Great Basin Naturalist

Volume 55 Number 1 Article 13

1-16-1995

Full Issue, Vol. 55 No. 1

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GREATR EAT BASINBAS I1 N naturalist

A

VOLUME 55 nigNIQ I1 JANUARY 19951995

BRIGHAM YOUNG university GREAT BASIN naturalist editor assistant editor RICHARD W BAUMANN NATHAN M SMITH 290 MLBM 190 MLBM PO box 20200 PO box 26879 brigham young university brigham young university provo UT 84602020084602 0200 provo UT 84602687984602 6879 8013785053801 378 5053 8013786688801 378 6688 FAX 8013783733801 378 3733 emailE mail nmshbllibyuedunmshbll1byuedu

associate editors MICHAEL A BOWERS PAUL C MARSH blandy experimental farm university of center for environmental studies arizona virginia box 175 boyce VA 22620 state university tempe AZ 85287

J R CALLAIIANCALLAHAN STANLEY D smiSMITHTH museum of southwestern biology university of department of biology new mexico albuquerque NM university of nevada las vegas mailing address box 3140 hemet CA 92546 las vegas NV 89154400489154 4004 JEFFREY J JOHANSEN PAUL T TUELLER department of biology john carroll university department of environmental resource sciences university heights OH 44118 university of nevada reno 1000 valley road reno NV 89512 BORIS C kondratieff department of entomology colorado state ROBERT C WHITMORE university fort collins CO 80523 division of forestry box 6125 west virginia university Morganmorgantowntown WV 26506612526506 6125

editorial board berranjerran T flinders chairman botany and range science duke S rogers zoology william hess botany and range science H duane smith zoology all are at brigham young university ex officio editorial board members include steven L taylor college of biology and agriculture stanley L welsh director monte L bean life science museum richard W baumann editor great basin naturalist the great basin naturalist founded in 1939 is published quarterly by brigham young university unpublished manuscripts that further our biological understanding of the great basin and surrounding areas in western north america are accepted for publication subscriptions annual subscriptions to the great basin naturalist for 1995 are 25 for individual sub- scriscribersbers 30 outside the united states and 50 for institutions the price of single issues is 12 all back issues are in print and available for sale all matters pertaining to subscriptions back issues or other busi- ness should be directed to the editor great basin 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

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copyright 0 1995 by brigham young university ISSN 001736140017 3614 official publication date 16 january 1995 1951 95 750 12935 the great basin naturalist PUBLISHED AT PROVO UTAH BY BRIGHAM YOUNG university

ISSN 001736140017 3614

VOLUME 55 31 JANUARY 1995 no 1

great basin naturalist ssi551 0 1995 appp 1 18 LIFE HISTORIES OF stonefliesSTONE FLIES plecoptera IN THE RIO CONEJOS OF SOUTHERN COLORADO

R edward DeWaltdewalt12dewa1ti212 and kenneth W StewartStewartiartl1

ABSTRACT thirty one stonstoneflyefly species representing eight families were collected during the march 1987 to may 1990 study period genera represented by more than one species included capniaacapnia Utacapnia taenionematahtaemonemaTaeniTaeonemamonema Suwallia triznakatnznakaTriznaka Isogenisogenoidesoides and isoperla peak species richness was recorded on or near the summer solstice in 1988 and 1989 climatic differences between years were reflected in nymphal development and emergence phenology of most species new or important corroborative life history data are presented for 11 stonstoneflyefly species of this assemblage the hyporheichyporheic nymphal development of most chlorochloroperlidperlid species limited the number of early instarsmstars sampled and our capacity to interpret voltinism limited nymphal data suggested a univoltine slowsiow cycle for plumiperlaPlumiperla diversa frison adults of walliasuwalliaSu pallidulapallidula banks and S wardi banks were present for an extended summer period but the bulk of their respective emergence times was temporally separated isogenoidesIsogen oides zionensis hanson pteronarcella badia hagen and pteronarcys californicacalifornica newport were all shown for the first time to have a 9 lomo10 mo egg diapause and all three species have a semivoltmesemivoltine life cycle sawalaskwala americanaamencanaamencana klapalekklapdlekKlapalek and isoperlafulvaisoperla folva claassen were further confirmed to have univoltine slow cycles univoltine fasthast and univoltine slowsiow life cycles are reported for the first time in I1 phaleratephalerataphalerata and I1 quinquepunctatequinquepunctata respectively regression analysis revealed that six of the eight abundant species had extended emergence patterns slopes of ad5d while only two had synchronous patterns warmer spring and summer tempera- tures in 1989 increased the slopes for five of the eight species studied but did not change their synchrony designation nine of I111 I1 abundant species advanced their median emergence date in 1989 over 1988 this and the higher slope values are consistent with a hurried nymphal development and narrower emergence period due to the warmer thermal regime of 1989

key words plecoptera life history diversitybiobiodiversity life cycle rocky mountains

Stonestonefliesflies plecoptera are one of the integral understood sheldon and jewett 1967 stewart and often dominant insect orders in stream and stark 1988 precise life histories are ecosystems therefore they are important as known for 5 of the more than 575 north biological indicators as fish food and as part american species and knowledge of stonstoneflyefly of the energy and nutrient economy of streams life histories and ecology in southern rocky stewart and stark 1988 taxonomy of the mountain streams is sparse this has limited north american fauna is now well known our ability to increase understanding of eco- however information on their life histories logical relationships between cohabiting stone local species richness and ecology is still poorly fly species in this region

1 departmentidepartment of biological sciences university of north texas denton TX 76203 2presentpresent address department of zoology and physiology louisiana state university baton rouge LA 70803

1 2 GREAT BASIN naturalist volume 55

one objective of this study was to deter- highway 17 and 4 km north of antonito at the mine richness of the stonstoneflyefly assemblage of colorado highway 285 bridge respectively the rio conejos of southern colorado a large stream temperatures varied from below drainage that has not been previously studied freezing during the winter months to near second we documented the important life 20c20gocgog C in august ice cover was common from history events of its dominant species for december through march snowbeltsnowmeltSnowmelt began which sufficient individuals and observations in april usually leading to peak flows in june could be gathered by intensive monthly sam- base flows were attained by late august and pling and by living streamside during spring continued through the winter water released and summer from platiroplatoro reservoir 48 km upstream aug- research was patterned after the classic mented river flow during summer low flow studies of harper 1973a 1973b and harper periods bottom substrates were characterized and hynes 1972 who studied a substantial by large boulders cobble gravel and sand portion of the eastern canadian fauna and these were covered by a thin layer of silt in addressed critical aspects of life histories such quiet water important organic substrates as egg development diapause and adult included the flooded coppicescoppicuscoppices of willows and behaviors that are often overlooked H B N cottoncottonwoodswoods and their entrained leaf packs hynes in an address to the international ple- willow salix sppapp cottoncottonwoodswoods and ashensaspens coptera symposium 1992 emphasized the populus sppapp and alder alnus sp con- need for more attention to these aspects to tritributed to the riparian corridor support the eventual development of a para- physical conditions digm of life history evolution within the ple- copteracoptera we have also adopted the approaches stream temperature was monitored at site of knight and gaufin 1966 harper and one from june through august 1988 using a ryan TM magnin 1969 sheldon 1972 barton 1980 continuous recording thermograph low ernst and stewart 1985a 1985b and high and mean daily stream tempera- from hassage and stewart 1990 in comparatively tures were calculated temperatures recorded at 0400 0800 1200 1600 2000 and studying an assemblage of species this report 2400 h water temperatures were not record- is the first to address on a large scale such an ed during 1989 due to equipment failure assemblage in a western american north however summer air temperature highs and stream since the works of knight and gaufin lows and rainfall were recorded 1300 h daily 1966 sheldon 1972 and stanford 1975 mountain time for both 1988 and 1989 at the conejos peak US weather service reporting METHODS station at site one flow data for site two were petsch study stream gathered from 1987 90 nymphal growth the rio conejos is located in the southern rocky mountains of south central colorado nymphs were collected monthly except the river flows east to west for 145 km from december due to poor weather conditions at its headwaters in the rio grande national all sites from march 1987 to may 1988 forest of the san juan range to the rio grande additional collections were made at irregular 32 km northeast ofantonitoof antonito CO three sam- intervals until march 1990 samples were col- pling sites were established along the rio lected by disturbing the substrate mineral conejos to ensure access to at least one of and organic upstream of a bioquip rectangu- them during the winter and to enhance collec- lar dionetdipnet until debris clogged the net the tion of stonstoneflyefly species that were not abundant net was composed of a coarse imm1 mm mesh at all sites these were located at elevations be- first stage modified by the addition of a coni- tween 2400 and 2600 m above sea level the cal second stage of 153 amtm mesh size the lat- primary site 10615w longitude 3703n ter collected even the smallest instarsin stars A latitude consisted of a 1 km stretch located 24 plankton bucket was attached to the second km west of antonito conejos county CO off stage to facilitate sample removal contents of colorado highway 17 sites two and three the plankton bucket and the coarse stage con- were located 22522.5 km west of antonito also on stituted a sampling unit and were stored in 199519951 STONEFLY LIFE HISTORIES 3

70 isopropyl alcohol the number of sam- the malaise trap was deployed among wil- pling units per month varied with the effort low and cottonwood coppicescoppicuscoppices where its olive necessary to secure approximately 50 nymphs drab coloration mimicked the surrounding of all abundant species vegetation flying or crawling adults inter- nymphs were separated from sample cepted by the trap ascended the screening debris with the aid of 4 lox magnification on into a dry apical collection chamber addition- a stereo dissecting microscope sorted to ally all adults on the trap mesh were collected species when possible and stored in 80 using an aspirator ethanol until measurement head capsule emergence traps were anchored over shal- width HCW greatest distance across the low fifflesriffles during the 1988 field season eyes was measured with a calibrated ocular natural diurnal changes in water level and micrometer fitted to a stereo dissection micro- erratic discharges due to water release from scope nymphs from all sites for the 3 yr sam- platiroplatoro reservoir rendered these ineffective pling period were pooled by species and at times therefore their use was discontinued month of collection to increase the number of in 1989 nymphs per month and to allow construction pitfall traps consisted of 283 cm2 modified of more robust growth histograms gender of aluminum soda cans that were buried flush in nymphs was assessed by a gap in the posterior streamside substrates A mixture of 70 setal margin of the eighth sternum of females ethanol and ethylene glycol the latter to stewart and stark 1988 and by developing retard evaporation was used as a preservative external genitalia of females sex specific kite in 1988 12 traps were installed I1 m from the diagrams were constructed by placing male stream at 1 m intervals on an open beach with and female nymphs into 010.1oi or 02 mm size nearby vegetation this was expanded in 1989 classes the frequency of these classes was to three transects each consisting of 30 cans converted to a percentage of the total number set 1 m apart in transects 1 m 5 m and 8 m of nymphs males females unsexed from the initial shoreline these traps moni- nymphs collected for that month polygons tored not only adult presence of ground tra were constructed for each month depicting versing brachypterous stonefliesstone flies but also the relative proportion of all nymphs at that their potential to move laterally from the size class stream sweepnettingSweepnetting was conducted over a 15 X 2 adult emergence m willow and cottonwood riparian zone the adults of winter and early spring ebergemerg entire area was methodically swept working ing stonestonefliesflies were collected from bridge abut- from the base of each clump of vegetation up- ments from shoreline debris and under the ward exuviae removal was the only method cobble at streamside to provide a general used to assess emergence of claasseniaClaassenia sabu emergence period for each species adults losa banks and was used for no other species were also reared from emergentpreemergentpre nymphs in 1988 exuviae were removed daily from the A combination of sampling methods and same 15 X 1 m area of cobble shoreline and observational procedures was used during the the frequency of each sex was noted in 1989 summers of 1988 and 1989 to evaluate emer- the removal area was expanded to 30 X 1 m of gence duration of adult presence and behav- shoreline area and up to 5 m into the water for ior of these species adult traps and methods collecting exuviae from emergent substrates included a 225 m2ma basal area bioquip malaise year and sex specific kite diagrams of adult trap two 025 m2ma basal area floating emer- presence were produced for all abundant sum- gence traps pitfall traps sweepnettingsweepnetting of mer stonefliesstone flies by pooling all methods and streamside vegetation exuviae collection and expressing daily catches as a percentage of the day and night transect walks pitfall traps were total catch duration of emergence of eteroptero emptied on alternate days and the others narcys californicacalifornica newport would be greatly were emptied daily between 0900 and 1100 h overestimated by including pitfall trap collec- all of these methods were used at site one tions due to its synchronous emergence and sweepnettingsweepnetting was employed at site three on since pitfall traps were emptied on alternate several occasions days 4 GREAT BASIN naturalist volume 55

dates of first capture 50 cumulative mm plastic petri dishes in an environmental catch and last collection plus total duration of chamber in both instances these were incu- adult presence were determined for the 11 bated at approximate stream temperature and most abundant species collected in the sum- light regime mers of 1988 and 1989 emergence synchrony fecundity was estimated from number of was estimated using linear regression of the egg batches deposited number of eggs per cumulative percentage catch all methods batch and for sawalaskwala americana klapaiekklapdlekKlapaieklek pooled versus days since first capture slopes only total number of eggs remaining in the generated for each species were used as an ovaovariolesrioles females were housed at streamside index of synchrony steeper slopes indicated a in screened glass containers and provided more synchronous emergence slopes 5dad with moist cotton balls as a source of water were chosen to be indicative of synchronous alternatively some species were reared in emergence since species with these slopes denton and held under simulated streamside emerged their entire population within a few conditions in large cotton stoppered shell days and had steep j shaped cumulative emer- vials gence curves differences between slopes for RESULTS 1988 and 1989 were tested using a modified t test zar 1984 common slopes were calculated physical conditions if no differences between years were noted mean daily stream temperatures in 1988 this was a purely descriptive approach de- increased from near IOCloc10 in early june to signed to detect and compare patterns there- 15c15 in mid july fig 1 the stream cooled fore it is not our aim to model emergence for dramatically between 8 and 12 july this coin- the purpose of prediction but only to describe cided with cool damp weather conditions patterns of emergence fig 2 summer air temperature highs rarely since most adult collection methods em- exceeded 30c30 in 1988 and rainfall occurred ployed in this study collected adults of unknown at regular intervals throughout the summer age results reflected adult presence rather fig 2 however 1989 was marked by many than in the strictest sense emergence no days above 30c30 with rainfall relegated to late attempt was made to discard old males and july and august fig 2 the mean monthly dis- females using any index of age however pat- charge of the rio conejos during 1987 1989 terns of adult presence should follow that ofI1 a fluctuated predictably peak discharge true emergence pattern and since longevity of occurred typically in june but occurred in may during the warm windy spring of 1989 most adults approached only 1 wk in the labo- 3 ratory we believe these results to be useful fig behavioral observations were made from species richness 0800 to 1300 h and from 2000 to 2300 h for sev- more than 13000 nymphs and adults were eral days during emergence of each species studied over the 3 yr period among these observations made during intervening hours adult behavior of adult produced little timing 20 activities their relative distance from the teanhean low hlj h stream and substrates on which activities took 18 place were monitored by walking the stream P 2 16

logs rocks J margin turning and and exposing v bases marginal W A leaf entrained of vegetation E of these observations have been details narra- E tively described for each species in this paper fecundity and egg incubation 10 eggs of several species were incubated in 8 confirm the laboratory to proposed voltinism 6 based on growth histograms eggs were 64 614 6124 74 7114 72247124 83 813 placed into I1 cm diameter dialysis tubing dates bags and reared in a frigid units living fig 1 daily mean high and low stream temperatures in stream or they were stored in 100 X 15 the rio conejos summer 1988 199519951 STONEFLY LIFE HISTORIES 5

36 262.625 including an early and a peaked 1988 waning more T so- distribution of species richness in 1989 e 2 R M 26 a P leuctridae e 20 high 16 r f paraleuctra ververshinavepshinashina gaufin and ricker a 16 low t this was the only leuctrid found at our sites U 10 r no nymphs were recovered from the stream 6 rain C e og indicating a probable hyporheichyporheic existence 060.6 M C 0 AA adults were abundant in riparian vegetation A 6 during june and july fig 5 no variation in 527627 66 616 626 76 716 726 86 815816 826 adult presence parameters was noted for P dates ververshinavepshinashina table 2 emergence was classified 40 2 as extended in both years although slopes of 1989 high T these cumulative emergence curves were sig- e 30 R M 16 nificnificantlyantly different over the 2 yr table 3 P 1 e 20 1 r low f a 1 a t iol10- 1 u I1 TABTABLELE 1 stonefliesStoneflies collected from the rio conejos r 0 og colorado march 1987 through march 1990 e 060.6 C M C rairaln Euholognatha 10 & CAPNIIDAE capniaacapnia coloradensis claassenlClaassenclaassensclaassen11 5161 611 621 631 610 620 630 710 720 730 89 dates capniaacapnia confusaconfuso claassen capniaacapnia vernalis newport fig 2 daily high and low air temperatures and rainfall isocapnia arinitacrinita needham & Claassen 1 for summer 1988 and 1989 Utacapnia logana nebeker & Gaufingaufinlgaufin11 Utacapnia poda nebeker & Gaufingaufin11 leuctridae paraleuctra ververshinavepshinashina gaufin & rickerl 60 nemouridaeNEMOURIDAE e3ea MEAN AMAXIMUMmaximumAMAXIM 0 O MINIMUM amphinemura bancsibanksi baumann & gaufinlgaufin1 1 F 50- Prprostoiaostoia besemetsabesemetsa riekerrickerlRickerricker1rickerd 1 1 0 zapadafrigida Claassenclaassenlclaassensclaassen1 40 taeniopterygidae m3ma taenionemathenionemaTaenionema pallipallidumdum Bankbankssl P taenionemathenionemaTaenionema pacificuspacificumpacificum banks I1 e 30 r doddsia occidentoccidentalisoccidentalistalis BankBanksbankslbankelbanks1sl1 a systellognatha 20 C chloroperlidaeCHLOROPERLIDAE 0 n paraperlafrontalisbaraparaperlaParapetlapetiaperla frontalis Bankbanksbankslbankelsl d 10 plumiperlaPlumiperla diversa prisonFrisofrisonnl Suwallia lineosafineosa Bankbankssl 0 Suwallia pallidulapallidula baniBankbankssl MONTH 10 12 2 4 6 8 10 12 2 4 6 8 10 12 2 4 6 8 Suwallia wardi kondratieff & Kirchkirchnernerlneri YEAR 1987 1988 1989 triznakaTriznaka pintado rickerriekerrickerl fig 3 mean minimum and maximum monthly stream triznakaTriznaka signata Bankbankssl discharge of the rio conejos during the study period PERLIDAE claasseniaClaassenia sabulosesabulosasabulosa BankBanksbankslbankelbanks1sl1 hesperoperla pacifica Bankbankssl perlodidae 31 1 were species table in eight families the isogenoidesIsogen oides zionensis hansonhansonl chloroperlidaeChloroperlidae perlodidae and capniidae isogenoidesIsogen oides prob colubrinuscolubrinus hagenhagenlhageni were the most speciose families with six isoperlafulva claassen seven and seven species respectively seven isoperla monnonamannonamormona banks by isoperla phalerataphaleratephalerata smithsmithl genera were represented more than one isoperla quinquepunctataquinquepunctate banks species capniaacapnia Utacapnia taenionemathenionemaTaenionema suwal sawalaskwala americana klapalekklapdlekKlapalek lia triznakaTriznaka Isogenisogenoidesoides and isoperla table 1 pteronarcyidae peak species richness occurred on or near pteronarcella badia hagen the summer solstice in both years fig 4 pteronarcys californicacalifornica newportNewportlporti pattern differences existed between years lneinenew drainage and county records 6 GREAT BASIN naturalist volume 55

12

s p lo10 e C i 8 e S

r 6 i C h 4 n e

2

0 519 529 68 618 628 78 718 728 87 817 dates

fig 4 temporal species richness pattern of adult stonestonefliesflies collected daily from the rio conejos during the summers of 1988 and 1989

Chlorochloroperlidaeperlidae data suggested a univoltine slow life cycle for this species from two subfamiliessubfamilies in- representatives suwalliaSu pallipalliduladula banks chloroper and mid wallia habited the stream the early instar linae only 59 nymphs of Suwallia sppapp were nymphs of the chloroperlinae genera could collected from the rio conejos even though not be reliably identified to genus this neces- adults were abundant nymphs were hyporhe sitated the illustration of a portion of the ic until immediately prior to emergence this nymphal growth of Plumiplumiperlaperla diversa frisonfrisonprison habitat preference and our present inability to and triznakaTriznaka signata banks as chloroperlinae distinguish convenerscongenerscongeners of Suwallia nymphs sppapp fig 6 growth of reliably identified mid precluded generation of meaningful his- to late instar nymphs was illustrated separately tograms and designation of voltinism for either Paraparaperlaperla frontalis banks paraperlinae species adults of Suwallia wardi kondratieff nymphs were collected infrequently among & kirchner were consistently larger than S marginal substrates during the colder months pallidulapallidula this trend followed in nymphs too of the year all were pale very thin and had with proposed female nymphs of S wardi in eyes set far forward as described for mature june peak emergence being 22022.0 larger nymphs stewart and stark 1988 these limit-

ed data are presented for the first year of the paraleuctra vershinaververslmnawerweevepshinashinasbona presumed semivoltinesernivoltine growth pattern of this M F large chlorochloroperlidperlid fig 6 less than 10 adults 25 of total catch 3658 were collected in early june during the 3 yr 1988 study no plumiperlaPlumiperla diversa chloroperlinae 2434 adults were collected on which to base specif- 1989 0 406waw ic identity however nymphs of this genus are distinctive and only P diversa has been col-

1 1 1 1 lected in this region baumann et al 1977 1 1 1 nymphs were identifiable to genus by march 20 30 9 19 29 9 19 29 females were readily distinguished from MAY JUNE JULY growth continued through males at this time fig 5 emergence of paraleuctra ververshinavepshinashina from the rio may when females attained a median HCW conejos 1988 and 1989 polygons indicate daily relative 969.6gg larger than males the limited nymphal proportion of total catch 199511995 STONEFLY LIFE HISTORIES 7

TABLE 2 range of dates for adult presence parameters for 11 summer emerging stonstoneflyefly species collected in 1988 appears first and 1989 from the rio conejos duration is mean SD of the number of days all parameters not avail- able for C sabulosasabulosesabulosa which emerged past our study period date ist date last date species n capture 50 catch capture duration d P dercervercershinavershinavepshinashina 94 2 june 12 june 5 july 35035.035 0 000.0oo0 0 58 1 june 17 june 7 july

S wardi 467 6 june 10 july 15 august 66066.066 0 7177.11 352 19 may 25 june 18 july

S pallidulapalhdulapallidula 276 30 june 28 july 23 august 44544.544 5 484.84 8 162 1 july 14 july 4 august

T signata 662 9 june 28 june 23 august 59059.059 0 24024.024 0 2697 2 june 19 june 12 july

C sabusabulosesabulosalosa 356 19 july 1195 16 july

I1 fulda 19 9 june 22 june 7 july 24524.524 5 787.87 8 61 9 june 18 june 28 june

I1 phalerataphaleratephalerata 12 20 june 28 june 24 july 22522 5 353 5 20 18 june 25 june 8 july

LI1 quinquepunctataquinquepunctateqmnquepunctata 9 24 june 14 july 27 july 30030.03004242424.2 12 19 june 5 july 15 july

I1 zionensisztonensis 200 8 june 19 june 28 june 15515.515iss 5 494.94 9 75 10 june 17 june 24 june

P badia 215 10 june 22 june 17 july 34534.534 5 353.53 5 480 7 june 20 june 7 july

P calicallcailfornicacalifornicacahfomica 55 6 june 8 june 12 june 606.0go6028282.8 21 4 june 5 june 13 june than the july peak emergence females of S august table 2 it had the longest mean pallidulapallidula only two proposed male nymphs of duration of presence 66 d for any stonstoneflyefly the latter were collected over the 3 yr period studied on the rio conejos table 2 like its adults of S pallidulapallidula were collected in july congener S wardi s 1989 date of median catch and august in both years fig 7 table 2 was advanced by 2 wk over that of 1988 table slopes from regression models were different 2 fig 7 emergence of S wardi was extend- between years t 64764.7 p 0001.0001 but ed and no significant slope differences were below the 5dad criterion we categorized this noted between years table 3 no egg data species as an extended embergeremerger table 3 the were collected for this species median emergence date was advanced by 2 riparian vegetation was used by this large wk in 1989 over that of 1988 table 2 the yellow green chloroperlidchloroperlid as a staging ground adult sex ratio over the two seasons was 13 6S for adult behaviors walliasuwalliaSu wardi was active 415 Y six field collected and laboratory throughout the morning on sunny days and maintained females produced only one egg again for 2 3 h before sunset if conditions batch table 4 were warm and dry during cool rainy days Suwallia wardi chloroperlinae this was the low vegetation was devoid of S wardi or the most abundant of the three walliasuwalliaSu any other stonstoneflyefly species species collected from the rio conejos adults triznakaTriznaka signata banks chloroperlinae were first collected in late may or early june identifiable late instar nymphs were collect- reached 50 cumulative catch by mid july ed during a 5smomo period in the spring and and disappeared from streamside by early summer nymphs of this univoltine slow 8 GREAT BASIN naturalist volume 55

TABLE 3 synchrony and linear regression statistics for 0 2 individuals 50 of monthly catch the years 1988 appears first and 1989 slopes between 10 08 i 4 f ovipsitionowposrtionoviposition n 83 years were tested significance 05010501.050105 01 001 06 04 emergence level or lower probability and NT not tested 02 V emergence 4 pareperlaParaParereraparapertareeananenanaperlapertapardapaddapaasapansa frontgronttrontalisalls species slope r2fi2ra P synchrony n 453 P ververshinavepshinashina 30 85 0001 extended 08 d K 0 35 90 0001 extended 04 IF E plumiperlaPlumi perlapeolapenta diversadeversa E f T signata 15 87 0001 extended s 4 14 t nn653653 30 91 0001 extended 12 yioylo510 08 ttlmakatriznakaTriztotznakamaermawr signals S pallidulapallidula 25 96 0001 extended 90 0001 extended 08 nn694 35 06 0 S wardi 84 0001 extended 02 22 v achloroperlinaeae sppapp 22 85 0001 extended 1I1 II11 liililIII111 IV V VI vilVII villVIII IX X XI months C sabulosasabulosesabulosa 28 94 0001 extended 99 0001 extended 41 fig 6 growth of Chlorochloroperhdaechloroperlidaeperlidae nymphs collected from the rio conejos 1987 1990 I1 zionensis 75 92 0001 synchronous 79 95 0001 synchronous they were collected in abundance during the P badia 36 81 0001 extended day adults inhabited marginal vegetation 44 92 0001 extended where males were observed actively searching willow stems and leaves for females no drum- P calicalffornica 133 84 004 synsynchronosynchronysynchronouschronouS was observed during the two summers 189nt 97 103 synchronous ming of intensive fieldwork large flights of adults of both sexes took place just before dark at species were largely full grown by april fig which time females were observed oviposit 6 with some degree of sexual dimorphism ing egg masses were dropped from up to 4 5 present at this time m above the stream first appeared early reached adults in june perlidae 50 cumulative catch 2 wk later and could no longer be collected by late august fig 7 claasseniaClaassenia sabulosasabulosesabulosa although two perlissperlids table 2 emergence was protandrous but were present in the rio conejos table 1 slightly female skewed sex ratios dominated in only C sabulosasabulosesabulosa was sufficiently abundant for both 1988 and 1989 fig 7 triznakaTriznaka signata growth and emergence interpretation nymphs displayed the greatest variation in last date of of this species were found among larger rub- capture and duration of presence of all stone ble of midstream the life cycle was seminolsemivol flies in the river table 2 it advanced its 1989 tine and appeared to require 3 yr of nymphal median emergence date by 9 d over that of growth fig 8 recruitment occurred 1988 regression slopes indicated an extended throughout the fall with possibly some addi- emergence in both years table 3 differences tional recruitment in march from overwinter between slopes for 1988 and 1989 were signif- ing eggs eggs containing eyespotseye spots were icant t 113511.35 p 0001 recovered from the stream in october and attempts during the entire study to obtain november sexual dimorphism in both size eggs from laboratory reared and mated and external genitalia occurred when nymphal females were unsuccessful the mean number size reached 222.22 2 mm HCW the size disparity of eggs from six females caught during ovipo- increased until the third year of growth when sition flights was lower than any first batches little overlap between the sexes remained for other stonestonefliesflies studied table 4 although A protandrous emergence began in mid these females were held for a prolonged peri- july in both years fig 9 table 2 exuviae of od of time no additional egg batches were this species were abundant throughout august laid possibly into september emergence of C adults were never seen emerging in the sabusabulosasabuloselosa was extended and slopes were signif- field despite many hours of observation along icantly different between years t 10710.710 7 p the shoreline day and night in habitats where 0001 table 3 199511995 STONEFLY LIFE HISTORIES 9

