with bush et al 1997 parasite lists table 2 can now be devel- algariaelgaria kingiikingbi harbored 2 species of cescestodescustodestodes oped for 3 of 4 species of elganaelgariaalgaria occurring in mesocestoides sp tetrathyridia and oochobocho north america namely E coeruleacoerulea E kingiikingbi risticamistica eumeciseurnecis harwood 1932 and 4 species and E multicannatamulticarinatamulticarinatemulticannata E panamintpanamintinapanaimntinainalna has not of nematodes cosmocercoides variavariabilisbilis yet been examined there is some overlap in

I1 department otof biology whittierwhittler college whittierwhittler CA 90608 21epartmentlepai tineiit of biology pennsylvania state university Sheshenangonango campus 147 shenangoShe nango avenue sharon PA 16146

198 1999 NOTES 199

TABLE 1 helminthshelminthes from elganaelgariaalgaria kingiikingbi N 31 from arizona prevalence mean intensity mean abundancece helminth N s range s site

CESTODA mesocestoides sp tetrathyridiatetrathyndia 47 3 470 1521 52 8448 44 coelom oochoristicaoochonstica eumecis 1 3 10 0030 03 0180 18 small intestine NEMATODA cosmocercoides variavariabilisbilis 8 13 202.02 0 202 0 1 5 026096ogg0 26 0930 93 large intestine spauligodon goldbergigoldbergegoldbergi 40 10 13313.313 3 10210 2 6 25 1291 29 4794 79 small large intestines physaloptera sp larvae 34 35 31 32 1 11 lioiio1101 10 2392 39 stomach small large intestines skrjabmopteraskrjabinoptera sp larvae 36 6 18018.018 0 18318.318 3 5 31 ilg1161 16 5615 61 stomach

TABLE 2 prevalence of helminthshelminthes in species of elganaalganaelgariaalgaria from north america species of elganaelgariaalgaria helminth location prevalence reference E coecoerulearulea mesocestoides sp tetrathyridiatetrathyndia contracontiacostacocacosta co CA not given voge 1953 cosmocercoides sp whitcomwhatcom co WA 22104921042 2104 coldgoldbergGoldbeigbelg and bursey 1991 oswaldocruzia sp whitcomwhatcom co WA 111041 1104 goldberg and bursey 1991 E kingiikingbi mesocestoides sp tetrathyridiatetrathyndia gila co AZ 31313 131 this study oochoristicaoochonstica eumecis santa cruz co AZ 31313 131 this study cosmocercoides variavariabilisbilis cochise pima cos AZ 1343113 431 this study spauligodon goldbergigoldbergegoldbergi cochise co AZ 1033110 331 this study physaloptera sp larvae cochise graham pima cos AZ 35113135 1131 this study skijabinopteraskrjabinoptera sp larvae cochise co AZ 62316 231 this study E multicarinatamulticannatamulticarinatemulticannata buenettaBaeriettaerlettaenetta gerrhonoti los angeles co CA 64162564 1625 telford 1970 mesocestoides sp tetrathyndiatetrathyridia riverside co CA 723079307 230 telford 1970 oochoristicaoochonstica sp los angeles co CA 11961 196 goldberg and bursey 1990 physaloptera retusaaretusa riverside co CA 1343013 430 telford 1970 physaloptera sp larvae los angeles co CA 11961 196 goldberg and bursey 1990 oswaldocruzia pipienspipkens los angeles co CA 229692962 296 goldberg and bursey 1990