2 ind 1 5 of total catch swsex ratioraboraho perlodidae I MF ti&1781ra triznakasignatasignata 1988 63446gg 9 c j7qcj5fcift422- jtxa gy Isogenisogenoidesoides zionensis hanson Perloperlodinaedinae C 432634461989 118115151181 1515 Perperlodmiperlodinilodini large range in size of nymphs libilibl the at july samples 10 could not be SuSuwallwalliamalliamagliata wardwaad from fig 4t 1988 US119348348 accounted for by nymphs hatching from eggs

1989 150191 laid by june mated females june eggs reared

suwatiasuwalliaSuwaliawatiawallia pallidulapallidula at simulated stream conditions hatched in A 1988 1325313263 aa A march and 9 10 mo after oviposition 6agm aw&wtwfysvvfftlmfalm i april therefore at least some individuals of this 1989 0162 speciesIs have a semivoltine life cycle with eggs diadiapausingpausing over their first summer and winter 20 30 9 19 29 9 19 29 8 18 28 may june july august early instar nymphs were missed in benthic samples during their second spring possibly fig 7 emergence of Chlorochloroperhdaechloroperlidaeperlidae from the rio due to high water or their occurrence deep in conejos 1988 and 1989 polygons indicate daily relative the substratum sexual dimorphism in size proportion of total catch and morphology was apparent by july of the second year when nymphs approached 181.8 mm HCW fig 10 this disparity increased claasseniaClaassenia sabulosasabulosesabulosa produced the greatest steadily throughout the rest of their growth mean number of eggs of any stonstoneflyefly species little overlap in size of the sexes existed by studied table 4 with females producing up to may prior to emergence four batches longevity of seven females was the adult presence parameters of I1 zionenzionek 393.9 191.9iglg d egg production lasted through sis showed little variation over the 2 yr studied 80 of the adult life several egg batches were table 2 emergence was not protandrous incubated but none hatched within 6 inomo of but the sex ratio was heavily skewed towards observation males fig 11 this species was one of two emergence occurred between 2000 and that emerged synchronously table 3 no dif- 2200 h nymphs crawled out of the water onto ference in slope was found between years t emergent cobble and boulders to transform 0820.82 p 2.2 therefore a common slope of the entire boltingmolting process taking less than 5 76d was calculated min hardened and newly transformed males laboratory reared females put nearly 75 ran over all emergent substrates searched for of their total egg complement into a first batch females in a circular pattern and drummed table 4 only one of four females produced mostly on large mineral substrates pitfall trap additional batches collections of 1989 caught a total of 115 male transformation of I1 zionensis took place adults in transect I1 and only 12 in transects 2 from 2030 to about 2200 h nymphs crawled and 3 only two females were collected in the away from the stream until they reached wil- pitfall traps presumably because of their less lows or other vegetation then ascended 1 m intensive and unidirectional movement pat- vertically where they boltedmolted daylight activity tern therefore excursions of great distance began by 0700 0800 h at the base of small wil- away from the water s edge for either sex were low coppicescoppicuscoppices where adults were often found infrequent females were often found in the in emergent leafleafpackspacks adults ascended stream- morning under dry cobble with abdomens de- side willows as the sun rose drumming mat- void of eggs or with large egg masses suspend- ing and egg batch formation took place from ed between the cerci several females were these perches females crawled to the tops of observed at night running over the surface of these willows and flew to the stream where the water but the cause of this behavior could they fluttered on the water to release their not be determined no females were actually black egg masses most activity ceased by observed ovipositing males were distinctly 1300 1400 h on days when air temperature cursocursorialriall which fits with their brachypterous reached near 25c25 C on cloudy cool days this morphology however females were never ascendance did not occur most adults could observed flying nor did they inhabit tall sub- then be found in the leaf entrained bases of strates even though they had full wings riparian vegetation drumming on willow stems 10 GREAT BASIN naturalist volume 55

TABLE 4 mean eggs per batch number of batches and mean total egg complement for nine species of stonestonefliesflies occurring in the rio conejos colorado eggs batch n species 1 2 3 4 total

S pallidulapalhdulapallidula 54754 7 26626.626 6 54754 7 26626.626 6 6 6

T signata 42242.242 2 17417.417 4 42242.242 2 17417.417 4 6 6

C sabulosasabulosesabulosa 2166216602166.0 0 7740774774.00 9020902.0902 0 2462246246.28462 2 1580158.0158 0 91091.091gio 0 40040.040 0 3188318803188.0 0 6130613.0613 0 7 5 5 1 7

LI1 zionensis 5880588.0588 0 86086.086 0 3270 1850 8432843843.22 1414141 4 4 1 1 4

LI1 fulvabulva 2315231 5 787.87 8 2315231.5231 5 787.87 8 2 2

I1 phalerataphaleratephalerata 7030 7030 1 1

S ameticanaamericanaametiamencanaamencana 8847884.7884 7 2673267.3267967 3 8847884884.77 2673a267 3311 6 6

P badia 3390339.0339 0 86086.086 0 58458.458 4 37237.237 2 56856.856 8 39739 7 3510351 0 loi1010101.0101 0 30 5 4 30

P californicacahfomicacalicalffornica 3930393.0393 0 1256125 6 1913191.3igligi191 3 1302130130.22 94394.394 3 49549.549 5 69869.869 8 24724 7 4 4 4 4 5 6 7

51351 3 29329.329 3 58558.558 5 23323.323 3 570 8453845.3845 3 90590.590 5 4 2 1 4 rotaltotal fecundity includes those eggs remaining in ovaovriolesanoles was observed at night even when tempera- fecundity was difficult to assess since few tures approached loc10 C mature nymphs were available for rearing isoperla fulvabulva claassen isoperlinae we one egg batch from each of two field oviposit collected this species in benthic samples only ing females was collected table 4 longevity occasionally but enough individuals were of three field collected adult females was 575.7st obtained to allow a tentative interpretation of t 0580.58 d voltinism recruitment of nymphs was first isoperla phalerataphaleratephalerata smith isoperlinae detected in august fig 12 these measured although the number of nymphs collected was 040.4 080.8 mm HCW and grew at a slow rate small no month supported more than one size throughout the fall until a winter decrease in class fig 12 therefore we have tentatively growth rate their size increased dramatically proposed a univoltine slow growth pattern for after february until emergence in june and this species adults were taken from mid june july this species conformed to a univoltine through mid july table 2 fig 11 no assess- slow growth pattern ment of synchrony was made for LI1 phalerataphaleratephalerata adults were collected for the first time on 9 due to low numbers of adults captured june in both years fig 11 table 2 sex ratios females did not produce eggs in captivity A for the small number of 1988 adults were single egg batch from a field collected individ- approximately equal but heavily skewed ual contained 703 eggs four field caught towards males in 1989 numbers ofadultsof adults col- females lived 11311.3 363.6 d past date of capture lected in both years were too small to warrant isoperla quinquepunctataquinquepunctate banks isoderisoper an analysis of synchrony linae this species was more common at site 199511995 STONEFLY LIFE HISTORIES 11

alaClaclaasseniaclaassemaClaasasseniasewasema sabulosesabulosasabulosa alaakaClacfaasseniaclaasseniaassenia sabusabulosesabulosalosa H 20 off anthlymonthlynthly catchwh 2 individual M F ebergemergemergencen 1 1 MF sanpan n 697 4 of total catch d 9 ow positionitioaitio 235129 6 1988 E E 6

803401 u3ua 1989 2

S y

I1 II11 iliIII111 IV V VI VII villVIII IX X XI months 9 19 29 8 18 28 july august fig 8 growth of ciaclaasseniaclaassemachaasClaaschaabClaasseniasema sabulosasabulosesabulosa nymphs collected from the rio conejos 1987 1990 fig 9 emergence of ciaclaasseniaclaassemaClaasClaasseniasema sabulosasabulosesabulosa from the rio conejos 1988 and 1989 polygons indicate daily relative three the data suggested that I1 quinquepunc proportion of total catch tata had a univoltine fast growth pattern recruitment occurred in january and february fig 12 and growth was rapid from march pteronarcyidae through may sexual dimorphism in nymphal pteronarcella badia hagen this species size was not as evident in this species as in its was found to have a semivoltinesernivoltine growth pat- convenerscongenerscongeners emergence began in mid june and tern recruitment of nymphs began in march lasted through much of july table 2 fig 11 and april from eggs laid the previous june no eggs were collected fig 14 many small nymphs were available sawalaskwala americana klapaiekKlapaklapdlekieklek Perloperlodinaedinae in benthic samples by mid april when they this species displayed a univoltine slow were at 020.2 040.4 mm HCW this scenario was growth pattern and grew faster during sum- corroborated by laboratory incubation of several mer and fall months than all other perlodids in egg batches that hatched in march and april the rio conejos fig 13 nymphs were re- after a 9 lomo10 mo diapause growth of nymphs cruited in june and increased their median was rapid throughout their first spring size HCW from 040.4 mm to about 282.8 mm by differentiation among sexes was not apparent january growth was nearly completed by this until august a full 14 mo after oviposition time sexual dimorphism was apparent as early median size of females just before emergence as august and female nymphs reached a medi- the following may was 21 greater than that an HCW before emergence that was 21421.4 of males greater than males female nymphs in april emergence began by early june with slight were found to contain fully sclerotized eggs in protandry and a preponderance of males being their oviductsoviducts hence this species is fully collected fig 15 median emergence occurred capable of mating and egg laying immediately in the third week of june in both years table upon emergence 2 emergence was extended table 3 and emergence was in april and early may when slopes were significantly different between our sampling was still on a monthly basis years t 222.2 p 05os05.05 therefore no detailed analysis of emergence females generally laid only single egg phenology and synchrony can be offered batches but a small number produced up to adults were collected mainly from emergent three egg batches table 4 most females laid logjam debris or under cobble at the stream their first egg batch within 24 h of mating and margin often waited 2 d intervals before laying others egg batches collected in mid april from longevity of seven females under simulated four laboratory reared females hatched syn- field conditions was 777.7 424.2 d chronously after a mean of 61061.0glogio 737.3 d this pteronarcella badia emerged just after dusk corroborates field collections of early instar and typically used willows cottoncottonwoodswoods and nymphs in june only a single egg batch was stream margin sedges as transformation sites collected from each of six laboratory reared males were observed actively searching the females table 1 willows and drumming for females at night 12 GREAT BASIN naturalist volume 55

6 isogenoldesIsogenoldes z1onens1s 1 IND perlodidae

emergence wposition 20 of total catch 5 M F E 5 of monthly catch isogenoidesisogenoldesIsogenoldesoides zbnensiszionensis E 9 1988 14655 4 individuals n 421 Ccf 1989 5318 c13 eulvafulwa isoperla fulka 1988000 1190 o 00 bp99136pfii3 CT 2.2 e 1989 537 laboratory recruitment isoperla abatahaiphaphalphaleratephalerataphalarataeretaeratalarataeneta 1 1988 48 of previous years eggs ag3ct 1 171 1989 713 isoperla quinquepunctatequinquepunctata 1 11 illiliiiilil111 IV V VI VII villVIII IX X XI 1988 months 36 I1 & 1989 66 fig 10 growth of Isogenisogenoidesoides zionensis nymphs col- lected from the rio conejos 1987 1990 9 19 29 9 19 29 june july even when air temperatures were near 101c10 C fig 11 emergence of perlodidae from the rio adults entered the leaf choked bases of wil- conejos 1988 and 1989 polygons indicate daily relative lows as the night progressed and were often proportion oftotalof total catch found the next morning in large mating aggre- these debris individuals gations under these adults utilized marginal vegetation much ascended the willows as the sun warmed the as did P badia as a staging ground for mating air at streamside drumming mate searching and ovipositing however they tended to select mating and egg batchinghatching took place in mid- the taller cottoncottonwoodswoods and engelmann spruce morning hours while most activity ceased by rather than the shorter willows for their activi- 1200 h when air temperatures reached ties salmonfliesSalmonflies positedovioviposited by flying over the 22 25c25 C females positedovioviposited by launching stream and dropping their salmon colored or themselves from the tips of tall riparian shrubs bluish egg masses dimorphism in egg color was toward the stream where they would jettison observed from as high as 10 in ovipositingOvipositing their white egg mass a few meters above the adults were heavily fed upon by opportunistic surface of fast flowing water eastern robins turdus migratormigratoriesmigratoriusius and pteronarcys californicacalifornica newport this steller s jays cyanocitta stelleristelspellerileri species is commonly known as the salmonsalmonflyfly recruitment began in april fig 16 after a discussion 9 lomo10 mo egg diapause nymphs grew to only about I1 mm HCW through their first year species richness differences size and morphology sexual in the rio conejos displayed a great diversity were apparent by june of their second year of plecoptera twenty of the 31 species were when they were nearly 151.5lsis mm HCW evenly distributed among the capniidae the nymphs grew for two more years by the end chloroperlidaeChloroperlidae and the perlodidae the only of which time emergentpreemergentpre females had north american family not represented was attained a 20 larger median HCW than the peltoperlidaePeltoperlidae which occurs transcontinen males these data suggest a semivoltinesemivolvoitine life tally but not in latitudes below the northern history ofofaof44 yr duration for this species rocky mountains baumann et al 1977 nearly adults were first found on 6 june during all species collected were adapted for a mon- both years fig 15 table 2 emergence was a tane existence and were characteristic of highly synchronous event table 3 slopes streams with high biotic integrity twentyfivetwenty five were not tested for significant differences due species were both new drainage and county to small sample size records baumann et al 1977 szczytko and most laboratory reared females produced stewart 1979 nelson and baumann 1989 table five egg batches but one individual produced 1 though all of them had been previously seven table 4 egg production lasted reported from colorado and neighboring new through 82 of the 15015.0iso 181.8 d n 4 aver- mexico this demonstrates that we have yet to age adult female life span adequately investigate the fine scale diversity 199511995 STONEFLY LIFE HISTORIES 13

isoperlinae sppapp 4 ebergemerg go amala americana 141 41 30 of monthly catch 41 T faponoviposibon positionositositionionlon 1 individual emergence vpon 3 6 C isoperaisopertafulvaguevaguava n 105 E eg 9 isopea E 3 E J a P 3 am lsopmlaisopertaIsotsopertaperdapedda obatphatphalphaleratephalerataerataaradaanadala T T EBB 2 2 p if ebn RIa tat3 nn2525 F i 30 of monthly catch 333 V soperlaboperlaisoperla qiqepwquinquepunctatequinquepunctata 3 individuals 2 n 227 n229n 229

11 111 I1 II11 liiIII111 IV V VI VII villVIII IX X AXI III IV VI VII villviliVIII IX X AXI months months

fig 12 growth of isoperla sppapp nymphs collected from fig 13 growth of skwalasawala americanaamencanaamencanocana nymphs collected the rio conejos 1987 1990 from the rio conejos 1987 1990 and distribution of this order of aquatic insects L ferrugineaferrugiferrugineannea walker snellen and stewart in at least some portions of the southern 1979 record univoltine fast cycles for rocky mountains zealeuctra claasseniclaassensclaasseni and Z hiteihidei in streams of responses to altered thermal regime north texas additionally ernst and stewart 1985a report leuctra tenuis as univoltine fast we became aware of substantial climatic in an ouachitaOuachita mountain stream differences fig 2 between the two summers when adults were intensively studied though chloroperlidaeCHLORO PERLIDAE no water temperatures were available for most chloroperlidaeChloroperlidae exhibit a univoltine 1989 air temperatures fig 2 and hydrologic slow or fast growth pattern haploperla brevis data fig 3 suggested that the stream warmed banks is widespread from oklahoma to more quickly and attained peak summer highs quebec and west to alberta canada ontario much earlier than in 1988 consequently harper and magnin 1969 quebec harper development of several species was hurried et al 1994 and oklahoma ernst and stewart which narrowed the window of time adults 1985a populations exhibited univoltine fast were present streamside at the assemblage growth with a 2 5smomo diapause while alberta level of organization this trend is demonstrat- populations were univoltine slow barton ed by the species richness pattern of figure 4 1980 european populations of chloroperlaChloroperla the 1989 pattern was more peaked and great- tripunctatatripunctate scopoli elliott 1988 siphonssiphono ly truncated over that of 1988 species level perla torrentitorrentiumum pichetPipictetctet elliott 1967 and S responses can be demonstrated by inspection burmeisteri pichetPipictetctet benedetto 1973 also of the flight diagrams for each species nine of exhibited univoltine slow growth species the 11 species presented in table 2 show in- with semivoltine growth include sweltsa creased median emergence dates additionally onkos ricker and possibly utaperlaUtaperla gaspe slopes produced by linear regression that siana harper and roy harper 1973a harper were different between years table 3 were et al 1994 S mediana banks cushman et al S always higher in 1989 this result was consiscansisconsis- 1977 and lateralis banks huryn and with a hurried nymphal development and wallace 1987 tent gaufin shorter emergence period for each species Paraparaperlaperla frontalis stanford and 1974 presented some evidence for seminolsemivol life history parameters tine growth of this species emergence for this species and for P wilsoniwilisoni ricker occurs from leuctridae may through july stewart and stark 1988 paraleuctra ververshinavepshinashina harper 1973b reports paraperlinae are rather robust chloroperlids that that most leuctra ferrugiferrugineaferrugineannea in an ontario tend to be hyporheichyporheic for most of their nymphal stream are semivoltine but that some univol- development their larger size the more sta- tine individuals exist huryn and wallace ble stream temperatures in the hyporheichypo rheic 1987 propose a 2 yr life cycle for a composite environment hendricks 1993 and the possi- of leuctra sppapp most of which were probably bly low availability of some nutrients in the 14 GREAT BASIN naturalist volume 55

emergence pteronarcella aadlabadla pteronarcyidae oviposition 20 of total catch 3 MFM F

E E t pteronarcys calftfnlcacalifcaliecatiearnicaornica 19881938 2332 2 C 1waw 1989 iilo111011 10 CL 4 pteronarcella badabadka n 1417 1988 13479134 79 40 of monthly catch

1989 308172 unsexed

1 II11 III111 IV V VI VII villVIII IX X XI months 30 9 19 29 9 19 29 fig 14 growth of pteronarcella badia nymphs collect- ed from the rio conejos 1987 1990 june july fig 15 emergence of pteronarcella badia and hyporheichyporheic habitat stanford and ward 1993 pteronarcys califcalifomicacahformcaomica from the rio conejos 1988 and may have contributed to a preponderance of 1989 polygons indicate daily relative proportion oftotalof total semisemivoltinismvoltinism in this subfamily catch plumiperlaPlumiperla diversa stewart et al 1990 re- slow cycle for this ported a univoltine species fact that 0 6 657 Y of the closely related S on the north slope of alaska emergence lineosa were caught during concurrent sam- may occurred from through september with pling on massey creek a tributary of the rio recruitment of nymphs from a direct hatch in conejos july growth occurred through the summer triznakaTriznaka signata hassage and stewart months with most nymphs attaining maximum 1990 studied the widely distributed T signata size before a winter quiescence this assess- in the rio vallecitos of northern new mexico ment compared well with our limited data they reported a univoltine slow growth pat- failure to collect adults was probably due to tern with which we concur no study of the our infrequent sampling during their pre- emergence of this species has previously been sumed early may emergence published Suwallia pallipalliduladula and Suwallia wardi no aspects of the life histories of either S pallidulapallidula PERLIDAE or S wardi have been reported the latter was claasseniaClaassenia sabusabulosasabuloselosa hassage and stewart recently described from a colorado front 1990 and barton 1980 report a merovoltine range springspringbrookbrook kondratieff and kirchner 2 yr growth pattern for new mexico and 1990 it was one of the most abundant chloro alberta populations of this species no egg perlissperlids in the rio conejos this suggests that batches from the rio conejos hatched in our its ecological tolerance is wide and that it may laboratory but this colorado population showed soon be found in a variety of streams in the some evidence of an extended hatch leading to southern rocky mountains cohort splitting stewart and stark 1988 eggs several explanations are possible for the may undergo a temperature dependent quies- heavily female skewed sex ratio 13 6425 Y cence as occurs in dinocrasDinodinocerascras cephalotescephacephalotuslotes curtis of S pallidulapallidula adults the most probable is a when fall temperatures decline to 8cac lille- combination of limited use of emergence traps hammer et al 1989 presence of first instar coupled with an inaccessible micromicrohabitathabitat of nymphs in the fall eyed eggs in october and adult males probably high in the vegetation november and more first instar nymphs in parthenogenesis may also be possible but it is march supported this contention exceedingly rare in stonestonefliesflies harper 1973a life histories have been reported for at least reported that a few eggs of a perlid parag one species in every genus in the tribe perlini netina media walker hatched without fertil- to which C sabulosesabulosasabulosa belongs all growth pat- izationization we did not attempt rearing of eggs terns involve 2 3 yr of development agnetinaagnesinaAgnetina from virgin females to check for parthenogen- flavescentflavescensflaves cens walsh from an ozark stream esis in either walliasuwalliaSu sppapp these sex ratios exhibits a 2 yr life cycle a short egg incuba- are a perplexing problem compounded by the tion period and an extended emergence period 199511995 STONEFLY LIFE HISTORIES 15

4 pterol7arcyscalifomicapteronarcys califomicacalifcalefcatieamicaomicaamaca occur in its convenerscongenerscongeners but this will be con- I1 A 20 of monthly catch ememergenceergen position 5 1 individual firmed only when detailed studies using small n622 mesh nets frequent sampling and egg rearing e4ea have been conducted E 1 fi isoperla sppapp of the three isoperla whose 3 d 9 partial growth patterns are presented here only I1 julda has been previously reported 2 NO hassage and stewart 1990 reported a anivolunivol tine slow cycle with a june emergence in the f t rio vallecitos of new mexico we concur with unsexed the new mexico study our results agree well 1 II11 III111 IV V VI vilVII villVIII IX X AXI of isoperla biology summarized months with reviews for 12 nearctic species through 1987 stewart fig 16 growth of pteronarcys califcahformcacalifomicaomica nymphs col- and stark 1988 ten species were univoltine lected from the rio conejos 1987 1990 slow while only two were univoltine fast in more recent literature stewart et al 1990 ernst and stewart 1985b agnetinaagnesinaAgnetina capitatecapitatacapitata reported univoltine slow growth for I1 petersolipetersonipetersoni pichetPipictetctet was shown to have a 3 yr cycle ex- needham & christenson ofalaska additionally tended emergence and a 40 80 d egg incuba- harper et al 1994 added as univoltine slow tion period in ontario harper 1973a this I1 francesca harper and I1 montana banks range of incubation coupled with a long emer- from quebec populations these and our rio gence promotes great differences in size of conejos work bring to 17 the nearctic isoperla nymphs that ultimately prevents the separa- species known to exhibit univoltine slow tion of cohorts and determination of voltinism cycles while only three species appear to be this was also a problem for C sabulosasabulosesabulosa in the univoltine fast isoperla grammatica poda rio conejos and 1 difdifformisformis klapalekklapdlekKlapalek palmqvistmalmqvistMalmqvist and perlodidae sjostrom 1989 and I17 obscuraobscure zetterstedt studied by elfstrandulfstrandUlfstrand 1968 are univoltine this family contains over 115 species stark slow in the palearctic et al 1986 stewart and stark 1988 in the up to seven species of isoperla commonly nearctic although life histories of only 26 occur in streams in north america stewart species are known a clear trend toward uni and stark 1988 conversely in scandinavia holtinevoltine slow cycles occurs among the subfami rarely more than two species occur simultane- lies isoperlinae and Perloperlodinaedinae stewart and ously palmqvistmalmqvistMalmqvist and sjostrom 1989 bongencongen stark 1988 growth and emergence had not erics of aquatic insects often partition resources previously been studied for three of the seven along one or more resource gradients grant perlodids in the rio conejos these include I1 and mackay 1969 though only small numbers zionensis I1 quinquepunctataquinquepunctate and I1 phaleratephalerataphalerata of adults were collected a pattern of succes- Isogenisogenoidesoides zionensis few detailed life his- sive emergence of I1 fulua I1 quinquepunctataquinquepunctate tory studies of the genus have been reported and I1 phaleratephalerataphalerata was clear in the rio conejos stewart and stark 1988 barton 1980 kupectsupect fifty percent cumulative catch dates for LI1 ed semisemivoltinismvoltinism for an alberta population of fuluajulua LI1 phalphaleratephalerataerata and LI1 quinquepunctatequinquepunctata were I1 colubrinuscolubrinus since two size classes of nymphs 22 june 28 june and 14 july respectively for were collected in early may flannagan 1977 1988 these dates for 1989 were 18 june 25 reported great body length variation in may for june and 5 july temporal segregation this species in another alberta watershed but brought about by a gradual change in domi- concluded a univoltine slow cycle hilsenhoff nance liliesillies 1952 of these species may have and billmeyer 1973 and dosdall and lehm- accounted for the presentpres ent coexistence of kuhl 1979 proposed univoltine growth pat- these stonestonefliesflies terns for the may june emerging I1 frontalis sawalaskwala americana two other studies re- in wisconsin and saskatchewan streams ported univoltine slow cycles with emergence respectively based on samples taken a few from february through april for this species in months of the year Semisemivoltinismvoltinism as reported northern new mexico and central colorado for I1 zionensis in the rio conejos may also short and ward 1980 hassage and stewart 16 GREAT BASIN naturalist volume 55

1990 sawalaskwala curvatecurvata hanson of california pteronarcys californicacalifornica the egg diapause also exhibited a univoltine slow cycle with plus 38 mo nymphal life span places total life emergence in april and may sheldon 1972 span of this population at 4 yr this is one of other arcynopterygini with univoltine slow the longest lived aquatic insects known to growth include frisoniafridoniaFrisonia picticeps hanson in occur in the nearctic additionally this california sheldon 1972 Megarmegarcyscys signata species is perhaps our most synchronously hagen in utah cathergather and gaufin 1975 emerging stonstoneflyefly and Perlperlinodesinodes aurea smith in california and two to 3 yr life cycles with a 9 lomo10 mo egg alberta radford and hartland rowe 1971 diapause occur in other pteronarcys such as P sheldon 1972 dorsatadoreata barton 1980 P proteus holdsworth sheldon 1972 estimated average total 1941a 1941b W perry et al 1987 and P fecundity of S curvatecurvata to be near 1780 eggs scothscotti in the southern appalachian mountains for emergentprepreemergent nymphs this is much greater folsom and manuel 1983 however lechleit- than that proposed for S americana from the ner and kondratieff 1983 detailed a 1 yr life rio conejos he used interocular width as an history for P doreatadorsata in virginia index to predict fecundity conversion of inter- multiple year life histories are common ocular width to HCW likely involves a factor among larger species of the pteronarcyidae of 2xax which would make S curvatecurvata the larger stewart and stark 1988 accompanying this of the two stonestonefliesflies this largely accounts for long nymphal growth and perhaps contribut- differences in fecundity mutch and pritchard ing to it is another life history trait long egg 1986 reported that S ameTiamericanacana as S paralle diapause univoltine growth patterns and la had a warm stenothermal egg development direct egg development are ancestral patterns most species in this family have conserved while the semivoltine growth and diapause of the life history traits that lillehammer et al P badia and P californicacalifornica are derived traits 1989 proposed as ancestral these traits lillehammer et al 1989 future studies of include univoltine slow cycles temperature egg incubation in lower latitudes of north dependent growth and direct egg develop- america will enable us to outline the range of ment isoperla quinquepunctatequinquepunctata and JL zionenzionek responses of which pteronarcys and pteronar sis have likely abandoned all of these except cella are capable temperature dependent growth unanswered questions pteronarcyidae several largely unanswered questions per- pteronarcella badia gaufin et al 1972 sist about the life histories of stonestonefliesflies in and reported that a 2 yr life cycle was possible for along the rio conejos we have found that this species in utah however S perry et al nymphs of many chloroperlids are not avail- 1987 and stanford 1975 reported a univol- able in surface sediments until just prior to tine life history in montana no eggs were emergence they must be hyporheichyporheic in their reared in either montana study and it is habitat choice second chloroperlids of the apparent from their growth histograms that present study did not readily produce eggs in early instars were missed entirely therefore captivity and those incubated never hatched semivoltine life history is most probable we can still ask many questions about their throughout its range life histories the answers would require a nymphs of this species are more likely to detailed study of the hyporheichyporheic habitat of an be found aggregated on filter paper leaf mod- open sediment stream like the rio conejos els than alone hassage et al 1988 we have this study should concentrate only on the also observed nymphs aggregating under mar- chloroperlids since they are generally abun- gin cobble immediately before emergence dant and diverse such a study would still fit adults aggregate in leaf debris at the base of within the comparative study approach of willow and cottonwood coppicescoppicuscoppices at the rio sheldon 1972 but the guild would involve conejos this behavior may be attributable to hyporheichyporheic chloroperlids the transformation and nighttime refuge sites to settle the dilemma of aberrant sex ratios being contagiously distributed hassage et al in this family studies must concentrate on the 1988 also postulated that aggregation in P presence of male nymphs in the stream in badia lowers individual risk to predation this way the search for adult males whose 199511995 STONEFLY LIFE HISTORIES 17 whereabouts are unknown need not take siphonoperlastphonoperia torrentitorrentiumum and chiotchloroperlachloroperiaChlorChloro periaperlaoperia tripunctatatripunctatetnpunctata place since both sexes of nymphs presumably plecoptera chloroperhdaechloroperlidaeChloroperlidae freshwater biology 20 11 18 a enjoy similar microhabitatmicro habitat if no male ERNST M R AND K W STEWART 1985a growth and drift nymphs are located then incubation of eggs of nine stonstoneflyefly species plecoptera in an oklahoma from virgin females should be conducted to ozark foothills stream and conformation to regression confirm the possibility of parthenogenesis models annals of the entomological society of an exciting observation we made during america 78 635 646 basking 1985b emergence patterns and an assessment of the study was that of in the sun of collecting methods for adult stonestonefliesflies plecoptera nearly all adults of summer emerging species in an ozark foothills stream canadian journal of most displayed a remarkably consistent pat- zoology 63 2962 2968 tern of ascendence of riparian vegetation be- FLANNAGAN J FE 1977 life cycles of some common aqua- ginning at about 0800 h activity usually ceased tic insects of the athabasca river alberta alberta oil sands environmental research program report by 1300 h when air temperatures were hottest 11 20 appp this ascendence culminated for females in FOLSOM T C AND K L MANUEL 1983 the life cycle of egg batching and oviposition flights while pteronarcys scotti plecoptera pteronarcyidae from males used these riparian staging grounds for the southern appalachians USA aquatic insects 5 mate searching drumming and mating 227 232 Stonestonefliesflies should be investigated for potential GAUFIN A R W E RICKER M MINER P MILAM AND R A HAYS 1972 the stonestonefliesflies plecoptera of to benefit from basking an unreported phe- montana transactions of the american entomological nomenon for plecoptera society 98 1 161 GRANT P R AND R J MACKAY 1969 ecological separa- acknowledgments tion of systematically related stream insects cana- dian journal of zoology 47 691 694 we thank the conejos peak district of the HARPER PR P 1973a emergence reproduction and growth of setipalpian plecoptera USU S forest service for providing lodging and in southern ontario oikosbikosoikos2424 94 107 laboratory space during the study special 1973b life histories of Nemournemoundaenemouridaeidae and leuctnleuctrileucon thanks go to J B moring for his help with dae in southern ontario plecoptera hydrobiologia sample collection and D ziegler for providing 4130941 309 356 some fecundity data for P badia this project HARPER P P AND H B N HYNES 1972 life histories of was partially funded by faculty research funds capniidaecapnndae and taeniopterygidae in southern ontario plecoptera archivarchev farmar hydrobiologie supplement of UNT and a national science foundation 4027440 274 314 grant BSR 8308422 to KWS HARPER P P AND E MAGNIN 1969 cycles vitauxvieaux de quelquesquelquerquesques plecopteres des laurentidesLaurentides insecta literature CITED canadian journal of zoology 47 483 494 HARPER P P M LAUZON AND F HARPER 1994 life BARTON D R 1980 observations on the life histories and cycles of sundry stonestonefliesflies plecoptera from biology of ephemeroptera and plecoptera in north- quebec review dentomoiogiedentomologie quebec 36 28 41 eastern alberta aquatic insects 2 97 111 HASSAGE R L AND K W STEWART 1990 growth and BAUMANN R W A R GAUFIN AND R F SURDICK 1977 voltinism of five stoneflystonefly species in a new mexico the stonestonefliesflies plecoptera of the rocky mountains mountain stream southwestern naturalist 35 memoirs of the american entomological society 31 130 134 BENEDETTO L A 1973 growth of stonstoneflyefly nymphs in HASSAGE R L R E DEWALT AND K W STEWART 1988 swedish lapland entomologistentomologisk tidsknfttidskrift 94 15 19 aggregation of pteronarcella badia nymphs and CATHERGATHER M R AND A R GAUFIN 1975 life history and effects of interaction with ciaclaasseniaclaassemaClaClaasasseniasema sabulosasabulosesabulosa ecology ofofmegarcysmegarcysMegarcys signata plecoptera perlodidae plecoptera oikosbikosoikos5353 37 40 mill creek wasatch mountains utah great basin HENDRICKS S P 1993 microbial ecology of the hyporhehyporbe naturalist 35 39 48 ic zone a perspective integrating hydrology and CUSHMAN R M J W ELWOOD AND S G hildebrand biology journal of the north american benthologi 1977 life history and production dynamics of allo cal society 12 70 78 petiapetlaperlaperia mediana and diplectrona modesta in walker hilsenhoff W L AND S J BILLMEYER 1973 perlo branch tennessee american midland naturalist didaedidace plecoptera of wisconsin great lakes ento- 9835498 354 364 momologistlogist 6 1 14 DOSDALL L AND D M LEHMKUHL 1979 stonefliesStoneflies holdsworth R P 1941a the life history and growth of plecoptera of saskatchewan quaestiones entomoantomo pteronarcys proteus newman pteronarcyidae logicallogicae 15 3 116 plecoptera annals of the entomological society of ELLIOTT J M 1967 the life histories and drifting of america 34 495 502 plecoptera and ephemeroptera in a dartmoor stream 1941b additional information and a correction con- journal of animal ecology 36 343 362 cerning the growth of pteronarcys proteus newman 1988 interspecific and intraspecific variation in pteronarcyidae plecoptera annals of the entomol- egg hatching for british populations of the stonestonefliesflies ogical society of america 34 714 715 18 GREAT BASIN naturalist volume 55

HURYNHURYNAA DANDJD AND J B WALLACE 1987 the exopterygote 1990 water resources data colorado water year insect community of a mountain stream in north 1989 vol 1 missouri river basin arkansas river carolina USA life histories production and func- basin and rio grande river basin water data report tional structure aquatic insects 9 229 251 CO 89 1 united states geological survey HYNES H B N 1992 some thoughts on unanswered department of the interior questions about stonestonefliesflies XI international sympos- RADFORD D S AND R HARTLAND ROWE 1971 the life lumium on plecoptera tomahawk WI cycles of some stream insects ephemeroptera ILLIES J 1952 die plecopterenplecopteran und das monardscheMonardsche plecoptera in alberta canadian entomologist 103 Prinpnnzipprinzipprinziozip berlin limnologischelimnologiscbe flubstationFlubstation freuden 609 617 thalthaithalathal33 53 69 SHELDON A L 1972 comparative ecology ofarcynopteryx KNIGHT A W AND A R GAUFIN 1966 altitudinal distribu- and dzura plecoptera in a california stream archivarcbivarchev tion of stonestonefliesflies plecoptera in a rocky mountain fur hydrobiologie 69 521 546 drainage system journal of the kansas entomological SHELDON A L AND S JEWETT 1967 Stonstoneflyefly emergence society 39 668 675 in a sierra nevada stream pan pacific entomologist kondratieff B C AND R F KIRCHNER 1991 new 43143 1 8 nearctic chloroperlidaecbloroperlidaeChloroperlidae plecoptera journal oftheodtheof the SHORT R A AND J V WARD 1980 life cycle and produc- new york entomological society 99 199 203 tion of sawalaskwala parparallelaparallelsallela frison plecoptera perlo lechleitner R A AND B C kondratieff 1983 the didaedidace in a colorado montane stream hydrobiologia life history of pteronarcys dorsatadoreata say plecoptera 6927369 273 275 pteronarcyidae in southwestern virginia canadian SNELLEN R K AND K W STEWART 1979 the life cycle journal zoology 61 1981 1985 and drumming behavior of zealeuctra claasclaasseniclaassemclaassensclaciaassemseni lillehammer A J E BRITTAIN S J SALTVEIT AND P S frison and zealeuctra hiteihidei ricker and ross NIELSEN 1989 egg development nymphal growth plecoptera leuctndaeleuctridae in texas USA aquatic and life cycle strategies in plecoptera holarctic insects 1 65 89 ecology 12 173 186 STANFORD J A 1975 ecological studies of plecoptera in MALMQVIST B AND P SJOSTROM 1989 the life cycle and upper flathead and tobacco rivers montana un- growth of isoperla grammatica and LI1 difformis published doctoral dissertation university of utah plecoptera in southernmost sweden intra and salt lake city interspecific considerations hydrobiologia 175 STANFORD J A AND A R GAUFIN 1974 hyporheicHyporheic com- 97 108 munimunitiesties of two montana rivers science 185 MUTCH R A AND G PRITCHARD 1986 developmental 700 702 rates of eggs of some canadian stonestonefliesflies plecoptera STANFORD J A AND J V WARD 1993 an ecosystem per- in relation to temperature journal of the north spectivespec tive of alluvial rivers connectivity and the american Benthbenthologicalological society 5 272 277 hyporheichyporheic corridor journal of the north american NELSON C R AND R W BAUMANN 1989 systematics Benthbenthologicalological society 12 49 60 and distribution of the winter stonstoneflyefly genus capniaacapniacapma STARK B P S W SZCZYTKO AND R W BAUMANN 1986 plecoptera capniidaecapnndae in north america great north american stoneiesstonestonefliesfliesiesles plecoptera systematics basin naturalist 49 289 366 distribution and taxonomic references great basin PERRY S A W B PERRY AND J A STANFORD 1987 naturalist 46 383 397 effects of thermal regime on size growth rates and STEWART K W AND B P STARK 1988 nymphs of north emergence of two species of stonestonefliesflies plecoptera american stonstoneflyefly genera plecoptera entomological taeniopterygidaetaemopterygidae pteronarcyidae in the flathead society of america thomas say foundation 12 river montana american midland naturalist 117 1460lugo1 460 83 93 STEWART K W R L HASSAGE S J HOLDER AND M W PERRY W B E F BENFIELD S A PERRY AND J R OSWOOD 1990 life cycles of six stonstoneflyefly species WEBSTER 1987 energetics growth and production plecoptera in subarctic and arctic alaska streams of a leaf shredding stonstoneflyefly in an appalachian annals of the entomological society of america 83 mountain stream journal of the north american 207 214 benthologicalBenth ological society 6 12 25 SZCZYTKO S W AND K W STEWART 1979 the genus PETSCH H E JR 1987 water resources data colorado isoperla plecoptera of western north america water year 1986 vol 1 missouri river basin morphologyholomorphologyholo and systematics and a new stonstoneflyefly arkansas river basin and rio grande river basin genus Cascadocascadoperlaperla memoirs of the american water data report CO 86 1 united states geological entomological society 31 survey department of the interior ULFSTRAND S 1968 life cycles of benthic insects in 1988 water resources data colorado water year lapland streams ephemeroptera plecoptera 1987 vol 1 missouri river basin arkansas river trichoptera diptera simuliidae oikosbikos 19 basin and rio grande river basin water data report 167 190 CO 87 1 united states geological survey ZAPZAR J H 1984 statisticalbiostatisticalBio analysis and2nd edition prentice department of the interior hall inc englewood cliffs NJ 718 appp 1989 water resources data colorado water year 1988 vol 1 missouri river basin arkansas river received 1 october 1993 basin and rio grande river basin water data report accepted 11 april 1994 CO 88 1 united states geological survey department oftheodtheofthe interior great basin naturalist ssi551 0 1995 appp 19 28 pollinator SHARING BY THREE SYMPATRIC milkvetches INCLUDING THE endangered SPECIES astragalus MONTH

S A geerl3 V J Tepedinotepedin024tepedino2424 T L Griswold 2 and W R bowlinlbowlind

ABSTRACT insects visiting flowers of the endangered heliotrope milkvetchmilkvetch astragalus montilmontr were compared with those visiting two common sympatric convenerscongenerscongeners A kentrophyta and A miser on three sites on the wasatch plateau of central utah for 2 yr we recorded 27 species of bees most of which were uncommon visiting the three species all three species were primarily visited by native bees of the genera osmiacosmia 15 species andor bombus 4 species most osmiacosmia species visited the three species of astragalus indiscnminantlyindiscriminantly bumblebees preferred A miser and avoided A montu our hypothesis that A monthmontu flowers would receive fewer total bee visits and be visited by fewer bee species than their common congeneisconvenersconeoncongcongenersgeneiseners was rejected A montiimontaimontn was intermediate to the two common species in its attractiveness to bees also rejected was our hypothesis that the greater similarity between A montit and A kentrophyta in flower size flower morphology and micromicrohabitathabitat would be associated with greater similarity of flower visitors than either had with A miser the data suggest that rather than competing with each other for pollinatorspollinators the three species of astragalus facilitate each other s visitation rates

key words astragalus milkmilkvetchvetch endangered plant reproduction pollination facilitation bee diversity conservation fabaceae osmiacosmia

many insects such as dipteransdipterans and lepidop sparse and likely to attract foraging bees only teranscerans use flowers only as fuel stations elton incidentally we expect such species to be pol 1966 they collect nectar and burn it as they linatorligator vulnerable and therefore to be highly search for suitable spots to lay eggs such self compatible and perhaps primarily self insects may merely pass through areas where pollinatingpollinating karron 1987 it is less clear flowers are sparse bees in contrast are central whether plants in other categories of rarity place forforagersagers orians and pearson 1979 that especially endemicsendemics rabinowitz 1981 kruck must consistently reap profits in both nectar and berg and rabinowitz 1985 are also pollinator pollen for they forage not simply to under- vulnerable endemicsEnde mics have narrow habitat write their own movements but to provide food specificity but may be locally abundant to rear their progeny as well stephen et al one such endemic the rare heliotrope 1969 because bees are under strong selective milkmilkvetchvetch astragalus montiimontai welsh is limited pressure to be profitable foragersforagers they are to a few isolated populations in limestone attracted to dense patches of flowers heinrich gravel outcrops on the wasatch plateau of 1976 1979 thomson 1982 bumblebees for central utah at about 3350 in there it grows example quickly recognize and exploit partic- with two common convenerscongenerscongeners A kentrophyta ularly rewarding flower patches heinrich var tegetariustegetarius S wats dorn hereafter A 1976 1979 other bees probably do so also kentrophyta and A miser var oblongifolius density dependent foraging behavior by redbrydb cron hereafter A miser in all three bees has important implications for certain rare species seed production requires or is in- plants rabinowitz 1981 distinguished seven creased by pollinator visits to flowers geer and types of rarity in plants using the following tepedino 1993 information on the identity three criteria 1 local abundance 2 habitat and biology of these pollinatorspollinators is important specificity narrowdarrownarrow or wide and 3 geographic for A montiimontai occurs on rangelands that are range large or small those species with both grazed by domestic livestock and sprayed with narrow habitat specificity and small local pop- insecticides to control grasshoppers successful ulationsulations regardless of geographic range are management of this rare species requires

departmentldepartment of biology utah state university logan UT 84322530584322 5305 2usdaSDA ARS bee biology and systematics laboratory utah state university logan UT 84322531084322 5310 31presentresent address wallowagallowa whitman national forest highway 82 box 88401 enterprise OR 97828 authurauthor to whom correspondence should be addressed

19 20 GREAT BASIN naturalist volume 55 knowledge of how such spraying may affect its than do those of its convenerscongenerscongeners the three pollinapollinatorstors species overlap in bloom time for about 3 wk in this report we compared composition fig 1 and abundance of pollinator fauna of A montiimontai heliotrope milkvetchmilkvetch is a acaulescentsubacaulescentsub with those of its two sympatric convenerscongenerscongeners plant 1 5 cm tall that arises from a branched because there may be wide variation in a caudex flowers are deep purple with white species pollinatorspollinators between years and sites wingtipswingtips there may be a dozen to a hundred tepedino and stanton 1981 herrera 1990 or more flowers ts78787.8 15151.5 mm long N 10 eckhart 1992 we censcensusedcensusesused pollinatorspollinators of A geer unpublished per plant two to eight per montiimontai and its convenerscongenerscongeners for 2 yr at three sites raceme barneby 1989 it does not appear to we hypothesized that A montiimontai would 1 reproduce vegetatively personal observation attract fewer individual pollinatorspollinators 2 have in 1989 and 1990 A montiimontai commenced flow- lower pollinator species diversity than its two ering with final snowbeltsnowmeltsnowmelt beginning as early common convenerscongenerscongeners and 3 share more species as june and continuing for about 4 wk until of flower visitors with A kentrophyta than mid july fig 1 with A miser because similarity in plant and the common species A kentrophyta started flower size flowering time and micromicrohabitathabitat to flower approximately I1 wk before A montiimontai is greater with the former than with the latter and continued to flower through early august it is prostrate with stems that fork repeatedly SPECIES AND STUDY AREAS and closely to form low convex cushions cov- ered with small blue white to purplish flowers all three species of astragalus are small 66gg666.6 121.2 mm long N 10 geer unpublished perennial herbaceous legumes A montiimontai is re- only two per raceme barneby 1989 strictstricteded to three mountainmountaintopstops on the wasatch the other common congener A miser com- plateau in central utah although isely 1983 menced flowering 1 2 wk after A montiimontai and proposed that A montiimontai be reduced in status continued flowering until september it is taller to a variety of A limnocharis barneby it was 2 20 cm than A montiimontai or A kentrophyta listed as endangered under the endangered flowers are larger 11411411.4 141.4 mm long N2vav species act in 1987 as A montiimontai and remains 11 geer unpublished and vary in number per so anonymous 1991 therefore we refer to raceme 3 15 barneby 1989 and in color this taxon as A montiimontai flowers may be white pink or lavender A kentrophyta and A miser are widespread all astragalus species have papilionaceous species that occur with A montiimontai at three sites blossoms composed of a showy standard or on two of the mountains the third mountain is banner petal a keel that protects the joined less accessible and was not included in the stamens and pistil and two wings that along study A kentrophyta is widespread and abun- with the keel typically serve as a landing plat- dant in the rocky mountains mostly between form kalin arroyo 1981 to trip A miser 2280 and 3650 m A miser one of the most flowers bees land on the keel and force their common species of astragalus in the rocky way under the banner personal observation as mountains is locally abundant from sagebrush they do for other species of astragalus green foothills to the spruce fir belt barneby 1989 and bohart 1975 faegrifaegre and van der fiji 1979 the three species co occur at 3250 to 3350 m visitors to A montiimontai or A kentrophyta spread in an engelmann spruce picea engelmanniiengelengelmanniamannii the wing petals with their midlegsmidlege and take parrysubalpineParry subalpine fir abies lasiocarpalasiocampa hook nectar or comb pollen from the anthers to nutt community A montiimontai and A kentrophyta their abdominal pollen baskets with their are intermingled in limestone gravel outcrop foreforelegslegs personal observation stylar hairs pings where A miser is found only occasional- termed a brush mechanism aid in the collec- ly A miser is most abundant nearby where tion of pollen by transporting it from the keel soil is deeper and less rocky A montiimontai and A outward kalin arroyo 1981 miser occur at similar local densities on sexual reproduction by A miser and A heliotrope mountain 93939.3 01m2 and 12612.6196 kentrophyta requires insects to transfer pollen 83m2 geer unpublished data there are A montiimontai is capable of unassisted self pollina fewer A kentrophyta 96gg262.6 08m2 geer un- tion autogamy however fruits produced published but individuals cover more ground autogamousautogamouslyly by A montiimontai may be inferior in 199519951 pollinatorsPOLLINATORS OF SYMPATRIC milkvetchesmllkvetches 21

wind and no precipitation initially sight identification of some taxa was attempted so as A miser to reduce impact on the pollinator community it soon became obvious that it was impossible to identify osmiacosmia and other individuals with-

A month out laboratory examination subsequently all flower visitors were collected whenever possi- ble few insects other than bees visited the

A kentrophyta flowers diversity of bee visitors to each astragalus

1 10 20 30 10 20 30 10 20 species was calculated using simpsinssimpsonsSimp sons diver- june july august 1 pa sity index D I Is I1 p2 where Ppi the proportion of individualssi that belong to each fig 1 blooming dates for three co occurring species of astragalus at the SSH site solid line 1989 dashed line bee species southwood 1978 simpson s 1990 index gives little weight to rare species and more weight to common ones similarity of the bee fauna visiting astragalus species was quality to those produced by geitonogamous estimated using Czekczekanowskfsanowski s similarity or xenogamous hand pollinationspollinations or open pol czekanowskichekanowski index cg ajab n where N is the linatedlanated control treatments there are fewer Cs njabNJ ab number of plant species being compared is seeds per fruit and seeds are smaller geer J the number of bee species shared by those and tepedino 1993 thus all three species plant species and a b etc are the total num- probably benefit from insect visitation ber of bee species visiting each plant species cg METHODS southwood 1978 Cs is based on species presence alone we also calculated cicp which insect visitors were collected for about 3 wk adjusts for the number of individuals per in 1989 and for 2 wk in 1990 at the following species southwood 1978 the indices range three sites starting when A montiimontai was in peak from 0 no similarity to 101.0io complete similari- bloom the head of mill stream on ferron ty they were calculated between pairs of mountain HMS south side of heliotrope species and among all three species mountain SSH and east end of heliotrope probable pollinatorspollinators of the three astragalus mountain EEH in 1990 collections from all species were ascertained by examining flower three astragalus species were made only at visitors and recording areas of their bodies on the SSH site because only two insect collec- which pollen was found specimens were then tors were available instead of four as in 1989 relaxed and pollen was removed using an we concentrated on the SSH site in 1990 to insect pin or by dabbing it with acid fuchsin make the number of collector hours there gel beattie 1971 the pollen was placed on a equivalent to the 1989 effort in 1990 visitors glass slide with acid fuchsin gel warmed until to A kentrophyta were collected at the SSH liquid and a cover slip applied modified from and HMS sites and visitors to A miser were faegrifaegre and iverson 1964 one slide per leg or collected at the SSH and EEH sites following two slides per abdomen were made for each are approximate direct distances between sites insect all slides were viewed at loox magni- HMS to SSH 363.6 kinkm HMS to EEH 242.4 ficationfi and the pollen compared to a pollen kinkm and EEH to SSH 121.2 kinkm reference collection of species in bloom at the Pollinapollinatorstors were collected with a standard study sites butterfly net and killed in cyanide jars cold temperatures strong winds and frequent pre- RESULTS cipitation snow and rain prohibited pollina tors from flying during all but brief windows bees were scarce at the study sites in both of calm sunny weather so opportunistic collec- years table 1 appendices I1 II11 bee visitors tion was necessary to ensure an adequate sam- per plant species ranged from about 050.5os to just ple size collections were made from all three over 3 per hour a small number considering species contemporaneously whenever weath- that many flowers of each species were being er permitted ie temperatures 13c little monitored bee numbers were higher in 1990 22 GREAT BASIN naturalist volume 55

TABLE 1 number of person hours spent collecting and number of bee individuals collected or observed visiting flow- ers of astragalus montu asmo A kentrophyta aske and A miser admiasmi at three sites on the wasatch plateau in 1989 and 1990 SSH EEH south and east side heliotrope mountain respectively HMS head of mill stream ferron mountain SSH EEH HMS asmo aske asmiadmi asmo aske asmiadmi asmo aske asmiadmi 1989 hours 24 8 10 30 24 22 8 16 16 individuals 28 9 10 30 19 10 5 11 18 individualshourIndividual shour 12 11 10 10 08 05 06 07 11 species 7 3 5 7 7 3 3 4 7

1990 hours 30 15 15 12 12 12 12 individuals 57 7 35 40 24 16 24 individualshourIndividual shour 19 05 23 33 20 13 20 species 10 5 11 5 5 6 3 when categorized by site and astragalus species visiting A montiimontai than the other two species visited six of seven categories had more indi- but this difference is probably because we col- viduals per hour in 1990 than in 1989 lected at three sites for A montil but at only the initial hypothesis that A montiimontai would two for each of the other two species have fewer individual flower visitors than calculations using species diversity D would its common convenerscongenerscongeners received little also failed to yield expected trends table 2 support table 1 appendices I1 II11 in 1989 in 1989 diversity of visitors to flowers of A there was little difference among species in montiimontai was very similar to diversity recorded for visitors per person hour at SSH at EEH A A kentrophyta and A miser comparisons for montiimontai flowers were visited more often than 1990 are more tenuous because of the differ- the other species conversely at HMS A ences among species in number of sites sam- montil flowers received the fewest visits in pled however diversity of flower visitors was 1990 comparisons of number of visitors among highest for A miser and similar for A montiimontai all three astragalus species could be made and A kentrophyta diversity in 1990 was only at the SSH site where A montil had an generally lower than in 1989 although num- intermediate number of visitors per hour at ber of individuals captured was greater EEH A montil again had more visits per hour the most frequent visitors to these astra- than A kentrophyta and at HMSH M S it had fewer galus species in both 1989 and 1990 were visits per hour than A miser esmia bees table 3 for the small flowered the prediction that species richness and A montiimontai and A kentrophyta in both years species diversity of bees visiting the three 70070070.0 of all visitors were osmiacosmia bees only astragalus species would be lowest for A montiimontai for A miser in 1990 did the percent osmiacosmia was also provisionally rejected the number of visitors drop below 50 A miser was more species captured on A montiimontai commonly ex- frequently visited by bumblebees especially ceeded those captured on the other species at SSH the abundance of bumblebees caused both when more hours were spent collecting SSH to have the lowest percentage of osmiacosmia from A montil than the other species 1989 individuals recorded at any site in both years SSH and when collecting hours were equal even so osmiacosmia bees were always more than 1990 HMS table 1 only once when fewer 60 of the total flower visitor fauna recorded hours were spent collecting on A montiimontai than in any site year on the other astragalus species 1989 HMS because of greater similarities in flower size was A montiimontai visited by the fewest species of color and microclimate we expected A montiimontai bees when all sites were considered total and A kentrophyta to have more visitors in number of species collected on A montiimontai in common than either did with A miser this 1989 exceeded those captured on A kentro was not true in either year the three pairings phytapayta and equaled those captured on A miser of astragalus did not differ much in the num- table 2 in 1990 more species were caught ber of bee species they shared though results 199511995 pollinatorsPOLLINATORS OF SYMPATRIC milkvetches 23