the helminth genera harbored table 2 but newfoundland occasional papers in biology 11 too few helminthshelminthes have been found to evaluate 1 325 BOLETTE P the helminth in species El DPD 1997 first record of pachysentispachijsentisbachyPachysentis canicola community of algariaelgariagaria acanthocephala oligacanthorhynchida and the except for the 64 1625 prevalence of the occurrence of mesocestoides sp tetrathyndiatetrathyridia ces- cestode baeriettamaeriettaBaerietta gerrhonoti reported by toidea cyclophyllidea in the western diamondback telford 1965 preprevalencesvalences of helminth species rattlesnake crotalus atrolatrox serpentes viperidae of algariaelgaria are low journal of parasitology 8375183 751 752 BURSEY CRC R AND CTC T mcalastermcallistermcalhster 1996 spauligodonspauhgodon goldbergigoldgoldbergebergi sp n nematoda pharyngodonidae and we thank charles H lowe department of other parasites of sonora semisemiannulataannulata serpentes ecology and evolutionary biology university colubridae from new mexico and texas journal of of arizona for permission to examine speci- the helminthological society of washington 636263 62 65 mens of algariaelgaria kingiikingbi for helminthshelminthes BUSH AOA 0 KD lailatLAFFERTYFERTYJMJM LOTZ andawshostakannAND AW SHOSTAK 1997 parasitology meets ecology on its own terms margolis et al revisited journal of parasitology 83 literature CITED 575 583 FLORES BARROETA L E HIDALGO AND RRR R BRENES BAKER MRM R 1987 synopsis of the nematoda parasitic in 1958 cestodosc6stodosCesgestodos de Vertebravertebradosdos IV revistadevista de BiBioloologiagiagfa amphibians and reptiles memorial university of tropical 6556 55 78 200 GREAT BASIN naturalist volume 59 goui3timgoigol oindinoln RC SRS R AND CRC R BURSEYBURSLY 1990 helminthsheirninthshelminthes of APPENDIX 1 thetiletlletite san diego alligator lizard gerrhonotus multi natus wehhiwebbi anguidae journalnalnai of wildlife caricarinatus wethi join dis- museum accession numbers for specimens ofElof elganaalganaofelgariagariagarfa 26 297 eases 26297 298 kingiikingitkingbi N 31 from the university of arizona UAZ 1991 gastrointestinal helminthes helminths oftheodtheof the northwest- listed by arizona county cochise UAZ 15501 15691 alligator gerrhonotus ern haidhaldlizard coercoeruleusuleus pnncipisprincipisprincipesprincipis 36871 37918 37920 39487 39711 39715 40032 40033 Anguidanguidaeciecle journal oftheodtheof the Helminthologichelminthologicaldl society 403084093840308 40938 40939 40941 40942 40944 40947 46845 of washington 5824658 246 248 47297 48012 gila UAZ 36731 40309 graham UAZ HAKWOODHARWOOD PD 1932 the helminthshelrninthshelminthes parasitic in the 36362 39710 43862 fimapimapuma UAZ 11248 19773 19774 amphibia and reptilia of houston texas and vicin-vicin santa cruz UAZ 119904917111990 49171 ity proceedings oftlicof the USU S national museum 81181 1 71 STEBBINSS 11 liiilns RCR C 1985 A field guide to western reptiles and amphibians houghton mifflin company boston 336 appp APPENDIX 2 tltiixoiwI1 I1 ornowdown SRS R jilJK 1965 A new nematotaenndnematotacniid cestode flom califoimacalifbrniacalifornia lizards japanese journal of experi- accession numbers for helminthshelminthes of elganaelgariaalgaria kingiikingnkingbi mental medicine 3530135 301 303 deposited in the USU S national parasite collection meso 1970 A comparative study of endoendopaiasitismendoparasitismparasitism cestcestoideacestoidesoides sp 87667 oochoristicaoochonstica eumecis 87664 cos among some southern california lizard populations tnocercoidesmocercoides variavariabilisbilis 87665 spauligodon goldbergigoldgoldbergebergi amelameiicanamericanAmei ican midland naturalist 8351683 sig516 554 87666 physaloptera sp 87668 skrjabinoptera sp voivolvocvoo 1 M 1953 new host records foiforbol mesocestoides ces- 87669 toda cyclophyllidea in california amerleanamericanametAmei icanlean midland naturalist 4924949 249 251

received 27 february 1998 accepted 11 may 1998 great basin naturalist 592 019991999 appp 201 203

CHARLES LYNN HAYWARD

herbert H frostlfrosti and wilmerwiimer W tannerlnannerltanner1

dr C lynn hayward professor emeritus of zoology and entomology brigham young university provo utah died 30 august 1998 he was born 10 july 1903 in paris idaho where he spent his early years graduating from fielding high school in 1923 later that same year lynn entered brigham young uni- versityversity receiving a BS degree in 1927 fol- A lowing his university experience he returned to his homehometowntown in idaho and taught biology and english at the high school for 3 years in may 1930 he returned to BYU on an assistant- ship in the zoology and entomology department on 6 august 1930 he married elizabeth 7 libbie cook to them were born 2 children margaret and gerald in 1931 he completed his master s degree in entomology at BYU and became a zoology instructor there lynn hayward entered BYU as the department of zoology was being established and though it was first designated C lynn hayward as the zoology and entomology department his role was to establish and teach basic zoo- logical classes pertaining to vertebrates from and mammal collections at BYU in the early 1931 to 1942 he taught in the department and 1930s with the depression in full force during the beginning years began research in equipment funds were limited having been preparation for a doctoral degree in 1935 he raised in a rural community he knew how to did graduate work at the university of califor- work with his hands and so obtained plans for nia at berkeley and in 1941 completed his dis- the construction of bird and mammal cases sertationsertation in ecology at the university of illi- and built several for the growing collections nois under professor victor shelford one of today the mammal collection contains 15621 the pioneer ecologists his phd graduate skins and whole mount specimens studies emphasized systematics and ecology of lynn had a good grasp of the english lan- birds and mammals guage thus his lectures and research writings during his early teaching years at fielding were clear and meaningful he engaged his high school lynn made a collection of 94 students in biological field studies that required bird skins and several sets of eggs these careful observations and ecological evaluations eventually became the nucleus of BYU s orni- of the environment many of his field classes thologicalth collection which now numbers 9551 involved preparing bird and mammal skins as bird skin specimens and 9000 sets of eggs the students became proficient in mounting because of his collecting background lynn their best specimens became part of the uni- assumed the position of curator of the bird versity s collections