TABLE 2 number of individuals number of species and species diversity D of bees found visiting three species of astragalus at three sites on the wasatch plateau in 1989 collections were made for each species at all three sites in 1990 collections were made at all sites for A montiimontai but at only two sites for the other two species for comparative pur- poses collection data for the latter two species are shown in 1989 for all three sites and for only the two sites collected at in 1990 D simpsonSimpsorssois s diversity index individuals species DI astragalus species 3 sites 2 sites 3 sites 2 sites 3 sites 2 sites 1989 montiimontai 63 13 0870.87ost kentrophyta 39 28 9 8 079 0810.81 miser 38 28 13 11 0880.88 0870.87

1990 montiimontai 113 13 0620.62 kentrophyta 31 7 0600.60ogo miser 59 12 0790.79

linildin 1989 only indiviualsmdiviiialsindividuals that were collected were used inin calculations because uncaptured osmiaoemia0smia individuals were not identifiable to species

TABLE 3 percent visitors that were osmiacosmia bees to the flowers of three astragalus species abbreviations as in table 1 data shown grouped by species across sites and by site across species for 2 yr for comparative purposes 1989 data are shown in entirety 3 sites or 3 species or only for the 2 sites or 2 species sampled in 1990 asmo aske asmiadmi SSH EEH HMS across sites across species

1989 3 sitesspeciessites species 88988.9 71871.8 737 623 88188.1 76576.5 2 sitesspeciessites species 78678.6 64364.3 85785.7 87087.087 0

1990 938 74274.2 475 62662.6 87587.5 95095.0

varied somewhat with year and with index used movements only two were interspecific in table 4 in 1989 the three pairings ofastragaof astraga- 1990 4 of 21 observed interplant movements lus species had about the same number of bee were between species interspecific visits species in common in 1990 A miser and A occurred most commonly where species grew montiimontai had about twice the number of species intermingled in common as did the other pairings neither coefficient of similarity Cscg or cicp consistently discussion supported the hypothesis in 1989 but not two hypotheses make predictions about the 1990 cg and C were highest for the A mon Cs ci abundance and diversity of visitors to the flow- tii A kentrophyta comparison ers of rare plants entomophilous plants many bees visiting astragalus flowers car- for levin and anderson 1970 straw 1972 and pollen 43 ried on their bodies of the bees karron 1987 proposed that pollinatorspollinators should captured primarily females of the genus osmia0smiaoemia be more flower constant to abundant plant pollen had been collecting pollen loads com- species than to rare ones that this differential prised primarily astragalus pollen all means flower constancy would result in more suc- 80 table 5 it is unknown whether loads cessful reproduction by majority species commonly contained more than one species of than by minority species and that over time astragalus because pollen grains could not be minority species would become extinct because distinguished to species with the light micro- of dwindling recruitment or would evolve scope some method of self reproduction levin 1972 our observations of foraging bees suggest A corollary of this hypothesis is that both the some interspecific movement in 1989 few number and diversity of visitors to the flowers osmiacosmia individuals flew between A montiimontai and of rare plants should be lower than they are to A miser or A kentrophyta of 74 interplant abundant ones 24 GREAT BASIN naturalist volume 55

TABLE 4 number of bee species S collected on each astragalus species and number of species shared C and simi- larity indices for each pairing for each year CsC czekanowskichekanowskiCzekanowski s similarity index for bee species presence absence ciC index weighted by individuals captured

1989 1990 astragalus species pairpan S C Cscg ciC S C CS ciC montu 13 6 050 034 13 7 056 037 miser 13 12 montu 13 6 055 043 13 4 040 035 kentrophyta 9 7 kentrophyta 13 5 045 043 12 3 032 053 miser 9 7 all three species 20 4 035 027 21 3 028 030

in contrast the facilitation hypothesis re- trips gross 1992 also reported that bees for- viewed by rathcke 1983 predicts that rare aging on closely related legumes commonly species growing with attractive more abun- moved between species thus there was no dant species may actually reproduce more detectable rare species disadvantage and no successfully because the latter draw many evidence that endemicsendemics at least those growing more pollinatingpollinating insects into the area than in close proximity to abundant convenerscongenerscongeners are would otherwise be present if so rare and pollinator vulnerable abundant sympatric species should have simi- the shared micromicrohabitathabitat and similarities in lar visitor diversity and visitor abundances flower size and morphology of A montiimontai and should reflect respective frequencies of the A kentrophyta led us to expect that facilitation plants this study indirectly assessed the would be more likely between these two species importance of facilitation and competition A and therefore that they would have more visi- direct assessment is difficult because 1 the tors in common than either would with A miser experiments necessary to distinguish between for example thomson 1978 1981 1982 found alternatives cannot be conducted when the that in two species mixtures the degree of 11plant protagonist is protected by the endan- intermingling and the similarity in structure gered species act and 2 A montiimontai did not and appearance of convenerscongenerscong eners flowers deter- occur in the absence of its convenerscongenerscongeners on our mined the importance of competition and study sites so visitation rates of facilitated mutualism the more similar the flowers the and unfacilitated populations could not be more likely that visitation rates to rare species compared would be bolstered by the presence of abun- our results supply consistent though indi- dant species and the more likely that visitors rect support for the facilitation hypothesis would be shared our data supported this except for bumblebees which foraged almost expectation for 1989 but not for 1990 table 4 exclusively from large flowered A miser bees in 1990 Cscg for the A montiimontai A kentrophyta did not discriminate against A montiimontai but comparison was intermediate to the other rather seemed to treat all three astragalus comparisons for ciC it was lower than the other species as one taxa first A montiimontai did not comparisons thus results for the similarity consistently attract fewer visitors per hour analyses also tend to support the hypothesis than did the other species indeed visitation that most bees do not distinguish among these rates to A montiimontai were higher than to the astragalus species when foraging and that the other species in three of six site years table 1 astragalus species tend to facilitate each second neither species richness nor species other s visitation rates diversity of pollinatorspollinators was consistently lower only bumblebees seem uninfluenced by for A montiimontai than for the other species table 2 astragalus flowers in the aggregate they in fact an equal or greater number of species clearly preferred flowers of A miser and visited A montiimontai than visited the others in avoided those of the other astragalus species both years and finally bees were observed flowers of A miser are large probably more moving between species on individual foraging rewarding and provide a landing platform from 199511995 pollinatorsPOLLINATORS OF SYMPATRIC milkvetchesmllkvetches 25

TABLE 5 percent astragalus pollen grains in pollen loads and location of pollen loads carried by bees collected on three astragalus species at three sites on the wasatch plateau in 1989 and 1990 location of mean pollen astragalus number of astragalus species pollen loads pollen SSEE abdomen legs montiimontai 45 82 4 42 3 miser 19 90 1 19 kentrophyta 5 95 1 5 which large energy demanding bumblebees plants land managers must eliminate losses can readily forage other large flowered astra- of bees to insecticide applications made for galus species also attract numerous large bees rangeland grasshoppers and minimize physical such as bumblebees bombus sppapp and antho damage to nest sites the present insecticide phorids green and bohart 1975 sugden 1985 free buffer zone currently 484.8 km around rare karron 1987 in comparison bumblebees plant populations should continue to be main- seemed unable to land on the small weakly tained areas where bees nest in soil should supported A montiimontai flowers which are borne also be protected from livestock trampling above the foliage they did occasionally exploit off road vehicle use and foot traffic sugden the tiny A kentrophyta blossoms while perched 1985 such diversity comparable to or greater on the foliage of that cushion plant than that of other subalpine areas in north factors other than flower abundance can america moldenke and lincoln 1979 is to influence the flight path of foraging bees be marveled at and preserved foragers because bees are central place foragers orians acknowledgments and pearson 1979 travel time and energy expended between flower patches and nest we are grateful to the many people who are also important thus bees may patronize a assisted in this study etta Securesechrestst and mike flower patch because of its proximity to their cram were reliable field and laboratory assis- nest even though flowers are more abundant tants john healey don riddle and bob elsewhere for example osmiacosmia bees mated thompson of the US forest service and larry and nested at the sheltered EEH site where england US fish and wildlife service helped relatively few A kentrophyta or A miser plants in a variety of ways from locating plant popu- grew the population of A montiimontai was small lations to putting a roof over our heads the but dense nevertheless bees visited flowers manuscript was constructively reviewed by M at least as frequently at EEH as at the other barkworth K harper and E sugden this more flower rich sites tables 1 2 thus suit- study was funded as part of the APHIS grass- ability of nesting habitat at EEH rather than hopper IPM project it is journal paper 4436 astragalus flower abundance may best account from the utah agricultural experiment station for the abundance of bees there the effect of wild bee nesting sites on seed production of literature CITED surrounding vegetation is poorly studied and ANONYMOUS 1991 endangered and threatened wildlife warrants additional attention and plants 50 CFR 171117 11 and 171217.1217 12 july 15 1991 rigorous subalpine communities of the publication unit USU S fish and wildlife service wasatch plateau with frequent high winds washington DC thunderstorms and below freezing tempera- BARNEBY R C 1989 fabalesbabales pages 12 167 in A A H Hohnholmgrengren N H holmgrenHohn gren J L a cronquist tures during the blooming season support reveal and P K holmgrenHohngren eds intermountain flora surprisingly rich bee fauna in 2 yr we collected 3 part B new york botanic garden bronx NY 27 bee species foraging on astragalus flow- BEATTIE A J 1971 A technique for the study of insect pan ers during 2 3 wk appendices I1 II11 these borne pollen pacific entomologist 47 82 ECKHART V 1992 spatiospatiotemporaltemporal in abun- bees are invaluable pollinapollinatorstors of native plants A variation in dance and variation in foraging behavior of the polli both rare and common their welfare must also natorsgators of gynodioecious phaceliaphacehaphacella linearis hydro- be considered in management plans for rare phallaceaephyllaceae oikosbikos 64 573 586 26 GRFATGREAT BASIN naturalist volume 55

ELTON C S 1966 the pattern of animal communities LEVIN D A AND W W ANDERSON 1970 competition methuen london 432 appp for pollinatorspollina tors between simultaneously flowering FAEGRI K AND J IVERSON 1964 textbook of pollen species american naturalist 104 45545567467 analysis hafner co NY 237 appp MOLDENKE A R AND P LINCOLN 1979 pollination ecol- FAEGRI K AND L VAN DER PIJL 1979 the principles of ogy in montane colorado a community analysis pollination ecology ard3rd revised edition pergamon Phytolphytologiaphytologicphytologia42ogiaogla 42 349 379 press oxford 244 appp ORIANS G H AND N E PEARSON 1979 on the theory of GEER S M AND V J TEPEDINO 1993 breeding systems central place foraging pages 155 177 in D J horn of the rare heliotrope milkvetchmilkvetch astragalus montu G R stairs and R D mitchell eds analysis of welsh fabaceae and two common convenerscongenerscong eners pages ecological systems ohio state university press 334 344 in R sivinski and K lightfoot eds proceed- columbus ings of the southwestern rare and endangered plant rabinowitz D 1981 seven forms ofrarityoflarityof rarity pages 205 218 conference new mexico forestry and resources in H synge ed the biological aspects ofrareof rare plant conservation division santa fe conservation wiley new york NY GREEN T W AND G E BOHART 1975 the pollination RATHCKE B 1983 competition and facilitation among ecology of astragalus cibariuscibanuscibacibariousrius and astragalus uta plants for pollination pages 305 329 in L A real bensishensis leguminosae american journal of botany ed pollination biology academic press new york 6237962 379 386 NY GROSS C L 1992 floral traits and pollinator constancy SOUTHWOOD T R E 1978 ecological methods and2nd edi- foraging by native bees among three sympatric tion chapman and hall london 524 appp legumes australian journal of ecology 17 67 74 STEPHEN W P G E BOHART AND P F TORCHIO 1969 the HEINRICH B 1976 foraging specializations of individual biology and external morphology of bees agricultural bumblebees ecological monographs 46 105 128 experiment station oregon state university 1979 majoring and minoring by foraging corvallis 140 appp bumblebees bombus dagans an experimental analy- STRAW R M 1972 A markov model for pollinator con- sis ecology 60 245 255 stancy and competition american naturalist 106 HERRERA C 1990 daily patterns of pollinator activity 597 620 differential pollinapollinatingting effectiveness and floral re- SUGDEN E A 1985 Pollinapollinatorstors of astragalus monoensis source availability in a summer flowering mediter- barneby fabaceae new host records potential ranean shrub oikos58oikosbikos 58 277 288 impact of sheep grazing great basin naturalist 45 ISELY D 1983 new combinations and two new varieties 299 312 in astragalus orophacaOrophaca and oxytropis legumino- TEPEDINO V J AND N L STANTON 1981 diversity and sae systematic botany 8 422 competition in bee plant communities on short grass KALIN ARROYO M T 1981 breeding systems and pollina- prairie oikosbikos 36 35 44 tion biology in leguminosae pages 723 769 in R M THOMSON J D 1978 effect of stand composition of insect polhill and P H raven eds advances in legume visitation on two species mixtures of hieracium systematics part 2 royal botanic gardens kew american midland naturalist 100 431 440 UK 1981 field measures of constancy in bumble- KARRON J D 1987 the pollination ecology of co occur bees american midland naturalist 105 377 380 ring geographically restricted and widespread species 1982 patterns of visitation by animal pollinatorspollinators of astragalusofastragalus fabaceae biological conservation 39 bikosoikos 39 241 250 179 193 kruckeberg A R AND D rabinowitz 1985 biological aspects of endemism in higher plants annual review received 29 april 1993 of ecology and systematics 16 447447479479 accepted 2 june 1994 LEVIN D A 1972 competition for pollinator service a stimulus for the evolution of autogamy evolution 2666826.66826 668ggs 674

APPENDIX I1 species of bees collected and observed visiting flowers ofaofA montiimontaimontn asmo A miser admiasmi or A kentro phytapayta aske at three sites in 1989 entries represent number of malesfemalesmales females collected observations are in parenthe- ses site abbreviations as in table 1 SSH EEH HMS 1 21june21 june 14 25 june 14 22 june bee species asmo aske asmiadmi asmo aske asmiadmi asmo aske asmiadmi andrenidae andrena transnigratransmgratransnigra vier 01 andrena sppapp 1 199511995 pollinatorsPOLLINATORS OF SYMPATRIC mllkvetchesmilkvetches 27

APPENDIX I1 continued SSH EEH HMS 1 2121junjune 1414 25 junjunee 1414 22 june bee species asmo aske asmiadmiasnn asmo aske asmiadmi asmo aske asmiadmi

APIDAE bombus bifarius cr 01 bombusflavifronsbombus flavifronsflavifrons cr 02 bombus huntnhuntiihunanhuntie greene 02 02 bombus nevadensis cr 01

halictidaehaliotidae evylaeus niger viereck 01 01 megachilidae anthidium tenuifloraetenwflorae callckll 210 1 illiiiiitilioill11211121 122 megachile sppapp 10 10 osmiaosimacosmia cyanopoda callckll 10 osmiacosmia hurdiihurdnhurdia white 01 osmiacosmiaosmw lingulalongula cr 01 osmiacosmiaosmianigrifronsnignnignfronsfrons cr 01 04 02 03 osmiacosmia aff nigrimgnfronsnigrifronsfronsarons 03 01 01 osmiaosimacosmia paradiparadisicaparadisicalsica sanh 10 22 osmiacosmia penstemonispenstemoms callckll 01 osmiacosmia mikeipikei ckllcall 01 osmiacosmia ausillapusilla cr 01 10 osmiacosmiaosmw sladenislademgladem sanh 20 10 40 30 20 osmiacosmia sladenislademgladem bor&ornor alpestrisalpestris 02 03 02 01 02 05 osmiacosmia tannetttannert sanh 13 12 01 01 osmiacosmiaosmiasppsppapp 59 14 03 89 11 13 11 01 01

APPENDIX II11 species of bees collected and observed visiting flowers of A montu asmo at three sites and A miser admiasmi and A kentrophyta aske at two sites each in 1990 entries represent number of malesfemalesmales females collected observations are in parentheses site abbreviations as in table 1 SSH EEH HMS 19 june 4 july 19 29 june 21 29 june visitor asmo aske asmiadmi asmo aske asmo asmiadmi andrenidae andrena mgnhirtanigrihirta asamashm 01 andrena transnigratransmgratransnigra vier 01

APIDAE apis mellifera L 015 bombus bifarius cr 01 bombusflavifronsbombusbombas flavifronsflavifrons cr 01 01 bombus huntzi greene 036 bombus nevadensis cr 035 megachilidae anthidium tenuifloraetenwflorae callckll 102 20 013 hoplitisfulgidahoplitichoplitisHoplitis fulgida cr 30 Megmegacilemegachileacile melanophaea smith 20 10 10 megachile penhpenhirtaperihirtaperihirta CO 10 10 28 GREAT BASIN naturalist volume 55

APPENDIX II11 continued SSH EEH HMS 19 june 4 july 19 29 june 21 29 june visitor asmo aske asmiadmi asmo aske asmo asmiadmi megachilidae continued osmiacosmia lingulalongula cr 20 osmiacosmia montana cr 10 osmiacosmia Aaff nignnigrinigrifronsnignfronsfronsbrons 01 01 osmiacosmia paradmcaparadparadisicaparadiparadisicalsicamca sanh 10 02 30 12 10 osmiacosmia penstemonis callckll 01 osmiacosmiaosrma ausillapusilla cr 01 osmiacosmia scullscullensculleniscullemeni callckll 20 10 osmiacosmia sladeni sanh 1913 40 10 816 78 36 121 osmiacosmia subaustralis callckll 40 10 osmiacosmia tannert sanh 92 10 10 91 10 01 great basin naturalist ssi551 0 1995 appp 29 36

FACTORS AFFECTING SELECTION OF WINTER FOOD AND ROOSTING RESOURCES BY porcupines IN UTAH

dave stricklanl2stncklan12 berranjerran T Flindersflindersl3flinders1313 and rex G catellcateslcatesi

ABSTRACT ecological and phytochemical factors potentially affecting winter dietary discrimination by porcupines erethizon dorsatumdorsatum inm the mountain brush zone of utah were studied porcupines utilized gambel oak quercus gambeliigambelitgambelligambeliieliteill as their primary winter food and roosting resource big tooth maple acer grandidentatum was the most common tree species in the study area but was rarely utilized by porcupines conifer species were used as a food and roosting resource significantly less often than they occurred in the study area despite thermal advantages provided by their rela- tively dense canopiescanopies oak feed trees were successfully separated from conifer feed trees by discriminant analysis 100 of the time oak trees were correctly classified as feed and nonfeed trees 71 of the time gambel oak contained higher amounts of crude protein fiber and tanninsfanninstannins but was lower in ether extract fractions and fatty acid content than conifers A layer of adipose tissue used as an energy reserve by porcupines may have relaxed energy intake demands sufficiently to permit them to concentrate on a diet of oak tissue which is high in protein rather than a high fat conifer diet A diet relatively high in protein may have facilitated digestion of food material high in fiber temperature did not affect selec- tion of tree species used for roosting rock and snow caves were utilized infrequently and the study population ranged widely three of 15 study animals were eaten by predators

key words porcupine erethizon dorsatumdorsatum gambel oak quercus gambellgambehigambelngambehielneinehl dietary selection mountain brush zone predation

porcupines erethizon dorsatumdorsatum roost and STUDY AREA feed in canopiescanopies of deciduous trees and shrubs for extended periods during winter in much of the study was conducted in the mountain western north america oveson 1983 craig brush zone near the mouth of spanish fork and keller 1986 sweitzer and berger 1992 canyon in north central utah elevations at apparent localized interspecific and intra- the study site range from 1650 to 2075 m the specific preferences for food and shelter general exposure is northern and terrain is resources by porcupines imply that chemical steep overstory woody vegetation is dominated andor physical advantages are available to by gambel oak quercus gambeliigambelligambelii and big them further since snow caves rock dens tooth maple acer grandidentatum aspen pop- and cover in canopiescanopies of coniferous tree ulus tremultremuloidesoides chokecherry prunus virgini species likely offer increased thermal advan- anaand douglas fir pseudostugapsendostuga menziemenziesiimenziesiasii white tages in the form of energy savings to porcu- fir abies concolor and mountain maple acer pines clarke and brander 1973 roze 1987 flabrumglabrum are also represented in the woody 1989 their dependence on a deciduous food flora the climate in spanish fork canyon and roosting resource which does not offer during the winter of 1984 85 was not atypical those advantages further strengthens the data fromhrom the spanish fork US climatological implication that chemical andor physical station located approximately 555.5 km from selective advantages are realized by dietary the study site indicate that temperatures were selection predator avoidance may also be an slightly colder and precipitation was slightly important force in food and roost tree selec- higher than average US climatological data tion the objective of this research was to for utah 1984 85 coyote caniscams latranslalatranotrans and investigate physical phytochemical and eco- mountain lion felis concolor tracks were fre- logical agents involved in selection of gambel quently encountered in the study area private oak by porcupines in south central utah access into the study area allowed observation

department of botany and range science brigham young university provo UT 84602 present address USDA forest service pleasant grove UT 84062 address correspondence and reprint requests to this author

29 30 GREAT BASIN naturalist volume 55 of a porcupine population relatively free from low temperature readings were taken daily at human disturbance an elevation of 1597 in as well as from the spanish fork climatological station METHODS laboratory and statistical methods fieldwork tissues from feed and nonfeed trees were we conducted fieldwork from late decem- analyzed for protein and phosphorus using the method 12 for ber 1984 through april 1985 at which time auto analyzer semiautomated the study population had shifted from a diet of feeds horwitz 1980 calcium magnesium inner bark phloem and cambium of woody potassium and sodium content were deter- vegetation to herbaceous vegetation the study mined by the atomic absorption method 2 area was systematically searched by researchers for plants horwitz 1980 sulphur content on snowshoes study animals were captured was determined by a wet ash process using by hand usually while they were still in tree nitric and perchloric acid crude fiber was canocanopiespies this was accomplished by grasping determined by the acid detergent fiber and distal guard hairs at the posterior end of the lignin 21 method horwitz 1980 an evalua- tail between thumb and forefinger and pulling tion of crude fat was made using the direct the tail taut the captured animal was then method horwitz 1980 on a lab con solletsoxlet secured by grasping the tail with the free hand extractor A limited number of tissue samples packard using a backward stroking motion to flatten were analyzed on a hewlett model the quills fifteen porcupines 10 females and 5 5995 gas chromatographchromatographmassmass spectrometer males were instrumented with radio transmit- GCMS for fatty acids and terterpenesterpeneypenes tannin ter collars telonics inc animals were located content was measured by the radial diffusion daily by triangulation and visual sightings were method hagerman 1987 with quebracho tan- made on each animal approximately weekly ninnln being the standard and by astringency percent occurrence of woody species was Gambgamalielgamblielliel et al 1985 soluble carbohydrates calculated from point quarter measurements were determined according to dasilveira using the feedroostfeed roost tree as the center point 1978 urine samples of captive porcupines cottam and curtis 1956 percent occurrence on a strict diet of gambel oak were analyzed of woody species vs percent utilization of each for calcium and phosphorus content when lab- feed tree species was compared using chi oratory results indicated the cap ratio in the square analysis to test whether feed tree selec- tissue of food materials was greater than ex- tion was random diameter at breast height pected eight oak tissue samples were chosen dahdbh species and distance from the feed tree at random and retested for calcium and phos- center point were recorded for the nearest phorus content according to horwitz 1980 woody stem in each quadrant point quarter on a beckman DU 30 spectrophotometer measurements were repeated using the near- differences between oak white fir and est neighbor nonfeed tree of the same species douglas fir feed and nonfeed trees were statis- as the center point tissues from feed and tically analyzed to help discern foraging pat- nonfeed trees were collected to investigate terns used by instrumented porcupines possible differences in chemical makeup chemical and ecological factors were evaluat- tissue samples from feed trees were collected ed for between species differences using two where fresh bark removal indicated the roost sample t tests and for within species differ- ing animal had foraged samples from nearest ences with paired t tests minitabMini tab 1982 neighbor nonfeed trees were taken from statistical results are reported at the p 05os05.05 branches at the same height and with a diame- and p 1.1 levels chi square analysis was ter similar to those from corresponding feed used to determine if utilization of feed tree trees bark samples were frozen and analyzed species by porcupines differed from the ex- for dietary components results from those pected discriminant analysis using backward analyses reasonably approximated values elimination and forward selection SAS 1985 reported for gambel oak smith 1957 kufeld was used to determine chemical and ecologi- et al 1981 welch 1989 location slope cal factors that best discriminate between tree aspect snow depth and climatic conditions species and between feed and nonfeed trees were recorded at each feed tree site high and of the same species 199519951 WINTERING porcupines IN GAMBEL OAK 31

TABLE 1 mean values for factors tested for possible effects on porcupine herbivoreherbherbivoryivory

oak 1 white fir 2 dougiadougladouglassfiresfir3firhirbir 3 nonfeed feed nonfeed feed nonfeed feed tree tree tree tree tree tree n 46 n 46 n 3 n3nan 3 n7nan 7 n7nan 7 distance from conifer m 207 0 0 distance to feed tree same sp m 3582 5431 377 wind speed mph 5535.53 373.7 9719.71 slope 3353 365 4241 elevation m 17792 19371193713 16802 dahdbh cm 13213213.2 1652316.523 254 404140.41 33333.3 345134.51 crude fiber 43343.3 4423 43643.6 4803 424 401124011.2 protein 49 5023502.3 40 4214.21 40 3913.91 phosphorus 00380.038 00390.039 00870.087 0064 0038 0042 ether extract fractions 909.0go 91239.123 15715.7 12713 16516.5 1891218 91291.2 water 41041.0 396233962.3 461 4971 534 5071 potassium 0390.39 03930.393 0360.36 0310.31 0160.16olg 01910.191 calcium 27 2732.73 282.8 272.7 17ltit1.7 iti1711.71 magnesium 0137 0142230.14223 00830.083 0 092130921.3009213 0068 0 065120651.2006512 sodium ppm 51151.1 540 537 600 71671.6 58058.0 sulfur 0200.20 019 070 0480.48 0140.14 0350.35 ph 474.7 4734.73 47 473 43 44124.412 tanninsfannins radius in cm 30530.5 2962329.623 175 1791317.91317 91391 3 26226.2 2502325.023 astringency aggmgg fw 858 830 487 66366.3 1043 958 sodium salts 292.9 303.0 30 33 29 34 soluble carbohydrates 160216.02 1620 173317.33 165816.58 FA GCMS count units 827905 399239 2609969 1259531 superscript values indicate differences between species at the p 1 level or less 1 oak 2 white fir 3 douglas fir values different between feed and nonfeed trees of the same species at or below p J1 values multiple locations in the same tree responsible for different n values used inin calculations of chemistry and climatic data climatic data n are the same as icportedreported in table 3 not comparable across species boundaries n values for factors below dashed line not as reported for rest of column not statistically comparable due to smaller sample size