onielmonteonte L bembean litelife science museum brigham young university fiovopiovo UT 84602

201 202 GREAT BASIN naturalist volume 59

he was always willing to work with his stu- 9 1935 observations on some breeding birds of dents and his door was open for them to come mount Timpanotimpanogosgos utah wilson bulletin 4716147 igi161 162 to him for advice he chaired 18 students graduate committees and was a member for 10 1935 A study of the winter bird life in bear lake and utah lake valleys wilson bulletin 67 many more his graduate students were well 278 284 trained and have become recognized in their 11 1936 some observations on shore birds at utah fields his research is best seen in two publi- lake during the summer of 1936 utah acad- cationscations biotic communities of the wasatch emy of science 1319113 igi191 193 chaparral and alpine biotic communities of 12 1936 A bibliography of utah mammalogy includ- the uinta mountains utah his bibliogra- ing references to names and type localities phy of the mammals of utah and birds of applied to utah mammals utah academy of science 1312113 121 146 utah have been used and will continue to be 13 vespa used as resource materials by many who 1937 Aaiecordofvespacrabrorecord of crabro linnaeus from north are dakota entomological news 4812048 120 conducting research in vertebrate zoology of 14 1937 some new and unusual bird records from western north america utah wilson bulletin 6930369 303 305 in addition to his teaching and research 15 1940 notes on the distribution of nightnighthawkshawks in lynn had over the years a number of admin- utah great basin naturalist 1931 93 96 istraistrativetive responsibilities at BYU for 7 years hebe 16 1940 feeding habits of the red squirrel journal of served on the admission and credits commit- mammalogy 2122021 220 tee a year on the curriculum committee and 17 1941 notes on the nesting habits of some moun- many years as chair of the premedical and tain dwelling budsbirds in utah great basin nat- predental committee he chaired the zoology uralist 212 1 8 department from 1958 to 1962 and was for 5 18 1941 three new mammals microtus and ochotona years curator of the life science museum fromflornhrom utah great basin naturalist 21052 105 108 with ERE R hall which was the forerunner of the monte L 19 1941 A bean museum on 30 august 1974 he was bibliography of utah mammalogy including references to names and type localities presented the BYU alumni distinguished first supplement great basin naturalist 2 service award for teaching and research dur- 125 136 ing his professional years he wrote and collab- 20 1942 biotic communities of mt Timpanotimpanogosgos and orated on 147 articles the following 42 western uinta mountains utah an abstract papers covering a 53 year period are consid- of a thesis 11 appp ered to be his most important contributions 21 1943 notes on the status of the red crossbill in utah auk 6027660 276 277 22 1944 additional records of uncommon birds in condor 46 1 1930 notes on utah vespidae entomological news utah 46205205 4120441 204 205 222 226 23 1945 biotic communities of the southern wasatch and uinta mountains utah great 2 1931 A pieliminaryprepreliminary list of the birds of the sub- basin naturalist 6 1 124 alpine and alpine zones of the uinta moun- 61 tains utah academy of science 81518 isilsi151 152 24 1945 occurrence of perognathus faciatusfaciatus in utah journal of Mmammalogyamm alogy 37 4511 3 1932 the paper wasps of utah including a descrip- 3745 tion of a new varietyvaival lety of polistes canadensis 25 1948 biotic communities of the wasatch chaparral linn utah academy of science 9859 85 101 utah ecological monographs 336 appp 4 1933 notes on the taxonomy and description of the 26 1949 the short tailed weasel in utah and colo- wasp genus polistes in the intermountain rado journal of Marnmamammalogylogy 3033630 336 west with description of two new varieties 27 1949 nesting behavior of nuttall s poorwillpoor will wil- utah academy of science 10 139 147 son bulletin 6118861 188 5 1933 distribution of polistes in canada with dotesnotes 28 1951 nature sanctuaries in the united states and on the genus hymenopHymenop canadian ento- canada living wilderness 364636 46 with sev- momologistlogist 6512665 126 128 eral other authors 6 1934 important heron rookeriesrookeriedrookeries in southeastern 29 1952 alpine biotic communities of the uinta Mmoun-oun idaho auk 513951 39 41 tains utah ecological monographs 22 7 1934 A biological study of the la sal mountains 93 120 utah academy of science 1120911 209 235 with 30 1956 pelage color changes in perognathus longi VM tanneitanneltanner membris journal of mammalogy 3745137 451 452 8 1935 the bi cedingbreeding status and migration of the 31 1958 additional notes on the purple martin in caspian tern in utah condor 3714037 140 144 utah condor 6040660 406 0it1jahy0111tuary 203

32 1958 distribution and variation of the utah popu- 37 1966 new and unusual budbird records from utah lation of the gleatgreat basin pocket mouse gleatgreat condor 6830568 305 306 basin naturalist 1826 30 with L merlin 38 1967 budsbirds of the upper killpack colorado river basin BYU science bulletin biological series 92 33 1958 new and unusual records of budsbirds from the 1 64 uinta basin utah great basin naturalist 18 39 1968 natural history of the desert woowoodratwoodraldrat neotonui 23 25 with merlin L killpack lepido american midland naturalist 80 34 1958 zoology of the upper coiocoloradoCololadoiddo river basin 458 476 with robertrobertc C stones I1 the biotic communities BYU science bul- 40 1970 vasco M tanneitanneltanner creatgleatgreat basin naturalist 30 letin biological series 74 with 137413 D elden 181 189 beck and wilmerwiimer W tanner 41 1976 birds of utah great basin naturalist mem 35 1963 budsbirds of the nevada test site BYU science oilairsoirss 1 bi ighambighambrigham young univeiuniveruniversitysitssity piprovoovo UT bulletin biological series 301311 1 27 with 3l131 229 appp with clarenceclaienceciarenceclarenee Cotcottamtarn AMA M wood- meilinmerlin L killpack and gerald L richards bury and heiberthelbertherbert frost 36 1965 mammals of the nevada test site BYU sci- 42 1983 the high uintascintas monte L bean life science ence bulletin biological senes 1 seriesserles 63163 81 museum brigham young provopi with clive jorgensen university ovo UT 101 appp GREAT BASIN biological RESEARCH conference 14 16 october 1999 university of nevada reno

the university of nevada reno will host the first great basin biological research conference 14 16 october 1999 the conference will feature con- tributedtri buted presentations and posters keynote speakers and roundtable discussions related to the biological study and management of the great basin and eastern sierra nevada ecoregionseco regions also several field trips have been planned

this conference will provide an exciting opportunity to present and discuss biological research in a regional context and to share ideas with other profes- sionals working throughout the great basin and adjacent blomes As a biogeo- graphical region this area possesses a varied and unique biotic landscape that presents many management challenges and that has been used as a model system for investigating numerous ecological phenomena

authors are invited to submit posters and presentations involving biological research on the great basin or eastern sierra nevada mountains topics for paper sessions include but are not limited to physiology evolutionary biology population and community ecology conservation biology biogeography biology and management of riparian areas cellular and molecular biology behavior genetics management of nevadas biotic resources and paleobiology

papers will be allotted 12 minutes each with an additional 3 minutes for questions 15 minutes total posters 5 x 6 max size allow authors to engage in more detailed and personal communication with attendees abstracts must be less than 250 words and submitted by 18 june 1999 electronically via the conference website see below