RESULTS was not comparable among species but was greater for white fir feed trees than nonfeed oak and white fir feed trees were larger than trees p 1.1 ether extract fractions were nonfeed trees of the same species p 05os05.05 lower in oak than in conifers p 05os05.05 and table 1 herbivoreherbherbivoryivory by porcupines in decidu- lower in white fir than douglas fir p 1.1 ous species occurred in the canocanopiespies of large tissue from douglas fir contained less trees or in shrubs where branch diameters crude fiber than tissue from oak and white fir were relatively small in coniferous species p 0505.05 and douglas fir feed trees contained herbivoreherbivoryherbivory was also concentrated in the canopy still less than nonfeed trees p 1.1 water con- rather than on the tree bole only two instances tent was lower in oak tissue than in conifer tis- of chipping bark off the bole to expose the sue p 05os05.05 oak contained higher levels of inner bark were noted in our study both on potassium and calcium than douglas fir p deciduous tree species there were no trends 0505.05 white fir was also higher than douglas fir correlating calendar date or temperature to in calcium p 0505.05 magnesium levels for oak selection of feed tree species douglas fir feed were greater than for either conifer species p trees contained greater amounts of crude pro- 0505.05 white fir and oak tissue had higher ph tein than douglas fir nonfeed trees p 05os05.05 values than tissues from douglas fir p 05os05.05 crude protein content of both conifer species oak feed trees were higher in sodium salts was less than that of oak trees douglas fir p than douglas fir feed trees p 1.1 calcium 0505.05 white fir p 1.1 total tanninsfannins as mea- phosphorus ratios for feed trees were higher in sured by radial diffusion were higher in oak oak than in douglas fir p 0505.05 the calcium than in conifers douglas firpfirhirbir p 1.1 white firpfir p phosphorus ratio for oak is well above accept- 0505.05 astringency protein binding capacity able limits for mineral absorption by mammals 32 GREAT BASIN naturalist volume 55

underwood 1966 high calcium phosphorus locating porcupines in station trees compared ratios have also been reported by masslich to locating porcupines in rock or snow dens 1985 for aspen populus tremultremuloidesoides tissue there were approximately 707.0to porcupines utilized by beaver after an independent test km2 in the study area radio collared animals of feed tree tissue confirmed the high ratio were far ranging and did not utilize a single we tested the mineral content of feces and den or station tree as a base from which to urine from captive porcupines on an oak diet launch foraging expeditions rather they calcium phosphorus ratios from fecal material roosted and fed in a single tree for one to sev- were 101 while ratios from urine were eral days and then moved to another roost and approximately 2222111 1 feeding tree death loss due to predation and tissue samples from feed trees were ana- other causes left only 3 of 5 male and 6 of 10 lyzed by GCMS primarily as a check on ether female porcupines instrumented with radio extract fractions the small sample size did transmitting devices for the entire winter this not permit statistical analysis but trends sample size made statistical analysis of home showing lower fatty acid content in oak than in ranges unreliable several animals spent the conifers concurred with our observation of winter in relatively small areas but most had lower ether extract fractions in oak the relatively large overlapping home ranges amount of fatty acids was lower in oak than in male home range extremes were 686.8gs and 47547.5 either conifer species ha extremes for females were 929.2 and 61861.8 ha discriminant analysis correctly classified one female s home range overlapped those of feed trees as either conifer or oak 100 of the three males and at least four other females time table 2 six factors were important con- movements of up to 400 500 m between relo- tributorstributors to the model conifer feed trees had cations of some of the larger mature animals higher amounts of phosphorus and a greater were not uncommon some juvenile animals ether extract fraction than oak feed trees had reduced home ranges and movements alternatively oak feed trees were higher in which generally agrees with observations by protein calcium tanninsfanninstannins and magnesium roze 1989 mean distance from oak feed although tanninsfannins entered into the model they trees to a potential conifer feed tree was sig- were not a significant contributor these dif- nificantlynificantly less p 05og05.05 than the distance of an ferenferencesces between oak and conifer feed trees average move by a porcupine from an oak feed generally are in agreement with differences in tree to any other feed tree table 1 table 1 the classification of oak feed and non three of 15 porcupines 20 were eaten feed trees was less successful 71 table 2 by predators in a 4 mo period tracks in the oak feed trees were significantly higher in snow indicated that one porcupine was pur- sodium and fiber than nonfeed trees while sued worried and killed by two coyotes the nonfeed trees were higher in water content other two porcupines eaten by predators died porcupines used gambel oak as a food source late in the season on south facing slopes bare more often than it occurred in the study site of snow neither the cause of death nor carni- p values listed in table 3 six of 15 animals vore species could be positively determined were found roosting and feeding exclusively in carcasses of two other porcupines that died oak while 9 roosted and fed in conifer species presumably of starvation andor exposure dur- at least once snow depths and temperatures ing the course of the study were not scav- were analyzed for the winter period before the enged by coyotes main snowbeltsnowsnowmeltmelt judged to be 18 march average snow depths at porcupine location discussion for that period were 0.60ogo m sites time 060 chemical factors maximum snow depth was 1201.20 m median 0650.65ogs m mean minimum temperature for the dietary alternatives in the form of different night previous to locating study animals was feed tree species with significantly different 10cloc10 C the extreme low was 27c27 C mean chemical makeup were available to the study temperature for the night previous to locating population in winter vegetative oils have the animals in rock or snow caves was 12c12 C potential to be the most important source of there was no statistical difference between energy for porcupines data from ether extract the minimum nightly temperature previous to fractions derived from feed tree tissues indicate 199519951 WINTERING porcupines IN GAMBEL OAK 33

TABLE 2 standardized canonical discriminant function coefficients for factors that discriminated between oak and conifer feed trees 100 correct classification and between oak feed trees and oak nonfeed trees 71 correct classifi- cation oak vs coniferr feed trees n 56 oak feed vs nonfeed H treestrees n 46 coefficient prob b coefficient prob b phosphorous 1241.24 00001.00001 water content 0620.62 006006.006 ether extract fractions 0600.60ogo 0001.0001 sodium ogi0610.61 0202.02 protein 1181181.18 0005.0005 fiber 0590590.59 ooi001001.001 calcium 0390390.39 019019.019 tanninsfannins 0290290.29 its175.175 magnesium 0240240.24 006006.006 that gambel oak the major food source of our high fiber diet would be expected to maximize animals had lower values of ether extract frac- the intake of crude protein to compensate for tions than tissues from conifers evaluation of a low digestibility rate implications of a diet fatty acids by GCMS confirmed that fatty acid high in calcium and tanninsfannins are less clear but content was higher in conifer tissue additional it is possible that porcupines may deal with research on known digestible fractions is high levels of calcium in their food material by needed but until data indicating otherwise concentrating calcium in the urine tanninsfannins are presented we will operate under the function as protein binding agents rhoades premise that for porcupines conifers provide a and catesgates 1976 it is now evident that some greater source of useable fats than do oaks insects can circumvent tanninsfannins through a discriminant analysis was used to determine higher gut ph and the presence of surfactants if when all variables were taken together bernays 1981 martin and martin 1984 there would be general support from this martin et al 1985 however ph values for analysis with the t test significant differences the mid caecum 66gg666.6 and the pyloric 18181.8 and found by these analyses comparing oak and esophageal 32323.2 regions of the stomach of a conifer feed trees were in agreement tables laboratory porcupine on a diet of oak were 1 2 phosphorus and the ether extract frac- consistent with gut ph for monogastrics of tion were higher in conifer feed trees com- comparable size hume 1982 pared to oak feed trees and protein calcium oveson 1983 measured subcutaneous adi- taitattanninstantaiminskaiminsninsmins and magnesium were higher in oak pose concentrations on the rump of porcu- feed trees discriminant analysis was less suc- pines and reported a thickness of lsiisi15115.1 mm cessful in classifying feed and nonfeed trees 262.696gg mm in early winter by late february and within oak table 2 an important reason for early march fat reserves were virtually non- this less successful classification was that the existent A similar phenomenon was observed cloning nature of oak was emphasized by the by sweitzer and berger 1993 in nevada point quarter method this method may have where porcupine body condition decreased resulted in selecting nonfeed trees from the significantly throughout the winter season same clone as the feed tree future research those authors suggested the change in body should involve delineating the boundary of the mass was an indication that porcupines deplet- clone and selecting a nonfeed tree from a ed energy reserves early in the winter and were clone different from the feed tree clone stressed nutritionally during late winter the conifer roost sites also offer greater thermal heavy accumulation of fat serves as an energy advantages than deciduous roost sites clarke reserve for porcupines to draw upon through- and brander 1973 roze 1989 despite multi- out the winter allowing them to concentrate ple options porcupines depended heavily on on a food source relatively high in crude pro- an oak diet low in fats and associated thermal tein the reduced capabilities of protein advantages but higher in tanninstanfanninsnins the advan- digestibility associated with a high fiber diet tage of the oak diet may well be that it is high- may have encouraged our study animals to er in protein high levels of crude fiber eg maximize dietary protein by selecting oak cellulose reduce the digestibility of crude porcupine herbivoreherbherbivoryivory was generally noted on protein in monogastrics glover and duthie small branches in large trees porcupines fed 1958a 1958b therefore herbherbivoresivores on a high in the canopy where limbs are smaller 34 GREAT BASIN naturalist volume 55

TABUTABLE 3 chighi square analysis ofpercent occurrence and utilization of trees by porcupines occurrence used chicbl square value p value oak 435 82182.1 3233.23 010 conifer 272.7 16416.4 maple 52152.1 15 524152.41 ooi0010.01 conifer 272.7 16416.4 oak 435 82182.1881 591459.14 ooiool0010.01 maple 52152.1 151.5lsis n values differ komhomhornbomm those reported inin table I1 due to the extended use of some feed trees by porcupines occupancy of the same feed tree during more than one sampling event counted as multiple utilization of oak but not double sampled foiroirolforoor chemistry data df 1 we observed only two instances in which por- observed using dens during the winter or fol- cupines chipped bark of large tree boles and lowing runways in feeding areas they re- fed on tissue from large dahdbh limbs or trunks mained in the tops of hawthorne crataegus selection of larger feed trees by porcupines dougladouglasiisii thickets or utilized other deciduous may be related to the texture of bark and ease food sources throughout the winter sweitzer of climbing roze 1989 rather than chemistry and berger 1993 identified buffalo berry shepherdia willow salix app bit resource argentea spp deciduous food and roostingboostingRoosting terterbrushterbruschbrush purshia dentatatritridentatatridentate and juniper roze 1989 discussed the thermal advan- juniperus osteosperma as primary winter tages of dens andor conifer roost trees in rela- food sources of porcupines in nevada we tion to maintenance of a core body temperature have also observed the extensive use of hack- citing livingirving et al 1955 and clarke 1969 he berry celtis occidentoccioccidentalsoccidentaleoccidentalisdentalsalisails and green ash fratifraxi- indicated that the critical external tempera- nus pennsylvanicapennsylvanica by porcupines as a food ture below which porcupines must increase and roosting resource in the sand hills of their metabolic rates to maintain a core body nebraska and the missouri river breaks of temperature is a range between 12 and 4cac south dakota caves and conifers except plan- he suggested dens are temperature averaging tation forests and eastern red cedar juniperus devices that protect porcupines against convec- virginiajavirginianavirgivirginianalvirgininianaanal are not available in the sand hills tional and radiational heat loss station trees swinehart 1989 oveson 1983 reported that provide thermal advantages to porcupines a porcupine remained virtually motionless clarke and brander 1973 and may serve as a while perched in a gambel oak tree for a 24 h substitute for rock caves and snow dens period when the ambient temperature was as however none of these are requisite to porcu- low as 37c37 C during a 13 d period from 30 pine survival roze 1989 noted that porcu- january through 11 february when the mean pines may spend winters in trees away from low temperature was 17c17 C 3 of 25 12 loca- dens and that in every report the tree species tions of our study animals were in conifers 4 have been everevergreensgreens 16 were in rock or snow caves and 18 our data conflict with this observation 72 were in oak although porcupines did porcupines throughout western north america select trees with a larger dahdbh as roostingfeed are able to survive using a variety of deciduous ing sites they were also often found in smallish species as food and roost tree resources shrubs even though large trees were readily despite the prominence of literature concern- available it is therefore difficult to link possi- ing dens and conifer station trees use of a ble benefits presumed to be available to porcu- deciduous food and ostingroostingboostingro resource without pines that roost in larger trees such as protec- dependence on caves or snow dens is not an tion from the elements or from predators to anomaly for porcupines craig and keller s the selection shown by animals in this study 1986 study site in southern idaho was at an despite the availability of snow caves dens elevation of 1525 2089 in in desert shrub and conifer species that could provide thermal habitat animals in this study were not advantages the study population was heavily 199519951 WINTERING porcupines IN GAMBEL OAK 35 dependent on gambel oak for a roosting and periods in the winter or that more subtle cues feeding resource considering that this re- were used to transfer the information liance was during a season of energetic stress long movements between feed trees in it is likely that remaining motionless in the dense oak cover by some study animals sug- canopy of oak trees to conserve energy while gest that predator prey relationships may have exploiting a high protein food source is an influenced movements sweitzer and berger adaptive strategy 1992 found that habitat use was related to the age or size class of porcupines presum- predation size movements and ably in response to increased risk of predation the availability of conifer feed trees was not to smaller porcupines our observations gen- limiting since the average distance between erally agree with their findings mountain lion locations of study animals was significantly and coyote tracks were seen regularly in the greater than the mean distance of a move from study area both species are known to prey on any roost tree to a conifer roost tree table 1 porcupines keller 1935 robinetteRob mette et al 1959 it does not appear that spatial relationships of toweill and meslow 1977 maser and rohweder the various feed tree species played a role in 1983 the strong urine scent at station trees or feed tree selection by our study population dens makes porcupines readily detectable the relatively large overlapping winter home mountain lions are capable of knocking porcu- ranges of animals in this study differ from pines from the canopiescanopies of trees taylor 1935 reports of other researchers home ranges for if long moves decreased the predictability of porcupines in northwestern minnesota were mountain lions locating porcupines in station small enough to be reported in square meters trees it would be an adaptive strategy how- tenneson and oring 1985 curtis 1941 ever long moves expose porcupines to terres- dodge 1967 brander 1973 roze 1987 trial predation by mountain lions coyotes and 1989 and others have documented that por- wolves canis lupis1upiscupis which are now extirpated cupines move short distances from dens to from the study area and would presumably be feed trees sometimes along permanent trails nonadaptivenon adaptive since ample forage exists in the snow craig and keller 1986 and throughout the study site and long moves to locate food do to be a smith 1979 also reported reduced ranges in resources not appear the winter however dodge and barnes dietary necessity long movements may be an adaptive strategy to avoid arboreal predation 1975 did not indicate a similar restriction in by lions winter movements roze 1987 suggested the mountain this hypothesis deserves further reason may be crusted snows that bear the examination weight of the animals porcupines in our study acknowledgments did adeptly toboggan on crusted snows down slopes an attempt avoid extreme in to capture we thank S H jenkins and two anony- however one female moved over 450 in in mous reviewers for helpful suggestions to this fresh snow trails in powdery snow were often manuscript direct and suggested that a destination may have been predetermined literature CITED common use of oak and conifer feed trees by different porcupines occurred several times BERNAYS E A 1981 plant tanninsfannins and insect herbherbivoresivores during the study sometimes concurrently an appraisal ecological entomology 6 353 360 BRANDER R B 1973 life history notes on the hedging in the canopiescanopies of gambel oak trees in- porcupine in a hardwood hemlock forest in upper michigan dicated that some trees were used consistently michigan academician 5 425 433 over time by porcupines while others were CLARKE S H 1969 thermoregulatoryThermoregulatory response of the not consistent foraging in common trees over porcupine erethizon dorsatumdorsatum at low temperatures special indicate a learned behavior such as report department of forestry and wildlife time may management university of massachusetts amherst that described by glander 1981 for howler CLARKE S H AND R B BRANDER 1973 radiometric monkeys but we hesitate to attribute it to determination of porcupine surface temperature such because porcupine young of the year under two conditions of overhead cover were usually separated from their mothers physiological zoology 46 230 237 COTTAM G AND J T gurtisCURTIS 1956 the use of distance during the winter it is possible that some measures in phytosociological sampling ecology 37 young accompanied their mothers for limited 451 460 36 GREAT BASIN naturalist volume 55

CRAIG E H AND B L KELLER 1986 movements and MASER C AND R S ROHWEDER 1983 winter food home range of porcupines erethizon dorsatumdorsatum in habits of cougars from northeastern oregon great idaho shrub desert canadian field naturalist 100 basin naturalist 43 425 428 167 173 MASSLICH W J 1985 aspen beaver relationships in the CURTIS J D 1941 the silvicultural significance of the strawberry valley of central utah unpublished porcupine journal of forestry 39 583 594 master s thesis brigham young university provo dasilveira A J FE F FEITOSAFFITOSA TELES AND J W STULL UT 34 appp 1978 A rapid technique for total nonstructural car- MINITAB 1982 release 82182.182 1 copyright penn state uni- bohydratebo determination of plant tissue journal of versityversity state college PA agricultural and food chemistry 26 771 772 OVESON M C 1983 behavioral and metabolic adapta- DODGE W E 1967 life history and biology of the porcu- tions of porcupines erethizon dorsatumdorsatum to winter pine erethizon dorsatumdorsatum in western massachusetts stress unpublished master s thesis brigham young unpublished doctoral dissertation university of university provo UT 20 appp massachusetts amherst 167 appp RHOADES D AND R G cafesCATESGATES 1976 toward a general DODGE W E AND V G BARNES 1975 movements theory of plant herbivoreantiantiherbivore chemistry pages home range and control of porcupines in western 168 213 in biochemical interaction between plants washington USU S department of interior fish and and insects J W wallace and R W marshall eds wildlife service leaflet 507 recent advances in phytochemistry volume 10 GAMBLIEL H A R G CATESGATES M K CAFFEY MOQUIN plenum press new york london AND T D PAINE 1985 variation in the chemistry of ROBINETTE W L J S CASHWILERGASHWILER AND 0 W MORRIS loblolly pine in relation to infection by the blue stain 1959 food habits of the cougar in utah and nevada fungus pages 177 185 in S branham and R thatcher journal of wildlife management 23 261 273 eds proceedings of the integrated pest Manmangementmanagementgement ROZE U 1987 denning and winter range of the porcu- symposium asheville NC USDAUS DA forest service pine canadian journal of zoology 65 981 986 southern experiment station new orleans LA 1989 the north american porcupine smithsonianSmith soman GLANDERGLANDEK K E 1981 feeding patterns in mantled howl- institution press washington DC 261 appp ing monkeys pages 231 257 in A C kamil and T SAS 1985 copyright SAS inc carygary NC D sargent eds foraging behavior ecological etho- SSMITHmith A D 1957 nutritive value of some browse plants in logical and psychological approaches garland winter journal of range management 10 162 164 STPM press new york NY SMITH G W 1979 movements and home range of the GLOVER J AND D W DUTHIE 1958a the nutritive ratio porcupine in northeastern oregon northwest science crude protein relationship in ruminant and nonrumi-nonrumi 5327753 277 282 nant digestion journal of agricultural science 50 SWEITZER R A AND J BERGER 1992 size related effects 227 229 of predation on habitat use and behavior of porcu- 1958b the apparent digestibility of crude protein pines erethizon dorsatumdorsatum ecology 73 867 875 by ruminantsnonruminantsnon and ruminants journal of agricul- 1993 seasonal dynamics of mass and body condi- tural science 51 289 293 tion in great basin porcupines erethizon dorsadorsatumtum HAGERMAN A E 1987 A radial diffusion method for journal of Mammomammologymammalogylogy 74 198 203 determining tannin in plant extracts journal of chem- swineharteswinehartjSWINEHARSWINEHARTTJJ B 1989 windblownwind blown deposits pages 43 56 ical ecology 13 437 449 in A bleed and C flowerday eds an atlas of the horwitzwedHORWITZ WWEDED 1980 official methods of analysis of the sand hills resource atlas no 5 conservation and association ofofficial analytical chemists association survey division institute of agriculture and of official analytical chemists washington DC natural resources university of nebraska lincoln 1018 appp TAYLOR W P 1935 ecology and life history of the porcu- HUME I1 D 1982 digestion physiology and nutrition of pine erethizon epixanthum as related to the forests marsupials cambridge university press cambridge of arizona and the southwestern united states MA 256 appp university of arizona bulletin 6 1 177 IRVING L H H KROG AND M MONSON 1955 the TENNESON C AND L W ORING 1985 winter food pref- metabolism of some alaskan animals in winter and erences ofporcupinesof porcupines journal of wildlifeofwildlife manage- summer physiological zoology 28 173 185 ment 49 28 33 KELLER F L 1935 porcupines killed and eaten by a coy- TOWEILL D E AND C E MESLOW 1977 food habits of ote journal of mammalogy 16 232 cougars in oregon journal of wildlife management KUFELD R C M STEVENS AND D C BOWDEN 1981 4157641 576 578 winter variation in nutrient and fiber content and in UNDERWOOD E J 1966 the mineral nutrition of live- vitro digestibility of gambel oak quercus gambelligambeliigambelii stock central press ltd aberdeen great britain and big sagebrush artemisia tntridentata from diver- 237 appp sified sites in colorado journal of range manage- U S climatological DATA UTAH 1984 and 1985 ment 34 149 151 national climactic center asheville NC 86 12 and MARTIN J S AND M M MARTIN 1984 surfactants their 871387 13 role in preventing the precipitation of proteins by WELCH B L 1989 nutritive value of shrubs pages tanninsfannins in insect guts oecologia 61 342 345 405 424 in C M mckell ed biology and utilization MARTIN M M D C ROCKHOLM AND J C MARTIN of shrubs academic press inc new york NY 1985 effects of surfactants on precipitation of pro- teins by tanninsfannins journal of chemical ecology 11 received 18 august 1993 485 494 accepted 30 september 1994 great basin naturalist 551 0 1995 appp 374537 45 HISTORIC EXPANSION OF JUNIPERUS occidentalsoccidentaleoccidentalisOCCIDENTOCCIdenialsDENTALSallsALIS WESTERN JUNIPER IN southeastern OREGON

richard F millerlmilleramiller1 and jeffery A roselrosei

ABSTRACT the chronology of juniperusJumperus occidentoccidentalisoccidentaltsoccidentalistalisaltsaitsailsalls western juniper expansion in eastern oregon the effect of plant canopy and interspace on J occidentoccioccidentalsoccidentaleoccidentalisdentalsalisails seedling establishment and growth rates and the age ofJ occidentoccidentalisoccidentalistalis maximum reproductive potential were determined measurements were recorded in twenty two 040 4 ha plots estab- lished in sagebrush grassland communities and six 010 iha1 ha plots in populus tremuloidestremuloides quakingquaionglang aspen communities JJ occidentoccidentalisoccidentalistalis began increasing during the 1880s in stands containing trees 130 yr old relatively steady establishment ensued into the 1950s and then began to progress at a geometric rate in the 1960s JJ occidentoccidentalisoccidentalistalis encroachment into aspen stands began between 1910 and 1920 the largest proportion of juvenile trees established beneath artemisia species in sagebrush grassland communities J occidentoccidentalisoccidentalistalis trees appeared to reach full reproductive potential at 50 yr of age the ratio of malefemalemalemaie female trees increased from 171.71 7 in scattered J occidentoccidentalisoccidentalistalis stands to 383.83 8 in closed stands the initiation ofaofjofJ occidentoccidentalisoccidentalistalis encroachment during the late 1800s coincides with optimal climatic conditions for juniperusjumperus berry production and establishment reduced fire return intervals and heavy livestock grazing the accelerated increase in JJ occidentoccidentalisoccidentalistalis expansion since 1960 may be due to the continued absence of fire abundant woody plant cover and the large increase mjin J occidentoccidentalisoccidentalistalis seed production

key words western juniperjumper juniperus occidentoccidentalisoccidentalistalis expansion great basin intermountain shrub steppe aspen populus tremultremuloidesoides succession

one of the most pronounced plant commu- nah like nichol 1937 west 1988 or confined nity changes in the 20th century has occurred to rocky surfaces or ridges cottam and stewart in the juniper and pinyon juniper woodlands 1940 barney and frishknechtfrischknechtFrishknecht 1974 hopkins a major vegetation type characterizing the 1979 johnson and simon 1987 J occidentoccidentalisoccidentalistalis intermountain region these woodlands began increasing in both density and distribu- sometimes described as pygmy forests cur- tion in the late 1800s burkhardt and tisdale rently occupy 17 million ha throughout this 1976 young and evans 1981 eddleman 1987 region west 1988 juniperus occidentoccidentalisoccidentalistalis sspasp invading Atteartemisiamisiamisla tridentata subsp vastyanavaseyanavaseyana occidentoccidentalisoccidentalistalis hook western juniper is consid- mountain big sagebrush arteAtteartemisiamisiamisla arbuscula ered the northwest representative of the pin low sagebrush populus tremultremuloidesoides quaking yon juniper zone in the intermountain region aspen and riparian communities although JJ franklin and dyrness 1973 and occupies occidentoccidentalisoccidentalistalis is long lived vasek 1966 lanner over I1 million ha dealy et al 1978 in eastern 1984 less than 3 of the woodlands in oregon oregon southwestern idaho and northeastern are characterized by trees 100 years old california cronquist et al 1972 this sub- USDIBLMUSDI BLM 1990 in 1825 ogden observed species ofaofjof J occidentoccidentalisoccidentalistalis is found primarily only occasional J occidentoccidentalisoccidentalistalis reported as north of the polar front gradient neilson cedars growing on hillsideshillsides while traveling 1987 parallel to the oregon and nevada border through the crooked river drainage in central latitude 42 where temperatures are cooler oregon rich et al 1950 today these hill summer precipitation decreases and winter sides are covered by dense J occidentoccidentalisoccidentalistalis precipitation increases mitchell 1976 woodlands in a nearby area J W Meldrumeldrumrrsirs s relict juniper woodlands tree age class dis- 1870 survey notes describe a gently rolling tributiontribution fire scars and historical documents landscape covered with an abundance of peren- indicate presettlement pinyon juniper and nial bunchbuncligrassesbunchgrassesgrasses and a wide scattering ofaofjof J juniper woodlands were usually open savan occidentoccidentalisoccidentalistalis trees caraher 1977 today JJ

easternmasterneastern oregon agricultural research center HC 7145171 4 51 hwyhviydwy 205 bums OR 97720 the eastern oregon agricultural research center including the burns and union stations isis jointly operated by the oregon agricultural experiment station of oregon state university and USDA agricultural research service

37 38 GREAT BASIN naturalist volume 55 occidentoccidentalisoccidentalistalis densities on this site range expanding on the majority of sites measured between 125 and 250 ha 1 in silver lake early observations on steens mountain indi- oregon J occidentalisoccidentoccidentalistalis density increased from cate the landscape contained only scattered 62 ha 1 in 1890 to over 400 ha 1 by 1970 stands ofofaofjJ occidentoccidentalisoccidentalistalis griffiths 1902 since adams 1975 on another site in central 1900 the abundance ofofaofjJ occidentoccidentalisoccidentalistalis pollen in oregon where trees were absent prior to the steens mountain area has increased five- 1880 JJ occidentoccidentalisoccidentalistalis increased to 1018 ha 1 by fold mehringer and wigand 1990 1980 eddleman 1987 recent expansion is plant communities characteristic ofofaofjJ occi similar to increases in other juniperus species dentalisdenmentalistalis woodlands are artemisia tridentata throughout western united states ellis and sspasp vaseyanafestucavaseyanalfestuca idahoensis idaho fes- schuster 1968 tausch et al 1981 west 1984 cue artemisia arbusculaarbusculararbusculafarbusculalflF idahoensis and P tausch and west 1988 tremuloidestremuloides P tremultremuloidesoides communities on the objectives of our study were to 1 steens mountain range in elevation from 1760 describe the chronology ofaofjof J occidentoccidentalisoccidentalistalis to 2400 m at lower elevations in the J occi expansion during the past several centuries in dendentalismentalistalis woodland belt P tremultmmuloidestremuloidesoidesoldes stands southeastern oregon 2 determine the effect form long narrow communities along north of plant canopy and interspace on J occidenoccident aspects which capture windblown snow and talis seedling establishment and growth rates runoff and 3 determine the age when occidentalisoccidentalistalis J occident plot reaches maximum reproductive potential layout plot locations were selected in an attempt METHODS to reflect sagebrush grassland communities in different stages ofaofjof J occidentoccidentalisoccidentalistalis invasion on study area the west slope of steens mountain old stands the study area is located on steens moun- on the rocky outcrops which make up only a tain in southeastern oregon approximately 80 small percentage of present day woodlands km south of burns this isolated volcanic were not measured sites selected support or fault block which lies in the extreme north- have the potential to support sagebrush grass west basin and range province feiFerfennemanmeman land communities currently these sites are 1931 is about 80 km long and oriented in a occupied by varying numbers and sizes ofofaofjJ northeast direction baldwin 1981 the eleva- occidentoccidentalisoccidentalistalis dominance creating a woodland tion of steens mountain ranges from 1268 to structure of dispersed intermediate and 2949 m with a steep east facing escarpment closed tree stands table 1 twenty two 04 ha and a gentle west facing slope climate is cool plots were located within the J occidentoccidentalisoccidentalistalis and semiarid characteristic of the northern belt of steens mountain they ranged from 1500 great basin annual precipitation at the lower to 2000 m in elevation and were distributed 32 elevations averages 220 280 mm increasing to km along the mountain range plots were situ- 700 mm at higher elevations NOAA 1993 ated along an elevation gradient representing most moisture is received as snow in novem- communities from the lower to upper eleva ber december and january and as rain in tion J occidentoccidentalisoccidentalistalis woodland belt dominant march through june understory vegetation in the dispersed and I1J occidentoccidentalisoccidentalistalis woodlands on steens moun- intermediate plots was A dentatatritridentatatridentate sppapp tain form a discontinuous belt between 1450 vastyanavaseyanavaseyana and festuca idahoensis 13 stands and 2100 m in elevation severe winter condi- A arbuscula and FE idahoensis 4 stands and a tions probably restrict J occidentoccidentalisoccidentalistalis from ex- mosaic of A arbuscula and A tridentata sspasp panding into higher elevations billings 1954 vastyanavaseyanavaseyana 2 stands understory vegetation in mehringer 1987 limited distribution below the closed stands n 3 comprised a few 1500 rn is possibly due to a combination of late remnant deep rooted perennial grasses skele- spring frosts billings 1954 and limiting mois- tons of dead A dentatatritridentatatridentate sspasp vastyanavaseyanavaseyana and ture tree canopy cover varies from open to 70 bare ground EOARC data file 30 cover except on mesic P tremultremuloidesoides sites an additional six 01olha ha plots were estab- where J occidentalisoccidentoccidentalistalis cover approaches 100 lished in six separate P tremultremuloidesoides stands however based on age structure and canopy three stands were in advanced stages ofoffJ leader growth tree canopiescanopies are still actively occidentoccidentalisoccidentalistalis invasion with few to no adult P 199519951 WESTERN JUNIPER EXPANSION 39

TABLE 1 juniperusJumpetusperus occidentoccidentalisoccidentahsoccidentalistalisallsails stand maturity classes 2 obvious but not abundant and 3 abundant in artemisia communities modified from blackburn and each plot a 10 subsample tueller 1970 in tree was ran- domly selected for aging in each of four height closed abundant adult trees generally 5 m tall classes 1 05os050.5 m 2 050.5os 181.8isls m 3 181.8 3 m and usually several trees 130 yr of age 4 3 with little understory particularly on and m in several of the dispersed plots south slopes sample size for trees 3 m was smaller than 10 due to a lack of trees we also sampled all intermediate abundantjabundant occidentoccidentahsoccidentalisoccidentalistalisallsails cfallofallof allailali age classes old trees on plots when they occurred n 0 5 with a more open canopy and tree an I1 understory beginning to decline trees ha old trees were easily identified by their 130 yr of age are rare growth form containing rounded tops and heavy limbs and lacking strong terminal dispersed abundant 2 m tall few young trees a leader growth burkhardt and tisdale 1969 A adult trees but old trees absent and a well developed understory cross section was removed approximately 30 cm above ground level from each tree 05os050.5 m tall and at ground level for trees 050.5os m and tremultremuloidesoides trees and dead P tremultremuloidesoides then brought back to the lab for aging two trunks on the ground the remaining three radii from each cross section were polished stands were characterized by a dominant P stained and counted age was estimated by tremultremuloidesoides overstory and an understory of averaging both radii and adding 10 yr to cor- youngyoungjyoungaJ occidentoccidentalisoccidentalistalis elevation for thetheathejI1 occi rect for the 30 cm base mean differences dendentalismentalistalis P tremultremuloidesoides plots ranged from 1930 between radii were 4 for trees 50 yr and to 2000 m all with a similar northeast aspect 1 for trees 50 yr of age adams 1975 measurements reported that growth ring characteristics ofaofj occidentoccidentalisoccidentalistalis are useful in dendrochronological prior to sampling string was stretched along studies the presence of false and missing the contour of each 04 ha plot at imI1 m intervals rings was similar to that for pinus ponderosa to keep track of measured trees J occidentoccidentalisoccidentalistalis over 1200 trees were aged and approximately 1 density trees ha was recorded for trees 14000 counted and measured in the six PR os0.5 tall defined as 0505 m adult across the entire tremultremuloidesoides stands density of both J occidenoccident plot tree height minimal and maximal crown talis and P tremultremuloidesoides and age and height for diameters and basal above area just the trunk J occidentoccidentalisoccidentalistalis were measured across the entire swell at the stem base the litter layer near 010 1 haplotha plotpiot were recorded hallot tree height was measured evidence indicated minimal occidentalisoccidentalistalis with a tape for trees 2 m and a clinometer J occident mortality has occurred on steens mountain for trees 2 m tall tree canopy cover was during the past 120 years we observed very estimated by adding crown area measure- few dead or dying trees for all age classes ments of all trees for each plot similar mea- excluding seedlings except where surementssurements were recorded on juvenile trees individual occidentalisoccidentalistalis trees had been cut or burned defined as trees 050.5os m tall but only those J occident mortality of juniperus species rapidly declines on the lower left quarter oi01010.10 1 ha of each 040.4 following the seedling stage van pelt et al ha plot current year JJ occidentoccidentalisoccidentalistalis seedlings any plant with cotyledons still attached were 1990 juniperus has few pests that prove fatal not recorded establishment location of each to the tree lanner 1984 we avoided recently juvenile tree was recorded beneath the canopy cut or burned stands which constituted a offof J occidentalisoccidentalistoccident alis artemisia other shrubs tus- small percentage off occidentoccidentalisoccidentalistalis occupied sock grass or in the interspace less than 1 of stands where remains of dead trees were juveniles were located beneath other shrubs or observed we noted they persisted for a long grasses therefore onlyonlyjJ occidentoccidentalisoccidentalistalis ariearteATteartemisiamisia period of time by recutting several stumps and interspace are reported adjacent to one of our plots and aging and J occidentoccidentalisoccidentalistalis is considered submonoecioussubmonoecious matching ring widths with adjacent live trees vasek 1966 male and female reproductive we determined these trees were harvested status was determined by estimating abundance around 1920 others have also observed the of cones and berries for each tree abundance persistence of juniperusofjuniperus stumps young and was ranked in four classes 0 absent 1 scarce budy 1979 40 GREAT BASIN naturalist volume 55

statistical analysis characterized by an abundance of adult trees 3 m tall a tree canopy cover of 18 28 height growth data for adult trees were table 2 and the presence of a few old trees analyzed using a randomized complete block 130 yr 2 to 5 hatihacihai J occidentalisoccidentalistalis densities design PROC GLM of SAS SAS 1986 J occident in began increasing in these stands between means were separated using duncan s 1878 and 1890 in the intermediate J occidenoccident multiple range test at p 05os05.05 level A split J talis stands trees 130 yr were rare tree plot design was used in the analysis ofjuvenile I1 canopy cover ranged from about 8 to 16 and height growth main plots were sites and subSUD densities of adult trees varied from 35 to 100 plots were location of establishment interspace hahaalha7l1 trees 3 m in height particularly juve- artemisia J occidentalisoccidentalistalis A duncanduneanduncans s multiple J occident niles were abundant J occidentalisoccidentalistalis expansion range test was used to separate the means J occident in these sagebrush grassland communities began between 1890 and 1910 in the dis- RESULTS persed stands few trees were 60 arsyrs old and we aged no trees 100 yr tree canopy cover little change in JJ occidentoccidentalisoccidentalistalis density usually 5 the stands and appeared to occur between the early 1700s was in dispersed 35 ha 1 and the 1880s fig 1 we encountered old densities of large adult trees trees standing trees 130 years old large invasion of J occidentoccidentalisoccidentalistalis into these sage stumps and burned out trunks on several A brush grassland communities began after arbuscula flats and A tridentata sspasp vastyanavasevaseyanayana 1930 communities however data indicated preset- greatest densities ofofaofjJ occidentoccidentalisoccidentalistalis trees tlement tree densities in these artemisia com- measured on steens mountain occurred in P munimunitiesties were 5 trees ha 1 suggesting very tremultremuloidesoides sites table 3 in the late stages of open JJ occidentoccidentalisoccidentalistalis stands the first evidence J occidentoccidentalisoccidentalistalis succession on these sites tree of an increase in tree densities occurred in the canopy cover approached 100 live P 1880s with relatively steady establishment tremultremuloidesoides occurred only on one of the three ensuing into the 1950s similar to that sites and almost all trees were 05os050.5 m tall in observed by tausch and west 1988 in the the remaining two stands only the remnants of 1960s J occidentoccidentalisoccidentalistalis establishment began large P tremultremuloidesoides trunks decaying in the occurring at a geometric rate understory were present J occidentoccidentalisoccidentalistalis inva- closed JJ occidentoccidentalisoccidentalistalis stands which once sion in these P tremultremuloidesoides sites began be- supported A tridentata sspasp vastyanavaseyanavaseyana were tween 1910 and 1920 no JJ occidentoccidentalisoccidentalistalis trees

7

J

6 F 0 F 5

LL

0 4

Z 3 LU L cc 2 LU

0 1700 1720 1740 1760 1780 1800 1820 1840 1860 1880 1900 1920 1940 1960 1980 YEAR

fig 1 years of establishmentofestablishment for Junijuniperuspetgpergperm occidentoccidentalisoccidentalistalis trees on steens mountain oregon n 1200 199519951 WESTERN JUNIPER EXPANSION 41

TABLE 2 general description of closed intermediate and dispersed juniperusJumperus occioccidentalsoccidentaleoccidentoccidentalisdentalsalisails stands on steens mountain in artemisia tndentata sspasp vastyanavaseyanavaseyana and A arbuscula communities and the percentage of juvenilesofjuveniles located beneath J occidentoccidentalisoccidentahsoccidentalistalisails artemisiaArtemma and interspace canopy cover basal area and density means are followed by range in paren- theses

establishmentEstestablisestablishablisihment site cvo density ha 1 for juveniles canopy basal area adults juveniles sites cover maham2ha 1 05050.50 5 m ht 05050.50 5 m ht f occidentoccidentalisoccidentalistalis aileAtleatlemisiaartemisiamisiamisla interspace A tntritrldentata sspasp vastyanavaseyanavaseyana closed 6 22 18 28 525.25 2 31313.13 1 9.8998 8 296 217217496496 580 118 1226 86186 1 ab9b ab5b intermediate 8 6 5 10 18 05 47 95 50 165 815 335 1423 291 5858a ly dispersed 2 2 1 3 04 02 06 52 31 70 188 96 280 ab3b 501soa 47a A arbuscula closed 3 15 12 20 353 5 18181.81 8 545 4 158 74 247 99 20 198 2727b 67 6cac intermediate 3 6 454 5 676.76 7 18 09 32 104 77 153 375 167 790 lib 61gia 2811 isitesmisitessitessltes of establishment means followed by similar lowercase letters are not significantly different between establishment sites within J occidentalisoccidentaltsoccidentalistoccident alisaltsaits stand maturity classes p 05

80 yr were encountered in stands with a P and female and 30 contained neither fruits tremultremuloidesoides overstory P tremultremuloidesoides density of nor cones JJ occidentoccidentalisoccidentalistalis trees producing abun- small shoots was greater than that ofofaofjJ occi dant crops of cones or berries were either dentalisdenmentalistalis however P tremultremuloidesoides size classes male or female dominant no trees were mea- between 050.5os m and large adults were absent sured which contained an abundant crop of indicating a lack of P tremultremuloidesoides stand reju- both berries and cones sixty five percent offof venation on these sites JJ occidentoccidentalisoccidentalistalis inva- occidentoccidentalisoccidentalistalis trees with an abundant crop of sion began between 1930 and 1940 berries contained no male cones the remain- height growth for young JJ occidentalisoccidentoccidentalistalis ing 35 contained only a scarce number of trees 20 yr across all sites averaged cones the majority of trees producing abun- 292.9 cm yiyllyliyr 1 based on growth rates and height dant crops of male cones contained only of trees between 10 and 20 yr of age n 200 scarce numbers of berries approximately 75 across all artemisia sites 90 of trees 15 yr of trees producing heavy crops of berries or old were 1 I1 m tall 64 were 05050.5 m tall cones were 50 yr old trees 20 yr old ex- surprisingly height growth rates of juvenileofjuvenile pressing reproductive effort were rare and trees did not significantly differ between A produced only a few cones or berries the arbuscula and A dentatatritridentatatridentate sspasp vastyanavaseyanavase yana ratio of trees producing large crops of cones communities however location of establish- versus berries conesberriescones berries increased from ment within communities significantly influ- 171.7 in the scatteredfscattered J occidentoccidentalisoccidentalistalis stands to 383.8 in the closed stands enced growth rates of young JJ occidentoccidentalisoccidentalistalis trees table 4 trees establishing beneath an artemisia canopy grew faster than young trees discussion growing in the interspace low densities and limited distribution shrub and tree canopiescanopies also significantly ofaofj occidentalisoccidentalistalis trees 130 yr and limited num- influenced location occidentalisoccidentalist seedling occident ofaofj occidentalis bers of dead trees or old stumps suggest establishment in artemisia communities J the occidentoccidentalisoccidentalistalis has greatly expanded on steensS teens largest ofjuvenile proportion ofjuvenile trees was usually mountain during the past 100 yr distribution located beneath canopiescanopies ofaof A tridentata sspasp of old trees was generally limited to rocky vastyanavaseyanavaseyana or A arbuscula and JJ occidentalisoccidentoccidentalistalis ridges and A arbuscula communities old table 2 less than 20 of juvenilesofjuveniles across all trees were found only occasionally growing in 22 artemisia sites established in the interspace deeper well drained soils such as A tridentata on steens mountain for trees 05050.5 m tall sspasp vastyanavasevaseyanayana grassland communities and 32 expressed predominantly only male or were absent in P tremultremuloidesoides communities in only female characteristics 38 both male northeastern california barbour and major 42 GREAT BASIN naturalist volume 55

TABLE 3 mean densities ha 1 followed by range in of populus tremultremuloidesoides and juniperusJumperuspeyus occidentoccidentalisoccidentahsoccidentalistalisallsails in P tremultremuloidesoides sites P tremultremuloidesoides occidentalisoccidentahsoccidentalistalisallsails stagee ofr J occident succession adult juvenile adult juvenile late n 3 17 1316 1392 9462 0 50 0 3952 929 2203 4327 18791 intermediate 1060 6553 1090 2816 nan3n 3 476 1670 5266 9480 63263217391739 622 5968

1977 found a similar distribution of old and support this hypothesis fires set by native young J occidentoccidentalisoccidentalistalis trees A tridentata sspasp americans also declined in the 19th century vastyanavaseyanavaseyana and A arbuscula communities which due to large reductions in their populations contained a low density ofaofjofJ occidentalisoccidentoccidentalistalis trees caused by european diseases thompson prior to settlement were the earliest sites to 1916 cressman 1981 and relocation to reser- initiate an increase in J occidentoccidentalisoccidentalistalis dates of vations in the 1870s initial establishment of closed and intermedi- the invasion of conifers into P tremultremuloidesoides ate stands were similar to periods of early communities is a common occurrence through- stand development reported by young and out the western US however conifers report- evans 1981 in northeastern california and ed to typically invade P tremultremuloidesoides stands are eddleman 1987 in central oregon species adapted to more mesic sites such as expansion ofofaofjJ occidentoccidentalisoccidentalistalis coincides with pinus confortacontortacontorta lodgepole pine P ponderosaponderosal euro american settlement in this portion of pseudotsuga menziemenziesiimenziesiasii douglas fir abies the great basin although no direct cause and concolor white fir abies lasiocarpalasiocampa sub- effect relationship can be drawn we hypothe- alpine fir picea engelmanniiengelmanniaengelmannii engelmann size that climate altered fire frequencies and spruce and picea pungentpungens blue spruce bartos grazing in the late 1800s were primary factors 1973 mueggler 1985 invasion of the more initiating the recent expansion ofaofj occidentoccidentalisoccidentalistalis drought tolerant J occidentoccidentalisoccidentalistalis into P tremuteemu following the end of the little ice age in the loides stands is not well documented mid 1800s bryson 1989 winters became more P tremuloidestremuloides is frequently considered a mild and precipitation increased above the fire induced species replaced by less fire tol- present longtermlong term average in the northern half erant conifers baker 1925 daubenmire 1943 of the great basin between 1850 and 1916 mueggler 1976 prior to settlement lightning antevs 1948 graumlich 1985 mild wet win- and human set fires probably helped maintain ters and cool wet springs promote vigorous many P tremultremuloidesoides communities however growth in J occidentoccidentalisoccidentalistalis earle and fritts the occurrence of fire in P tremultremuloidesoides stands 1986 fritts and Xiangxiangdigxiangdingdig 1986 in the rocky mountains has been greatly presettlement fire return intervals in A tri reduced since the late 1800s jones and debyle dentata sspasp vastyanavaseyanavaseyana communities have been 1985 mueggler 1985 suggested the combi- reported to vary from 15 to 25 yr houston nation of fire suppression and heavy grazing in 1973 burkhardt and tisdale 1976 martin and P tremultremuloidesoides communities may favor the estab- johnson 1979 burkhardt and tisdale 1976 lishmentlishment of conifers concluded that fire frequency intervals of an increase in artemisia cover may also 30 40 yr would be adequate to keep I1J occi enhance the invasion of J occidentoccidentalisoccidentalistalis As a dendentalismentalistalis from invading a sagebrush grassland sagebrush grassland community shifts towards community following settlement frequency a greater dominance of shrubs the number of of fire in sagebrush grasslands has greatly safe sites for J occidentoccidentalisoccidentalistalis seedling establish- declined the reduction of fine fuels by high ment increases others have also reported the densities of domestic livestock greatly reduced majority ofJ occidentoccidentalisoccidentalistalis seedlings established the potential for fire in the intermountian beneath artemisia canocanopiespies burkhardt and shrub region burkhardt and tisdale 1976 tisdale 1976 eddleman 1987 in west texas west 1988 griffiths 1902 observations of J pinchopinchotiipinchottitii frequently establishes beneath the overovergrazedgrazed landscape on steens mountain mesquite plants mcpherson et al 1988 199511995 WESTERN JUNIPER EXPANSION 43

TABLE 4 mean growth rates for juvenile juniperus occi acknowledgments dentalisdenmentalistalis trees 2 30 yr old in three different establish- ment sites this is technical report 10494 of the establishment site cm yr 1 eastern oregon agricultural research center oregon artemisia 33a33. state university

JJ occidentoccidentalisoccidentalistalis 2727ab literature CITED

interspace 24b ADAMS A W 1975 A brief history of juniper and shrub means followed by similar uppercase letters are not significantly different p populations in southern oregon wildlife research 05 report 6 oregon state wildlife commission corvallis ANTEVS E 1948 climatic changes and pre white man great shading by nurse plants may benefit occi pages 168 191 in the basin with emphasis on J glacial and postglacial times university of utah dendentalismentalistalis seedlings johnsen 1962 by reducing biological service bulletin 107 summer surface temperatures by 45 57 of BAKER FE S 1925 aspen in the central rocky mountain bare ground surface temperatures burkhardt region USDA bulletin 1291 and tisdale 1976 enhanced growth rates of BALDWIN E A 1981 geology of oregon kendalhuntKendal Hunt publishing co dubuque IA young trees growing beneath A tritridentatatridentate dentata ST BARBOUR MGM G AND J MAJOR 1977 terrestrial vegeta- vastyanavaseyanavaseyana suggest micromicroclimatesclimates beneath shrub tion of california wiley interscience new york NY canopiescanopies are more beneficial than conditions BARNEY M A AND N C frishknechtfrischknechtFRISH KNECHT 1974 vegetation in the interspace burkhardt and tisdale changes following fire in the pinyonpryonpmyon juniperjumper type of central journal of range management 1976 reported occidentalisoccidentalistalis seedling growth west utah J occident 74 91 96 rates were correlated positively with artemisia BARTOS D L 1973 A dynamic model of aspen succes- and correlated negatively with bare ground sionslon pages 13 25 in proceedings IUFRO biomass JJ occidentoccidentalisoccidentalistalis approached full reproductive studies IUFRO potential near 50 yr As occidentoccidentalisoccidentalistalis densities BILLINGS W D 1954 temperature inversions in the J zone of nevada range increased the proportion of trees became pre- pinyon juniperjumper a mountain butler university botanical studies 12 dominantly male across sites highly fecund BLACKBURN W H AND P T TUELLER 1970 pinyon and female trees appeared to be most important in juniper invasion in black sagebrush communities in open stands where J occidentalisoccidentoccidentalistalis was actively east central nevada ecology 51 841 848 expanding in central oregon eddleman BRYSON R A 1989 late quaternary volcanic modulation of milankovithmilankomilankovitchvith climate forming theoretical applied 1984 observed that trees in the interior climatology 39 115 125 woodlands were strongly dominated by male BURKHARDT J W AND E W TISDALE 1969 nature and cone production while trees growing in the successional status of western juniper vegetation in open produced more female cones he also idaho journal of range management 22 264 270 reported trees did not produce significant 1976 causes ofjuniperof juniper invasion in southwestern quantities of fruit until 50 70 yr of age idaho ecology 76 472 484 CARAHER D L 1977 the spread of western juniper in conclusion central oregon pages 3 8 in R E martin J E dealy and D L caraher eds proceedings western optimal climatic conditions around the juniper ecology and management workshop turn of the century reduced fire return inter- USDA forest service general technical report pnw4 vals and the effect of livestock indirect COTTAM W P AND G STEWART 1940 plant succession as through the reduction of fine fuels and an a result of grazing and of meadow desiccation by increase in artemisia cover are probably pri- erosion since settlement in 1892 journal of forestry mary factors that have contributed to the rapid 3861338 613 626 of occidentoccidentalisoccidentalistalis in southeast CRESSMAN L S 1981 the sandal and the cave oregon expansion ofaofjJ state university press corvallis oregon 1800s early 1900s during the late and CRONQUIST A A H HOLMGREN N H HOLMGREN AND the accelerated increase in J occidentoccidentalisoccidentalistalis J L REVEAL 1972 intermountain flora vascular density and invasion during the last 30 years plants of the intermountain west USA volume 1 into new communities is probably largely due hafner publishing company new york NY absence of fire abundant daubenmire R F 1943 vegetational zonation in the to the continued rocky mountains botanical review 9 325 393 woody plant cover and the large increase in J DEALY J E J M GEIST AND R S DRISCOLL 1978 occidentoccidentalisoccidentalistalis seed rain westeinwestern juniper communities on rangeland of the 44 GREAT BASIN naturalist volume 55

pacific northwest pages 201 204 in D E hyder grasslands american midland naturalist 102 ed proceedings first international rangeland 391 397 congress denver CO MEHRINGER P J JR 1987 late holocene environments EARLE C J AND H C FRITTS 1986 reconstructing on the northern periphery of the great basin final river flow in the sacramento basin since 1560 report bureau of land management portland OR report california department of resources agree- MEHRINGER P J JR AND P E WIGAND 1990 compari- ment DWR B 55395 laboratory of tree ring son of late holocene environments from woodratwoodral research university of arizona tucson biddensmiddens and pollen diamond craters oregon pages EDDLEMAN L E 1984 ecological studies on western 294 325 in J L betancourt T R van devender juniper in central oregon pages 27 35 in T E bedell and PR S martin eds pankratpackrat biddensmiddens the last ed proceedings western juniper management 40000 years of biotic change university of arizona short course oregon state university and extension press tucson service corvallis MITCHELL V L 1976 the regionalization of climate in 1987 establishment and stand development of the western united states journal of applied western juniper in central oregon pages 255 259 in meteorology 15 920 927 R L everett ed proceedings pinyon juniper MUEGGLER W F 1976 type variability and succession in conference USDAUSDA forest service general technical rocky mountain aspen pages 16 19 in proceedings report INT 215 utilization and marketing tools for aspen manage- ELLIS D AND J C SCHUSTER 1968 juniperjumper age and dis- ment in the rocky mountains USDA forest service tributiontribution on an isolated butte in garza county texas general technical report RM 29 southwestern naturalist 13 343 348 1985 vegetation associations pages 45 55 in N V FENNEMAN N M 1931 physiography of the western debyle and R P winokur eds aspen ecology and united states mcgraw hill new york NY management in the western united states USDA FRANKLIN J F AND C T DYRNESS 1973 natural vegeta- forest service general technical report RM 119 tion of oregon and washington USDA forest NEILSON R P 1987 on the interface between current service general technical report PNW 8 portland ecological studies and the paleobotany of pinyon OR juniper woodlands pages 93 98 in R everett ed FRITTS H C AND W XIANGDIG 1986 A comparison proceedings pinyon juniper conference USDA between response function analysis and other re- forest service general technical report INT 215 gressiongres sion techniques tree ring bulletin 46 31 46 NICHOL A A 1937 the natural vegetation of arizona GRAUMLICH L 1985 longtermlong term records of temperature university of arizona technical bulletin 68 and precipitation in the pacific northwest derived NOAA 1993 national climatic data center federal from tree rings unpublished doctoral dissertation building asheville NC university of washingtonofwashington seattle RICH E E A M JOHNSON AND B R BAKER EDS 1950 GRIFFITHS D 1902 forage conditions on the northern peter skene ogden s snake country journals 1824 25 border of the great basin bureau of plant industry and 1825 26 the hudson bay society london USDA bulletin 15 SAS 1986 SASSTATSAS STAT user s guide release 603 SAS HOPKINS W E 1979 plant associations of the fremont institute inc carsgarscarygary NC national forest USDA forest service pacific TAUSCH R J N E WEST AND A A NABI 1981 tree age northwest region r6ra ECOL 79 004 and dominance patterns in great basin pinyon HOUSTON D B 1973 wildfiresWildfires in northern yellowstone juniper woodlands journal of range management national park ecology 54 1109 1117 3425934 259 264 JOHNSEN T N 1962 one seed juniper invasion of north- TAUSCH R J AND N E WEST 1988 differential establish- ern arizona grasslands ecological monographs 32 ment of pinyon and juniper following fire american 187 207 midland naturalist 119 174 184 JOHNSON C G JR AND S A SIMON 1987 plant associa-assoriaassocia THOMPSON D 1916 david thompsonsthompsodsThompsodssons narrative J B tions of the wallowagallowa snake province wallowagallowa tyrrel ed the champlain society toronto ontario whitman national forest USDA forest service canada pacific northwest region report r6ra ECOL TP USDIBLMUSDI BLM 1990 the juniper resources of eastern 255138625513 86 portland OR oregon USDA bureau of land management infor- JONES J R AND N V DEBYLE 1985 fire pages 77 81 inm mation bulletin OR 90 166 N V debyle and R P winokur eds aspen ecology VAN PELT N R STEVENS AND N E WEST 1990 survival and management in the western united states and growth of immature juniperusJumperus osteospermaosteospenna and USDA forest service general technical report finnsfinuspinus adulisedulis following woodland chaining in central RM 119 utah southwestern naturalist 35 322322328328 LANNER R M 1984 trees of the great basin a natural VASEK F C 1966 the distribution and taxonomy of three history university of nevada press reno western junipersjunijuniperuspers bnttoniabrittaniabrittoniaBrittonia 18 350 372 MARTIN R E AND A H JOHNSON 1979 fire management WEST N E 1984 successional patterns and productivity of lava beds national monument pages 1209 1217 of pinyon juniper ecosystems pages 1301 1332 in in R M linn ed proceedings first conference of developing strategies for range management science and research in the national parks USDI westviewWestview press boulder CO national park service transactions proceedings 1988 intermountain deserts shrub steppes and serial 5 woodlands pages 209 230 in M B barbour and W D MCPHERSON G R H A WRIGHT AND D B WESTER billings eds north american terrestrial vegetation 1988 patterns of shrub invasion in semiarid texas cambridge university press cambridge MA 199511995 WESTERN JUNIPER EXPANSION 45