registration is via the conference websitewebsite wwwbrrcedugbbrccom early registration deadline is 16 july 1999 40 for students 60 for non students late registration 55 and 75 respectively information FOR AUTHORS the great basin naturalist welcomes previously VOUCHER SPECIMENS authors are encouraged to unpublished manuscripts pertaining to the biologi- designate properly prepare label and deposit cal natural history of western north america high quality voucher specimens and cultures docu- preference will be given to concise manuscripts of menting their research in an established permanent up to 12000 words simple species lists are dis- collection and to cite the repository in publication coucouragedraged references IN THE TEXT are cited by author and SUBMIT manuscripts to richard W baumann date eg martin 1989 or martin 1989 multiple editor great basin naturalist 290 MLBM PO box citations should be separated by commas and listed 20200 brigham young university provo UT in chronological order use et al after name of 84602020084602 0200 an accompanying cover letter must first author for citations having more than two include phone numbers of the author submitting authors the manuscript and FAX number and emailE mail acknowledgments under a centered main address when applicable the letter must also pro- heading include special publication numbers when vide information describing the extent to which data appropriate text or illustrations have been used in other papers literature CITED also under a centered main or books that are published in press submitted or heading lists references alphabetically in the fol- soon to be submitted elsewhere authors should lowing formats adhere to the following guidelines manuscripts not mack GD and LD flake 1980 habitat relation- so prepared may be returned for revision ships of waterfowl broods on south dakota manuscript general great preparation in the stock ponds journal of wildlife management follows basin naturalist recommendations in 44695 700 style and CBE Manu por scientific format the manualformanualalforalnorainorfor sousa WPWE 1985 disturbance and patch dynamics authors editors and publishers ath6th edition on rocky intertidal shores pages 101 124 in cambridge university press 110 midland avenue STA pickett and PSES white editors the ecolo- port NY 10573 USA TOLL chester TOLLFREEFREE PHONE gy of natural disturbance and patch dynamics 1 800 872 7423 we do however differ 18008727423 in our academic press new york treatment of entries in authors literature cited coulson RN and witter 1984 forest ento- may consult great JA the most recent issue of the mology ecology and management john wiley basin naturalist for formatting guidelines and sons inc new york 669 appp TYPE AND DOUBLE SPACE all materials including literature cited table headings and figure legends TABLES are double spaced on separate sheets and avoid hyphenated words at the righthandright hand margins designed to fit the width of either a single 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per page is made for articles than 15 words names and addresses of authors a published the rate for individual subscribers will running head of fewer than 40 letters and spaces be 35 per page however manuscripts with com- footnotes to indicate change of address and author plex tables andor numerous photographs may be to whom correspondence should be addressed if assessed an additional charge reprints may be pirpur- other than the first author chased at the time of publication an order form is ABSTRACT states the purpose methods results sent with the proofs and conclusions of the research it is followed by FINAL CHECK 612 key words listed in order of decreasing cover letter explaining any duplication of importance to be used for indexing information and providing phone numbers TEXT has centered main headings printed in all FAX number and emailE mail address capital letters second level headings are centered 3 copies of the manuscript 5 copies of fish in upper and lowercase letters third level head- papers and WordwordperfectPerfect diskette ings begin paragraphs conformity with instructions photocopies of illustrations issn0017ISSN 001736140017 3614 GREAT BASIN naturalist vovolwolwoi 59 no 2 april 1999 CONTENTS articles reproductive ecology of bison on antelope island utah michael L wolfe milan PE shipka and john FE kimball 105 remarkable waxing waning and wandering of populations of mimulus guttatusgustatusguttatus an unexpected example of global warming robert K vickery jr 112 biogeography and physiological adaptations of the brine fly genus ephydra diptera ephydridae in saline waters of the great basin david B herbst 127 stream temperatures and elevational distribution of redband trout in southwest ern idaho bruce W zoellick 136 effect of salinity on seed germination of triglochin maritinamarimaritimatima under various tem- peratureperature regimes M ajmal khan and irwin A ungar 144 an irvingtonianIrvingtonian species of brachylagus mammalia lagomorpha from porcupine cave park county colorado colleen N ramos 151 A new species and new synonym in the genus psychoroniaPsychoronia limnephilidaelimnophilidae with significant records for caddisfliescaddisflies trichoptera from western north america david E ruiter 160 emergence patterns of large stonefliesstoneflies plecoptera pteronarcys calineuriaCalineuria hesperoperldhesperoper1a in a montana river andrew L sheldon 169 competition and niche partitioning among pseudoroegneria spicataspicatespicata hedysarum boreale and centaurea macumaculosamaculoselosa james S jacobs and roger L sheley 1751 75 A new species of aleciaplecia from the creengreen river formation and new combinations of fossil bibionidae diptera scott J fitzgerald 182 assortative mating in soldier beetles cantharidae chauliognathus test of the mate choice hypothesis ruth bernstein and stephen bernstein 188 notes body size dynamics of cougars felis concolor in oregon stephan G kohlmann and richard L green 193 helminthshelminthes of the lowland burrowing treetreefrogfrog ptyernohyla fodientfodiens hylidae from southern arizona stephen R goldberg charles R bursey and guillermo galindo 195 helminthshelminthes of the madieanmadrean alligator lizard algariaelgaria kingiikingbi sauria anguidae from arizona stephen R goldberg charles R bursey and hay cheam 198 obituary charles lynn hayward herbert H frost and wilmer W tanner 201