YOUNG J A AND J D BUDY 1979 historical use of received 7 february 1994 nevada s pinyon juniper woodlands journal of forest accepted 8 june 1994 history 23 113 121 YOUNG J A AND R A EVANS 1981 demography and fire history of a western juniper stand journal of range management 34 501 506 great basin naturalist 551 C 1995 appp 46 57

RANGELAND ALPHA diversities HARVEY VALLEY LASSEN NATIONAL FOREST californiaCALIFORNIAl raymond D Ratliffratlifffratliff22

ABSTRACT monitoring diversity usually begins by estimating alpha diversity of a plant community on a specific site the objectives of this study were to provide alpha diversity benchmarks and to determine whether rangeland com- munity basal cover characteristics explained variation in diversity estimates plant and surface component cover per- cencentagestages were estimated on 51 plots representing four vegetation types on the lassen national forest CA each plot was sampled with 30 random 102 basal point transects jackknife procedures were used to compute means and standard errors for margalef s diversity index drnarnD which stresses species richness and simpsonSimpsorssotss index DJdg which stresses species dominance within vegetation types Ddu and dgD did not rank all plots in the same order highest dmD values occurred with the most species highest dgD values occurred with comparatively few species but more uniform cover with either index average diversity declined from the meadow to grassland to open shrub grass to timber bunchgrass types all possible subset regressions of diversity on the basal cover characteristics were computed portions of the vari- ance accounted for by the best models were too low to allow prediction of dmD and Ddg the relation of alpha diversity to rangeland health is discussed

key words ecology plant communities Margmargalefalefss index simpson s index monitoring basal cover

biological diversity hereafter called diver- diversity may or may not follow traditional sity involves ecological processes structures concepts of succession and increase from pio- and functions and may occur at any spatial scale neer to climax plant communities or decrease society of american foresters 1992 diversity with rangeland deterioration over large areas refers to variety and abundance it is variety or diversity may be higher if communities are at multiformity of different forms or kinds several seraiseral stages than if the entire area is at stein and urdang 1966 there are alpha a single seraiseral stage within specific sites physghys beta and gamma diversities whittaker 1972 icalchemical factors or intense competition or alpha diversity is the variety that occurs with- both may work to reduce diversity odum in a plant community of a specific site A site 1959 absence of an expected species may be or stand is defined as an individual unit that is due to frequent disturbances a low immigration homogeneous in vegetation soil topography potential an immature soil or an inhospitable microclimate and history west 1993 beta moisture regime del moral and wood 1988 diversity is the variety of communities along a nevertheless because it may change with gradient eg topography soil acidity or mois- the kind of management diversity should be ture regime or on a given site through time assessed as part of range health evaluations gamma or large scale diversity is the variety diversity indices provide information that of plant communities or the total number of may not be immediately apparent from basic species present or both in a specific geo- measures of the plant community such as graphic area eg grazing allotment or water- cover and composition high diversity of plant shed species is important in maintaining processes diversity has two components richness and flow pathways for energy and nutrients and evenness ludwig and reynolds 1988 within and among communities higher diver- Mamagurranmagureangurran 1988 richness refers to variety sity implies a greater number of occupied numbers of species for example evenness niches whittaker 1972 refers to equality abundance or numbers of protecting or enhancing diversity or both species botanical composition for example are goals commonly set by policy or law west

ithisathishis articlearticie was written and prepared by USU S government employees on official time it is therefore in the public domain and not subject to copyright pacific southwest researchRe seaich station USDA forest service 2081 E sienaslenasierra fresno CA 93710

46 199511995 ALPHA DIVERSITY 47

1993 gave four reasons for having diverse the objectives of this study were 1 to pro- plant communities a sense of moral obligation vide local rangeland managers with indices of to living things an aesthetic appreciation of alpha diversity from plant communities to use nature economic benefits possible from them as guides of expected diversity for similar eg the gene pool for cultivated crops and sites and 2 to question whether variation in the important array of services they provide basal cover percentages of common and eg maintaining oxygen levels and cycling important indicators of rangeland health could nutrients explain variation in diversity although the A major cause of rangeland deterioration is findings are specific to the study area it is selective grazing of preferred plants and sites hoped they may assist others dealing with in similar patterns each year hormay 1970 questions of plant species diversity on range- even with conservative grazing populations lands of preferred plants on preferred sites may dis- appear thereby reducing the overall diversity METHODS of vegetation if such populations are ecotypesecotypes odum 1959 the ability of the species to study plots recapture site resources is reduced during 1964 and 1965 51 plots were estab- because nature abhors a vacuum other lished on the harvey valley and neighboring species may increase or invade as those pre- grazing allotments of the lassen national ferred by livestock decrease in abundance forest CA ratliffratliffetRatliffetet al 1972 the plots were dyksterhuis 1949 As a result plant species either 010.1oi ha or 020.2 ha and unevenly distrib- diversity may be higher rather than lower uted among meadow 8 open grassland 13 under grazing at least initially As preferred open shrub grass 12 and timber bunchgrass species decrease and less preferred ones in- 18 vegetation types these plots were used crease their abundances tend to become more for evaluating range condition health at even dyksterhuis 1949 with continued harvey valley relative to the neighboring deterioration species not previously able to allotments compete tend to invade and become established meadows ranged from ephemeral lake sites and thereby increase species richness the new with hardhardpanspans to deep organically rich soil of plant community though possibly comprising drainage bottoms open grasslands included more species that are more evenly abundant those dominated by shorthair sedge carex may cover less total area and higher diversity exsertdexexsertaexsertedserta and those where shorthair sedge had may be associated with greater amounts of been replaced by grasses open shrub grass bare soil areas included silver sagebrush artemisia increasingly land managers are asked to canacand black sagebrush A arbuscula big sage- monitor and determine change in diversity brush A tridentata and bitterbitterbrushbrush purshia monitoring diversity usually starts with an esti- tridentata subtypes the timber bunchgrass mation of alpha diversity for plant communities types were all in second growth ponderosa on specific sites such estimates are rare for pine pinus ponderosa some of them had bit rangelands to derive the greatest benefit from terterbrushterbruschbrush and big sagebrush along with grasses monitoring efforts managers must know what in the understory constitutes high and low diversity in given situa- tions they need to know how diversity changes data collection when other commonly estimated properties of data used to estimate alpha diversity on the site change eg litter cover and amount each plot were actual point contacts hits with of bare soil plant bases or soil surface components gravel seldom will examples of pristine or climax litter rock bare soil and large woody debris plant communities be available for developing and shrub crown area A hit on a shrub was diversity guides current plant communities recorded when a point contacted the shrub represent the sums of all past influences crown or was within its projected crown area current vegetation and site characteristics at the soil surface for each plot 3060 hits were therefore must serve as benchmarks from recorded consisting of 102 points in regularly which to develop guides and evaluate future spaced 3 point quadratsquadquadransrats on each of 30 ran- change domly placed transects points in a quadrat 48 GREAT BASIN naturalist volume 55

were at 23 cm centers and projected vertical- overall estimate a pseudovaluepseudovalue related form ly within transects quadrat spacing was was computed from the pseudovaluespseudovalues means either 060.6og m or 090.9og m depending upon plot and standard errors for the two indices were width basal cover percentages proportions of derived for each plot use of the jackknife pro- the surface occupied by different plants and cedure to improve estimates of diversity and surface components were calculated from the provide a way of calculating confidence inter- hits and summarized ratliffratliffetRatliffetet al 1972 vals was suggested by Mamagurranmagureangurran 1988 diversity indices basal cover relationships two indices of diversity were used 1 contributions of basal cover of various char- margalefmargalefsMarg alefss fDdn S llnallnlin N I1 where S is acteristicsacteristics to the variance in estimates of alpha the number of species and N is the total num- diversity were examined characteristics for ber of individuals hits for all species and 2 each plot were basal covers of grasses grass- simpsinssimpsonsSimpsons like herbaceous plants forbs shrubs and soil s surface components all possible subset re- Dsdg 1dad where D I1 ininianinin I1 gressionsgressions of dmD and Dsdg on the characteristics il were computed using the mallow s cp criteri- NN IJ1 and nin is the number of individuals on of the REG procedure a multiple linear here the percentage cover of the ith species regression program of the SAS institute inc Mamagurranmagureangurran 1988 dmD was selected for its sim- 1982 subset regression models explaining plicity and because it stresses the species rich- most variation in the indices were selected for ness component Dsdg was selected because it is study the pearson correlation matrix was com- well known and stresses the species evenness puted using the correlations CORR module dominance component in addition these of SYSTAT wilkinson 1989 to help assess indices were selected because they do not the influence of individual characteristics on require testing assumptions regarding the the indices underlying distributions of species abundance an overall estimate of diversity was com- RESULTS for each each index 30 puted plot using then alpha diversity indices new diversity estimates were computed using the jackknife procedure this procedure con- diversity indices and basal cover values are sisted of deleting each transect in turn from available for all 51 plots here only those plots the data set from each new estimate and the within each vegetation type ranking lowest

TABLE 1 numbers of species dominant species and percentage composition and jackjaekjackknifedjacklmifedknifed means and standard errors SE for margalef s and Simpsimpsonsorss diversity indicindicesindiceslindices1eslesi1 for vegetation typedtype2 benchmarks in 1964 65 eagle lake ranger district lassen national forest CA dominant diversitydiversi ty index

margalefmargaletietlef s Simpsimpsorssimpssorson s vegVPPmegyek no of composition type species species percentage mean SE mean SE MD 6 eleocharis papalustrislustris 52 101.0iolo 020.2 282.8 020.2 19 deschampsia caespitosa 62 404.0 040.4 242.434 020.2 14 juncus balticbaltitusbalticusbalticnsbalbaiticusns 16 232.3 020.2 999.9gg 070.7ot GR 9 carex exserta 71 161.6lgig 020.2 191.9ig 010.1oloi 9 C exserta 78 161.6iglg 020.2 161.6iglg 010.1oloi 19 C exserta 46 323.2 030.3 383.8 030.3 11 festuca idahoensis 26 202.0 030.3 585.8 030.3 SG 5 artemisia tntritrldentata 97 080.8 020.2 101.0iolo 000.0oo 17 A arbuscula 60 363.6 050.5os 262.6 040.4 11 Leptodactylleptodactylonon pungentpungens 34 191.9lg 030.3 505.0so 030.3 TB 7 purshia tridentata 84 080.8 000.0oo 141.4 010.1oi 13 artemisia tridentata 72 323.2 040.4 181.8lbib 020.2 6 carex rossiirossai 28 liii111.1 020.2og 474.7 040.4

highlighted values are the highest and lowest for each index within vegetation types vegetation2vegetation types follow Ratlifratliffratliffetfetet al 1972 MD meadow GR open grassland SG open shrub grass TB timber bunchgrass 199511995 ALPHA DIVERSITY 49

11711.171 P a

azzzn 411911921

c ug V

AL

61 awdAWA

fig 1 meadow diversity benchmarks a eleocharis papalustrislustris b deschampsia caespitosa and c juncus baltitusbalbalticusticus dominated plots eagle lake ranger district lassen national forest CA 50 GREAT BASIN naturalist volume 55

and highest for dmdj and D are specifically dis- the composition total live plant cover was cussed those plots are considered diversity below average but percentages of litter and benchmarks for their vegetation types in and soil cover were well above and below the near the harvey valley allotment averages respectively MEADOW dmD in the meadows was lowest GRASSLAND both djdm and Dsdg were lowest on a plot with just six species and demonstrat- on grassland plots with nine species figs 2aaa ed the effect of lack of richness table 1 the ab2b respectively shorthair sedge was the site was an ephemeral lake meadow fig la main contributor to the composition in the where dominant species covered 333.33 3 of the case of DMD three species each contributed 5 surface among the meadow plots percentage or more and five species each contributed 1 litter cover was lowest and percentage bare or less in the case of DsD only one species soil was highest table 2 other than shorthair sedge contributed as djdm was highest but DsD was lowest on a much as 5 of the composition for the plot meadow with 19 species that finding demon- with low dmD the evenness component was strated the effect of good variety with uneven better litter cover was higher and bare soil abundance the site was a basin meadow pos- cover was lower than for the plot with low Dsdg sibly an ancient lake fig ib there the domi- dmD was highest on a plot with 19 species nant species covered 575.75 7 of the surface only fig 2cac shorthair sedge idaho fescue festuca one species among the others contributed as idahoensis 20 and sandberg bluegrass poa much as 5 to the composition percentages sandbergsandbergiisandberghii 9 were main contributors to the of litter and bare soil were higher and lower composition sixteen species contributed less respectively than averages for the meadow plots than 5 each among the grassland plots this table 3 plot hadbad the highest live plant cover and was D was highest on a plot with 14 species well above average in litter cover and well be- the site was a groundwater fed meadow fig low average in percentage of bare soil lcic evenness in species abundance with mod- idaho fescue dominated the plot with high- erate variety was demonstrated four species est Dsdg fig ad2d four of the other 10 species including the dominant each constituted present each made up more than 10 of the more than 10 of the composition but less composition two species each made up about than 1 of the basal cover only one species 6 while the evenness component of diversi- among the others contributed less than 1 to ty was good and total live plant cover was

taboTABUTAB li 2 beibelPeipeicentapercentagespercentacentasehzehzes of basalb isalisai covecover r for plantpiantplapia nt groups and surfacesui face coarcomrcomponentsonents forhorborfoir vegetationvegetalvegetationlon type benchmarksbenchicbenchmcirks in 1964 65 eagle lake ranger distnctdistrictdistinct lassenI1assenassen N ationalnationalactional forestforest CA

percentageperce niagentage basabasal il cover

plant jgroupslaioups1 surfacsurface e componentScomponents2compoments2mentse2 vegetation typedtype3 g- gl bl sh dp IPlp gr li ro so wo meadow ill 46 06 tat4 62 471 466 61 22 09 07 92 01 802 98 13 26 12 51 903 01 45 open grassland 15 54 05 02 150 76 32 571 171 07 47 04 02 120 60 60 506 05 249 48 53 06 03 76 111 13 601 t 199 t 44 25 03 30 17 102 113 261 26 481 open shrub grass 07 254 162 261 189 284 104 18 05 08 46 82 76 40 380 t 421 36 34 11 45 31 125 86 210 12 537 timbeltimber bunchbunchgrassbuncbgrassgrass 12 06 216 118 233 12 540 05 65 27 10 04 01 45 56 59 29 762 40 46 07 10 06 06 31 21 28 623 94 138 66

1gragr I K grassgrassesK ssesasescs gl1 grasslike herbaccOUSherbaceous plants bl11 broadleafbiobro idleafheibaceoiisbroadleafherbaceousherbaceous plants forbs sh shrubs dp dead attached cover iplp live plant cover gigr glgi 11 sh 2rarci pgraveivelve li littclittalittzilittci ro roekloeklockrock so soil wo laigelarge woody debris 31pesslypesscypes follow kathffetliatlget al 1972and1972 and plot order isis the samesarne itsilsas in table 1 4tatt lesslesslosslevs thantilan 010 1 ofhasalof liaslyas il coverove 199519951 ALPHA DIVERSITY 51

TABLE 3 average diversity indices and percentages of basal cover for plant groups and surface components by vege- tation type eagle lake ranger district lassen national forest CA 1964 65 percentage basal cover diversity indellindexlindex1 plant groups2 surface components3components3 vegetation type D Ds gr gl bl sh dp IPlp gr li ro so wo meadow 229 438 21 36 08 03 64 02 772 01 158 grassland 228 363 33 36 12 08 56 89 93 390 15 357 t open shrub grass 228 278 16 13 06 144 107 180 72 302 02 338 timber bunchgrass 175 239 11 111 1 01 77 40 100 25 602 30 164 39 ad1dD Margmargalefalefss index DsD SimpSimpso sonsrrsrCs index agr 2grr grasses gl grasslike herberherbaceousbaccous plants blbi broadleafbioddlettfheibaceouiiherbaccousherbaceous plants forbs sh shrubs dp dead attached cover ip live plant covelcosercover gigr gl bl sh 3gragrr gravel li litter ro rock so soil wo voodmoodwood above average idaho fescue covered only 3 Ddj was highest on a plot with 13 species of the surface and litter cover was well below fig ab4b nine of them contributed 1 or less but bare soil was well above average of the composition thereby demonstrating SHRUB GRASS both indices were lowest that high evenness is not required when vari-varivarl on an open shrub grass plot where big sage- ety is the main component of diversity consid- brush contributed over 95 of the composi- ered big sagebrush dominated the understory tion fig 3aaa only one other species bottle and covered 434.34 3 of the surface litter cover brush squirreltail sitanionSitanion hystrix made up was well above and bare soil was well below as much as 1 and only five species occurred average for the timber bunchgrass plots on that plot this finding demonstrates the by contrast Dsdg was highest on a plot with effects of both low variety and low evenness just six species fig 4cac ross sedge carex diversity among on the shrub grass plots this rossiirosszirossia contributed most of the composition plot was highest total live second in plant 06og060.60 6 of the surface cover three species con- cover nearly all sagebrush highest gravel in tritributedbuted 15 26 each and two species con- cover and lowest bare soil in this suggests tritributedbuted 3 each thereby demonstrating that soil loss and formation of pavement high variety is not required when evenness black sagebrush dominated the plot with is the main component of diversity considered highest D fig ab3b the 17 species on that dn of percentages of soil and litter cover were near plot 12 of them each contributed less than 3 average for the timber bunchgrass plots of the composition the plot was above aver- age in both litter and soil cover but lowest in beta diversity indices total live plant cover statistical of diversity among the plot with highest Dsdg fig 3cac had just comparisons 11 species and was dominated by false phphloxox communitiescomeom unities and vegetation types were not Leptodactylleptodactylonon pungentpunpungensgens five other speciesspeespec es made nevertheless average values for both combined contributed nearly 62 of the ccom- indices declined from meadow to grassland to position among the shrub grass plots this open shrub grass to timber bunchgrass types plot was well below average in litter cover but table 3 highest in bare soil relative plot ranking high to low diversity TIMBER bunciigrassbunchgrass both indices were depends on the index used and inconsistent lowest fig 4aaa on a timber bunchgrass plot ranking by duD and DDs was expected among with seven species bitterbrushBitterbrush contributed the open shrub grass and timber bunchgrass over 80 of the composition three species types only two plots ranked the same those contributed 2 or more and three species with lowest diversity by both indices rankings contributed less than 1 of the composition by dmD and Dsdg were the same for 3 of the 8 while total live plant cover was above aver- meadow plots and 2 of the 13 open grassland age litter was near average and bare soil was plots well below average there were few species and they were unevenly abundant this plot basal cover relationships was similar in diversity to the shrub grass plot meadow and grassland plots had higher with djdm and Dsdg both low average diversity indices than open shrub grass 52 GREAT BASIN naturalist volume 55

9

WWI M X m7ma

fig 2 open grassland diversity benchmarks a b c carex exserta and d festuca idahoensis dominateddommsited plots eagle lake ranger district lassen national forest CA

or timber bunchgrass plots but lower average sobs w6b6wobbwobg error and Dsdg a shbjhb percentages of live plant cover table 3 total grbogrb2arbgrb error where a gr gl sh gr so and live plant cover was largely a property of shrub wo are explained in table 4 and the bib s are cover because projected crown hits were in- the coefficients corpcorporatedorated into the data base although gravel and bare soil were includ- significant portions of variances in the ed in the model for dmD they did not signifi- cantly with while the diversity indices all 51 plots included were correlate D also in model for dg gravel was not significantly cor- accounted for by variation in percentages of Ds related with Dsdg some basal cover characteristics forty seven individually correlation with dmD was posi- percent of the variation D and 27 of the in dn tive for grasses r 471471.471 but negative for grass- by variation in DsD were explained the best like plants r 014014.014 shrubs r 320320.320 and models table 4 wood r 348348.348 correlation of Dsdg with dmD a arbgrbgrbi albaglbaglb2 shb3 grb4grbo shrubs was negative r 507507.507 also 199519951 ALPHA DIVERSITY 53

k

d

fig 2 continued

discussion stress richness and those that stress evenness tend to be poorly correlated Mamagurranmagureangurran 1988 alpha diversity beta diversity many diversity indices are available to tthe data used in this study represent single land manager although a particular diversidiversity time samples and were not designed to esti- index be preferred is generally to may it best mate beta diversity testing for differences in use one that stresses species richness and one diversity using such data was not considered that stresses evenness dominance such as dmD reliable west and reese 1991 and Dsdg respectively doing so allows the man- nevertheless diversity indices for different ager to consider both components of diversity but closely similar plots or communities when the richness component of diversity may in- computed by the same methods should be crease at the expense of the evenness compo- nearly equal with time or different treatment nent or vice versa also those indices that wide divergence of the indices may occur 54 GREAT BASIN naturalist volume 55

f orfi

dflor fflorm 7

fig 3 open shrub glassgrass diversity benchmarks a artemisia dentatatntritrltndentatatridentatatridentate b A arbuscula and c Leptodactylleptodactylonon pungentpungens dominateddommAted plots eagle lake ranger district lassen national forest CA 19951 ALPHA DIVERSITY 55

d

do

illallalf

fig 4 timber bunchgrass diversity benchmarks a purshia tritrldentata b artemisia trltridentata and c carex rossiirossai dominated plots eagleeagie laielake ranger district lassen national forest CA 56 GREAT BASIN naturalist volume 55

TABLE 4 best model multiple linear regression coefficients tests of significance T and probabilities of significance P for margalef s and simpsinssimpsorssimpsonsSimpsorssonssots diversity indices eagle lake ranger district lassen national forest CA 1964 65 diversity index

margalef s simpsinssimpsonsSimpsons variable symbol coeffloefflboeffcoefflI1 T P coeffboeff T FP constant a 2696 8436 000 3584 12372 000 grasses gr 0238 3522 001 grasslike plants gl 0161 3285 002 shrubs sh 0036 2722 009 0102 4194 000 gravel gr 0026 1505 139 0033 0970 337 bare soil so 0007 1197 238 wood wo 0128 3173 003 lregressionlRegression coefficient

permanent plots represent a resource for dominant and the others contributed little assessing beta diversity responses to land man- plant cover the influence of evenness in abun- agement practices although sampling a site to dance on Dsdg was clearly evident Dsdg tended to include within and between season variation be highest when species were more or less is desirable doing so is seldom possible given evenly abundant frequently that occurred with time and monetary constraints As an alternative relatively few species few species with one one might restrict sampling to times when contributing a high percentage of the compo- selected species indicators are in specific phe sition produced low values of both indices nologicneologic stages eg budding or flowering situations with many species all contributing basal cover equally to the composition were not encoun- tered but such situations should give high val- because of the usual dominance of a single ues of arnD and dg species and because species tends to drn Ds that higher diversity did not necessarily mean occupy high proportions of an area reductions greater plant cover or greater forage cover or in diversity indices with increases in shrub less bare soil rela- cover may be expected more litter or while some tionshipsships both diversity indices may be related posi- tion between diversity and basal cover tively or negatively to characteristics of basal values were significant coefficients of deter- cover or to soil properties nevertheless D minationmi were too low to allow either of the was related to a greater number of characteris- best models to be used to predict diversity tics than DsD suggesting that dmD may be the neither index should be relied on apart more desirable index for comparing plant com- from other information for evaluating range- munitiesmuni ties of different sites or plant communities land health nevertheless plants capture the present through time on a given site sunsuns s energy and pass it as food for other orga- nisms and a high degree of plant diversity conclusions may equate with high diversity in other parts of the biotic community for similar communities we can expect plant species diversity to be highest in the literature CITED meadow and lowest in the pine bunchgrass types high and low values of margalef s and DEL MORAL R AND D M WOOD 1988 the high elevation simpsoisimpson s diversity indices are available for flora of mount st helens washington Madromadronofioflo 35 benchmark plots of different vegetation types in 309 319 dyksterhuis E J 1949 condition and management of and near the harvey valley allotment diversity range land based on quantitative ecology journal of indices for and averages among 51 plots are range management 2 104 115 available by vegetation types HORMAY A L 1970 principles of rest rotation grazing the influence of species richness on dmD and multiple use land management training text was clearly evident to be highest 42200 USDA forest service washington DC dmD tended 26 appp with the greatest numbers of species frequently LUDWIG J A AND J F REYNOLDS 1988 statistical ecology that occurred when one species was clearly john wiley & sons new york NY 199519951 ALPHA DIVERSITY 57

MAGURRAN A E 1988 ecological diversity and its mea- WEST N E 1993 diversitybiodiversityBio of rangelands journal of surementsu princeton university press princeton NJ range management 46 2 13 ODUM E P 1959 fundamentals of ecology W B saunders WEST N E AND G A REESE 1991 comparison of some co philadelphia PA methods for collecting and analyzing data on above- RATLIFF R D J N REPPERT AND R J MCCONNEN 1972 ground net production and diversity of herbaceous rest rotation grazing at harvey valley range vegetation in a northern utah subalpine context health cattle gains costs USDA forest service vegetatiovegetationvegetatio9696 145 163 pacific southwest experiment station research WHITTAKER R H 1972 evolution and measurement of paper PSW 77 24 appp species diversity taxon 21 213 251 SAS INSTITUTE INC 1982 SAS user s guide statistics 1982 WILKINSON L 1989 SYSTAT the system for statistics edition SAS institute inc carsgarscarygary NC SYSTAT inc evanston IL SOCIETY OF AMERICAN FORESTERS 1992 biological diver- sity in forest ecosystems a position of the society of received 26 july 1993 american foresters journal of forestry 90 424342 43 accepted 26 may 1994 STEIN J AND L URDANG EDS 1966 the random house dictionary of the english language random house new york NY great basin naturalist ssi551 0 1995 appp 58 65 EFFECTS OF SALINITY ON establishment OF POPULUS FREMONTHFREMONTII cottonwood AND TAMARIX ramosissima SALTCEDAR IN southwestern UNITED STATES

patrick B Shafroth 1 jonathan A Friedmafriedmanlfnedman1friedmannnl and lee S Ischischingerlischmger1ingerlingeri

ABSTRACT the exotic shrub tamarix ramosissima saltcedarsaltcedar has replaced the native populusfremontiipopulus fremonfremontefremontntn cottonwood along many streams in southwestern united states we used a controlled outdoor experiment to examine the influence of river salinity on germination and first year survival of P fremonfremontiifremontnaremonfremontiatiitn var wislizeniiwislizenn rio grande cottonwood and T ramosissima on freshly deposited alluvial bars we grew both species from seed in plantersplanpianteisters of sand subjected to a declin- ing water table and solutions containing 0 1 3 and 5 times the concentrations of major ions in the rio grande at san marcial NM 12121.21 2 10010.010loo 0 25725.725 7 and 37437.437 4 leqmeq 1 1 0 11 0970.970 97 2372.372 37 and 3453.453 45 ds m 1 germination of P fremonatemonfremontufrearemonfremontiifremontiapremontutiitil declined by 35 with increasing salinity P 008 germination of T ramosissimaramostssima was not affected there were no significant effects of salinity on mortality 01or above and belowgroundbelowground growth of either species in laboratory tests the same salini ties had no effect on P fremonaremonfremontiifremontufrefremontiapremontutii germination P fremonatemonstemonfremontitfremontiifrefremontiamontittiitil germination is more sensitive to salinity outdoors than in cov- ered petri dishes probably because water scarcity resulting from evaporation intensifies the low soil water potentials associated with high salinity river salinity appears to play only a minor role in determining relative numbers of P fre montu and T ramosissima seedlings on freshly deposited sandbars however over many years salt becomes concentrat- ed on floodfloodplainsfloodplamsplamsplains as a result of evaporation and salt extrusion from saltcedarsaltcedar leaves T ramosissima is known to be more tolerant of the resulting extreme salinisalinitiesties than FP fremonfremontufremontiifrefremontiabrepremontutiitil therefore increases in river salinitiessalinities could indirectly con- tribute to decline of P fremonstemonatemonfremontufremontiifrefthfrhfremontiamontutiitil forests by exacerbating salt accumulation on floodfloodplainsplains

key words exotic species tarnarixtamanetamanxTarnarix ramosissima populus fremonbremonfremontnfremontiifremontiatiitiltn river salinity seedling establishment rio grande riparian vegetation basquebosque del apache national wildlife refuge

in the last century the exotic shrub saltcedarsaltcedar channelchannelizingizing southwestern watercourseswatercourses tamarix ramosissima Ledebour has spread reductions in the magnitude of high flows and throughout southwestern united states where associated reductions in channel movements it now dominates many riparian ecosystems decreased the formation of bare moist alluvial bowser 1958 robinson 1965 in many areas bars which provide ideal P fremonfremontiifremontiatii seedling T ramosissima has replaced stands dominated habitat ohmart et al 1977 stromberg et al by the native fremont cottonwood populus 1991 smaller peak flows have also reduced fremonfremontiifremontiatii wats campbell and dickdickpeddiepeddie leaching of salts from floodfloodplainplain soils busch 1964 ohmart et al 1977 decreasing the habi- and smith in press perhaps favoring the salt tat of neotropical migrant birds anderson et al tolerant tamarix everitt 1980 brotherson 1977 cohan et al 1978 and altering fluvial and winkel 1986 jackson et al 1990 flow processes graf 1978 blackburn et al 1982 regulations that have altered the historical understanding the factors controlling estab- timing of peak flows may have inhibited P fre lishmentlishment of T ramosissima and P fremonfremontiifremontiatii can montil regeneration because of its short period aid in managing these species of seed dispersal and viability in early summer successful invasion by tamarix in the south- horton 1977 everitt 1980 but they have west has been attributed to many factors much enhanced tamarix regeneration because of its of the early spread probably resulted from the abundant seed production throughout the coincidental timing of clearing of P fremonfremontiifremontiatii growing season merkel and hopkins 1957 stands by early settlers and the availability of tomanek and ziegler 1962 warren and turner tamarix seed campbell and dickdickpeddiepeddie 1975 horton 1977 finally successful inva- 1964 harris 1966 horton and campbell sion of T ramosissima has been attributed to 1974 ohmart et al 1977 subsequent spread its superior ability to resprout following fire resulted largely from effects of damming and busch and smith 1993

rational biological survey midcontinentmidcontinedtMidcontinent ecological science center fort collinicollins CO 80525340080525 3400

58 199519951 SALINITY EFFECTS ON POPULUS AND TAMARIX 59

we conducted experiments to examine the filled to 92 cm with washed coarse sand influence of river salinity on germination sur- approximately 6 gravel 2000 lumjom 78 vival and growth of populuspopulusfremontiifremonbremonfremontiifremontiatiitil var wis sand 300 2000 tkmtamim 16 fine sand lizeniilizenie rio grande cottonwood and T ramo 75 300m300tkml and 1 silt and clay sissimamissima on freshly deposited alluvial bars the four salinity treatments were each replicat- principal habitat for seedling establishment of ed in three tanks 12 tanks total each tank both species field observations have suggest- contained three planters of P fremonfremontiifremontiatii var ed that P fremonfremontiifremontiatii is more negatively affected wislizenii and three of T ramosissima thus by high salt concentrations than T ramosissi the experimental unit for each species was a ma brotherson and winkel 1986 anderson group of three planters within a tank to avoid 1989 laboratory studies have confirmed this pseudoreplicationpseudo replication responses were measured difference by exposing seedlings and cuttings as the mean value of the three planters the of these species to varying concentrations of results for the two species were analyzed as nacl and cac12caccag jackson et al 1990 siegel separate completely randomized experiments and brock 1990 two factors potentially con- with four treatments and three replicates per found the relationship of results of laboratory treatment studies to field conditions first the mix of the tanks were filled with water from the salts found in riparian ecosystems typically cache la poudre river a snowbeltsnowsnowmeltmelt stream low includes many constituents other than na ca in dissolved solids and solutions containing and cl in many plants salinity effects result multiples 0 1 3 and 5 times of the mean con- from toxicity of specific ions as opposed to centrationcentration of all major ions in the middle rio osmotic stress greenway and munns 1980 grande were made these four solutions con- second moisture availability is lower and stitute treatments ox ix 3xax and 5xax mean ion more variable in the field than in these labora- concentrations were derived from eight mea- tory studies this factor is important because surementssurements from the conveyance channel at low soil water potential caused by high salinity san marcial NM between october 1989 and is exacerbated by low soil moisture content september 1991 US geological survey 1991 we addressed these concerns by exposing T 1992 the following salts were added to make ramosissima and P fremonfremontiifremontiatii seedlings to four treatment ix 3099309.9 mg I1 1 cas042h20cas042h2o 3024302.4 different concentrations of a mix of salts mg I1 1 nahc03 1220122.0 mg H1 1 mgclghaomgc1261120 designed to mimic ion concentrations in the 70170.1 mg I1 1 nacl 13913.9 mg I1 1 KSOk2s04 because rio grande the experiment was conducted the coarse sand substrate was low in nutrients outdoors in planters subjected to a controlled cf segelquist et al 1993 15 mg I1 1 of fisonsbisons water table drawdown experimental condi- technigroTechnigro fertilizer 16 N 17 P 17 K was tions were designed to simulate alluvial bars added to every tank along the rio grande in central new mexico at the time of planting and for I1 wk there- where once extensive P fremonfremontiifremontiapfremontiitii forests have after the water level was 10 cm below the soil largely been replaced by T ramosissima thick- surface A 35 cm week 1 drawdown rate was ets campbell and dickdickpeddiepeddie 1964 our applied for the remainder of the growing sea- outdoor experiments were supplemented by son 17 june to late september water table studies of germination under similar salinity drawdownsdrawdowns are associated with summer treatments in the laboratory declines in discharge along western streams the 35 cm week 1 drawdown rate was select- METHODS ed because a previous study segelquist et al 1993 indicated that it is within the optimal seedling establishment experiments were range for establishment and growth of plains conducted outdoors in 1993 near fort collins cottonwood populus deltoidesdeltoides sspasp monilifera CO at latitude 40 35 north longitude 105 5 flowering papaniclespaniclednicles of T ramosissima were west and elevation 1524 m twelve 122 X 92 collected on 17 may at the bosque del apache cm diameter X depth epoxy lined steel tanks national wildlife refuge latitude 334633 46 contained six 30 X 100 cm planters made of north longitude 10654106log 54 west elevation 1375 PVC pipe holes 1261.26 cm in diameter were m the paparticlesnicles were air dried for 48 h to drilled into the lower 10 cm of each planter to enhance opening of seed capsules collected allow water exchange and the planters were material was sifted through a series of soil 60 GREAT BASIN naturalist volume 55 screens until clean samples of seeds were ob- the substrate column and seedlings in the tained catkinsbatkins of P fremonfremontiifremontiatii were collected basin we gently separated seedlings from the at the bosque del apache on I1 june the cat sand and water and measured total length of kins were air dried for 72 h to enhance open- every harvested seedling mean root lengths ing of seed capsules capsules were placed were determined by subtracting the mean between soil screens and seeds were separat- shoot length for a planter from the mean total ed from the cotton and capsules using forced length in that planter roots and shoots were air seeds of both species were sealed in plas- separated for both species and P fremonfremontiifremontiatii tic containers and refrigerated at VC zasada leaves were stripped from the stems roots shoots goc and densmore 1977 on 10 june 100 P fre and leaves were dried at 60c60 C for 72 h weighed montiimontai seeds were planted in each of three and one analysis of variance SAS institute planters per tank and 200 T ramosissima way 1990 was used to assess the significance seeds were planted in each of the other three inc of differences within the two planters treatment species for five variables percent of planted seeds alive electrical conductivity EQEC and tempera- at the end of the experiment end of season ture were measured using a yellow springs survival shoot length root length per plant co inc model 33 S C T meter instrument SCT aboveground biomass and per plant root bio- and was measured using a corning 105 ph mass for all variables the mean value of the hand held meter in with a ph conjunction three planters in a tank was the unit of analysis ATC temperature probe and a coming coming arcsine transformation was applied to end general electrode EC the purpose combination of season survival values to meet the equal vari- weekly tank beginning was measured in every beginingbegining ance assumption snedecor and cochran 1980 12 june 17 measuring dates whenever EC data from the colorado climate center was measured a representative water temper- were used to determine the difference be- ature for that day was determined by averag- tween precipitation and open pan evaporation ing the temperature values from five randomly adjusted with pan coefficient 0730.73 for the selected tanks all EC measurements were period I1 june 30 september 1993 in fort corrected for temperature and reported at collins evaporation at fort collins exceeded 25c25 C fourteen weekly measurements of ph precipitation by 26226.2 cm during this period the were made beginning 30 june on 16 june 14 same calculation was made for the bosque del july 18 august and 17 september water sam- apache using data from the western regional ples from one randomly selected tank per climate center for the years 1975 through treatment were analyzed to determine con- 1990 precipitation data are from the bosque of mg na K coogoo centcentrationsrations ofcaca 0 6000 hloihcoihc03J cl del apache national wildlife refuge and s04 and n03 ca mg na and K were deter- open pan evaporation data are from socorro mined by inductively coupled plasma emis- NM latitude 34534 5 north longitude 10653106log 53 sion spectroscopy JCPICP EPA method 2000200.0 west elevation 1399 in pan coefficient 0730.73 united states environmental protection growing season evaporation at the bosque del agency 1983 c0303 and hc03 were deter- apache exceeded precipitation by an average mined by titration EPA method 3101310.1 united of 40640.6 cm n 16 maximum 510sio51.0 cm and states environmental protection agency minimum 32332.3 cm during these 16 years 1983 cl s04 and n03 were determined by we performed laboratory germination exper- ion chromatography concentrations are iments in january 1994 five 25 seed repli- reported in leqmeq 1 1 to facilitate comparison of cates of five salinity treatments were com- our solutions to solutions in other studies and pletely randomized for both T ramosissima and because leqmeq 1 1 can be related easily to elec- P fremonfremontiifremontiatii seeds were sowedbowed in 75 cm petri trical conductivity which is commonly report- dishes containing a whatman 3 filter and 7 ed in the context of salinity studies ml of a treatment solution petri dishes were on 29 september 1993 day 112 we mea- placed in a percival model 1351 35 biological sured the shoot length of every living seedling incubator after sealing the dish tops with para we harvested all live seedlings in early october film temperature in the incubator was 20c20 C to harvest we lifted a planter and laid it hori- throughout the experiment and petri dishes zonzontallytally in a water filled basin the planter were exposed to 16 h of light and 8 h of dark- was then slowly lifted upside down leaving ness each day four of the treatment solutions 199511995 SALINITY EFFECTS ON POPULUS AND TAMARIX 61 were the same as those used in the establish- 217c standard error 080.8os n 17 concen- ment experiment 0 1 3 and 5 times the con- trationstrations of measured chemical constituents in centrationcentration of the rio grande at san marcial different treatments did not increase propor- NM the fifth solution contained 7 times the tiotionallynally to the quantities of salt originally concentration of the rio grande Germigerminantsnants in added indicating that salts especially cac03 every petri dish were counted after seven days precipitated at higher concentrations table 1 A seed was considered germinated if it exhibited nevertheless concentrations increased across expanded cotyledons and an elongated radicle treatments with total concentrations ranging the arcsine transformation was applied to per- from 070.7 leqmeq I1 1 oli0110110.11 ds m 1 in treatment ox cent germination values to meet the equal to 37437.4 leqmeq I1 1 3453453.45 ds m 1 in treatment 5xax variance assumption and one way analysis of table 1 variance was performed on the transformed for P fremonfremontiifremontiatii there was a significant values SAS institute inc 1990 when germ- treatment effect P 003003.003 on end of season ination equaled 100 the proportion was survival but not on any of the four measured counted as n 025n where n the num- growth variables table 2 end of season sur- ber of seeds planted snedecor and cochran vival was negatively associated with increasing 1980 salinity survival was greatest in treatment ox and lowest in treatment 5xax because the end RESULTS of season survival variable combines germina- tion and mortality we analyzed the arcsine EC and ph in the tanks varied little within transformed number of seedlings 7 d after plant- treatments over the course of the experiment ing germination and the arcsine transformed table 1 mean temperature in the tanks was difference between germination and end of

TABLE 1 chemical analysis of tank water for four treatments in the outdoor establishment experiment in fort collins CO for ionlon concentrations n 4 minimum and maximum values are presented in parentheses below treatment means for electrical conductivity n 51 and ph n 42 means t I1 standard error are presented treatmenttreatmentament factor ox ix 3xax 5xax ca amolmmol 111 1 036 182 402 454 0200200.200 20 0520.520 52 1711711.711 71 2002.002 00 3493493.493 49 4834.834 83 3023023.023 02 7027.027 02

mg amolmmol 111 1 oiioli011oil0.11 060 165 262 0080080.080 08 0160.160olg16 0460750460750460.750 46 0 75 1471971471971471.971 471 97 2282972282972282.972 28 2 97 na amolmmol 11 017 485 1387 2224 0090280090.280 09 0 28 441511 11911549 193319 33 246524.6524 65 K amolmmol 11 008 026 051 079 0060060.060 06 0090.090 09 02002000.2020 0340.340 34 0440 44 0550.550 55 07207200.7272 090ogo0 90 hc03 amolmmol 11I1 1 104 392 834 960 0621440621440621.440 621 44 3243 24 4444 44 7299967299967299.967 29 9 96 587157458715.745 87 15 74 cl amolmmol 11 010 247 710 1212 0070140070140070.140 07 0 14 1882821882821882.821 88 2 82 6967316967.316 96 7 31 10881321108813.2110 88 13 21

804 amolmmolmm01 11 004 166 506 773 0040040.040 04 0050.050 05 1321321.321 32 1861.861 86 4764 76 5325 32 7137137.137 13 8338 33

n03 amolmmol 111 1 003 003 005 006 000200020.0020 002 0080.080 08 000600.006006 00900.0909 ooi0010 01 0080.080 08 0020 02 0150 15 total cationscanions leqmeq 11 12 100 257 374 07160716071.60 7 1 6 92108921089210.89 2 10 8 23823 8 26726.726 7 34541534541.534 5 41 5 EC ds m 1 1091.09log1 09 0030.030 03 0970.97097011011oiioli0.11 2372.372 37 0230.230 23 34533.4545 0390.390 39 ph 7547.547 54 0030.030 03 8108.108 10 0020 02 8298.298998 29 0020 02 8058 05 0030 03 62 GREAT BASIN naturalist volume 55

TABLE 2 survival and growth of populus fremonfremontfremontiifremonttzfremontiatiitiltz and tamarixramany ramosissima seedlings exposed to fourbour different salini- ty treatments for one growing season outdoors in fort collins CO high and low replicate means are given in parenthe- ses below the treatment means n 3 treatment effects were analyzed by completely randomized one way ANOVA survival ANOVA was performed on arcsine transformed data treatmenttreatment species variable ox lx1xdx 3xax 5xax F P cottonwood survival 570 493 466 290 114 003 of plantedofplanted seed 50050 0 63063 0 45745 7 54054 0 41041 0 51051.051sio 0 2072072020.77 35035.035 0

shoot height mm 339 363 396 383 26 13 32832832.832 8 34534.534 5 34534534.534 5 38538.538 5 36536 5 43943 9 34734 7 40840 8 root length mm 2392 2809 2869 2474 21 17 2271227.1227 1 2584258 4 25782578257.8257 8 3093309.3309 3 2536253 6 3117311 7 906gog2063206 3 2746274.6274 6

per plant shoot 141 146 214 198 28 11 biomass mg 13713713.713 7 14414.414 4 112 166 1891891818.99 25825.825 8 14314314.314 3 25925.925 9

per plant root 268 196 318 312 111 1 41 biomass mg 21221221.221 2 35535.535 5 16416416.416 4 21321.321 3 21621621.621 6 43243.243 2 17217217.217 2 42942.942 9 saltcedarSaltcedar survival 423 378 373 295 16 26 of planted seed 29551629551.629 5 51 6 33833833.833 8 42042.042 0 31831831.831 8 40840.840 8 22822822.822 8 35235.235 2 shoot height mm 181 177 182 183 004 99 17318817318817318.817 3 18 8 15519815519.815 5 19 8 15615.615 6 22222.222 2 18218318218318.218318 2 18 3 root length mm 1744 1736 1790 1620 015 92 166418491664184.9166 4 184 9 154819291548192.9154 8 192 9 12811281128.1128 1 2436243.6243 6 147216951472169.5147 2 169 5 per plant shoot 55 55 63 62 022 88 biomass mg 48624 8 6 2 41644164416.44 1 6 4 43964396439.64 3 9 6 58645864586.45 8 6 4 per plant root 77 73 99 95 074 56 biomass mg 71897189718.97 1 8 9 559255925.5925 5 9 2 701477 014 7 761217612.17 6 12 1 season survival mortality there was a signif- ds m 1 is consistent with results of earlier icant treatment effect on germination P studies jackson et al 1990 found that P fre 008008.008 but not on mortality P 4545.45 indicat- montiimontai germinated in the laboratory at salini ing that the effect on end of season survival ties of 0 27 and 106 leqmeq 1 1 using a mixture was predominantly due to lower germination of nacl and cacl2cac but not at 319 leqmeq I1 1 or at higher salt concentrations for T ramosissima above siegel and brock 1990 observed high- there were no significant treatment effects er percent germination of P fremonfremontiifremontiatii in the table 2 laboratory in nacl solutions of 0 25 and 50 although P fremonfremontiifremontiatii germination in out- leqmeq 1 1 than at 100 leqmeq 1 1 and above there- door tanks was significantly decreased at high fore P fremonfremontiifremontiatii is no more sensitive to the salinity laboratory germination was not simi- mix of salts present in the rio grande than to larly affected even at seven times the salinity nacl and cac12caclg solutions of equal strength of the rio grande total concentration 48448.4 tests at higher salinitiessalinities with the same ionic leqmeq 1 1 4564564.56 ds m 1 table 3 there was a sig- ratios were not possible with our rio grande nificantnificant positive effect of increasing salinity on mix because of low solubilities of some of the T ramosissima germination P 0303.03 table 3 constituent salts the decrease in T ramosissi ma germination at low salinity in the laborato- discussion ry table 3 is consistent with the finding by jackson et al 1990 that germination increas- the absence of a negative effect of salinity es between 0 and 106 leqmeq 1 1 on P fremonfremontiifremontiatii germination in the laboratory at our results indicate that a given water salin- concentrations as high as 48448.4 leqmeq 1 1 4564564.56 ity may negatively affect germination of P 199519951 SALINITY EFFECTS ON POPULUS AND TAMARIX 63

TABLE 3 percent germination of populus fremonfremontiifremontiatii and tamarix ramosissima seedlings exposed to five salinity treat- ments in covered petri dishes high and low replicate values are given below the treatment mean n 5 treatment effects were analyzed by completely randomized one way ANOVA using arcsine transformed data treatment species ox ix 3xax 5xax 7xax F P cottonwood 904 960 960 928 960 12 35 8001000800100.080 0 loo100 0 9201000920100.092 0 loo100 0 9201000920100.092 0 100 0 84084084.084 0 96096.096 0 9201000920100.092 0 loo100 0

saitsaltcedarsalteedarSaltsaiteSalteedarcedar 696 688 784 848 840 33 03 600880 56056080056056.0 80080.0 68068092068068.0 92092.0 76076092076076.0 92092.0 76076092092092.0 fremonfremontiifremontiatii seeds under ambient conditions but del apache john taylor bosque del apache not under laboratory conditions this may have national wildlife refuge personal communica- resulted from an interaction between the tion and soil salinity levels as high as 60000 effects of salinity and soil moisture content or mg 1 1 occur on floodfloodplainplain sites along the from vapor pressure deficit differences in lower colorado river jackson et al 1990 outdoor planters but not laboratory petri dish- soil EC above 202.090go ds m 1 can reduce the es evaporation of water may have resulted in growth of P fremonfremontiifremontiatii pole plantings anderson lower soil moisture and higher salt concentra- 1989 T ramosissima has been shown to be tion at the soil surface these factors would less susceptible than P fremonfremontiifremontiatii to many of both tend to reduce soil water potential the negative effects of higher salinitiessalini ties thereby increasing plant water stress because brotherson and winkle 1986 jackson et al the difference between evaporation and pre- 1990 tamarix species avoid harmful effects of cipitation is somewhat greater at the bosque del salts through extrusion from leaves and cellu- apache than in fort collins the effect of salin- lar compartmentationcompartmentation berry 1970 kleinkopf ity might be stronger at the bosque especially and wallace 1974 waisel 1991 in dry years finally greater vapor pressure our results could be applied to efforts to deficits in the field relative to the laboratory revegetate riparian areas from seed riparian may have exacerbated plant water stress revegetationrevegetation in the southwest has largely con- salinity appears to be a relatively minor fac- sisted of planting poles or potted shoot cut- tor regulating numbers of P fremonfremontiifremontiatii and T tings although these approaches have been ramosissima seedlings on freshly deposited successful in some areas anderson et al 1990 sandbars along the rio grande the only signif- they can cost up to 10000 per hectare ohmart icant effects of increasing salinity were a small et al 1988 furthermore they require the decrease in P fremonfremontiifremontiatii germination in out- destruction of parts of existing trees and often door planters and a small increase in T ramo entire trees or stands finally these approach- sissimamissima germination in the laboratory there es may require importing cuttings or poles were no significant effects on survival after adapted to different site conditions one alter- germination or above or belowbelowgroundground growth native is regeneration of native cottoncottonwoodswoods for either species even at water salinitiessalinities sev- and willows using natural seedseedfallweedfallfall friedman eral times that of the rio grande the presence 1993 john taylor personal communication of abundant seedlings of P fremonfremontiifremontiatii and T this approach generally involves clearing and ramosissima on sandbars along the rio grande irrigating an area so that seeds from nearby in most years is consistent with our results trees can colonize it our results suggest that although salinity may play only a minor role water as saline as 37437.4 leqmeq I1 1 EC 3453.45 ds in the colonization of newly deposited alluvial m 1 can be used to grow P fremonfremontiifremontiatii from bars by T ramosissima and P fremonfremontiifremontiatii this seed on sand tables 1 2 however care must factor can become more important over time be taken to prevent longtermlong term salt accumula- over many years salt becomes concentrated tion through evaporation eg through period- on some floodfloodplainsplains as a result of evaporation ic flooding to flush salts and to avoid sites and salt extrusion from T ramosissima leaves with preexisting high salinity use of water EC readings as high as 10010.0loo ds m 1 have been with low salinity can help prevent negative reported in floodfloodplainplain sediment at the bosque effects on P fremonfremontiifremontiatii and may decrease the 64 GREAT BASIN naturalist volume 55 germination rate of T ramosissima table 3 tion of the united states and comments concerning however in a restoration effort along the the plants influence upon the indigenous vegetation pages 12 16 in symposium on phreatophytes poudre T m ameri- cache la river ramosissima became can geophysical union sacramento CA established in large numbers along with P del brotherson J D AND V WINKEL 1986 habitat rela- toidesboides in spite of use of water of low salinity tiontionshipsships of saltcedarsaltcedar tamarix ramosissima in cen- douglas gladwin national biological survey tral utah great basin naturalist 46 535 541 personal communication therefore low salin- BUSCH D E AND S D SMITH 1993 effects of fire on water and salinity relations of riparian woody taxa ity will not prevent establishment of T ramo oecologiaoecologia9494 186 194 sissimamissima from seed when moisture a bare sedi- in press decline persistence and competition of ment and a seed source are present woody taxa in riparian ecosystems of the southwest- ern USU S submitted to ecological monographs CAMPBELL C J AND W A dickpeddieDICK PEDDIE 1964 compari- acknowledgments son of phreatophyte communities on the rio grande in new mexico ecology 4549245 492 502 G T auble D E busch and an anonymous COHAN D R B W ANDERSON AND R D OHMART 1978 reviewer provided constructive comments on avian population responses to salt cedar along the pages the manuscript we thank E R auble G T lower colorado river 371382371 382 in R R johnson and J F mccormick technical coordinators auble J back E D eggleston M jordan strategies for protection and management of flood and M L scott for invaluable assistance with plain wetlands and other riparian ecosystems pro- the experiments D smeltzer B upton and ceceedings of the symposium USDA forest service the colorado division of wildlife generously general technical report WO 12 washington DC EVERITT B L 1980 ecology of saltcedarsaltcedar a plea for provided access to the bellvue watson fish research environmental geology 3 77 84 rearing unit where the outdoor experiment FRIEDMAN J M 1993 vegetation establishment and was conducted T kern and PE soltanpourSoltanpour pro- channel narrowing along a great plains stream fol- vided useful advice regarding the salinity lowing a catastrophic flood unpublished doctoral treatments of ions solutions dissertation university of colorado boulder concentrations in GRAF W L 1978 fluvial adjustments to the spread of were measured by the soil plant and water tamarisk in the colorado plateau region geological testing laboratory at colorado state university society ofamericaofamerica bulletin 89 1491 1501 fort collins CO GREENWAY H AND R MUNNS 1980 mechanisms of salt tolerance in nonhalophytes annual review of plant physiology 31 149 190 CITED literature HARRIS D R 1966 recent plant invasions in the and and semisemiaridarid southwest of the united states annals of ANDERSON B W 1989 research as an integral part of the association ofamerican geographers 56 408 422 revegetation projects pages 413 419 in D L abell HORTON J S 1977 the development and perpetuation of technical coordinator proceedings of the california the permanent tamarisk type in the phreatophyte riparian systems conference protection manage- zone of the southwest pages 124 127 in R R ment and restoration for the 1990s USDA forest johnson and D A jones technical coordinators service general technical report PSW 110 importance preservation and management of ripar- berkeley CA ianlan habitat a symposium USDA forest service ANDERSON B W A HIGGINS AND R D OHMART 1977 general technical report RM 43 fort collascollmscollins CO avian use of saltcedarsaltcedar communities in the lower colo- HORTON J S AND C J CAMPBELL 1974 management of rado river valley pages 128 136 in R R johnson phreatophyte and riparian vegetation for maximum and D A jones technical coordinators importance multiple use values USDA forest service research preservation and management of riparian habitat paper RM 117 23 appp a symposium USDA forest service general JACKSON J J T BALL AND M R ROSE 1990 assessment technical report RM 43 fort collins CO of the salinity tolerance of eight sonoran desert ANDERSON B W E R MILLER AND J P washington riparian trees and shrubs final report USU S bureau of 1990 revegetation on the kern river preserve reclamation contract no 9 CP 30 07170 biological 1986 1989 research report prepared for the sciences center desert research institute univer- nature conservancy and california department of sity of nevada system reno 102 appp fish and camegame revegetation and wildlife manage- KLEINKOPF G E AND A WALLACE 1974 physiological ment center inc blythe CA 19 appp basis for salt tolerance in tamarix ramosissima plant BERRY W L 1970 characteristics of salts secreted by science letters 3 157 163 tamariztamarix aphyllyaphylla american journal of botany 57 MERKEL D L AND H H HOPKINS 1957 life history of 1226 1230 salt cedar tamarix fallicagallica L transactions of the BLACKBURN W H R W KNIGHT AND J L SCHUSTER kansas academy of science 60 360 369 1982 saltcedarSalt cedar influence on sedimentation in the OHMART R D B W ANDERSON AND W C HUNTER brazos river journal of soil and water conserva- 1988 the ecology of the lower colorado river from tion 37 298 301 davis dam to the mexico united states internat- BOWSER C W 1958 introduction and spread of the un- ional boundary a community profile USU S fish and desirable tamatamarisksrisks in the pacific southwestern seesec wildlife service biological report 85719857857.1919 296 appp 199519951 SALINITY EFFECTS ON POPULUS AND TAMARIX 65

OHMART R D W 0 DEASON AND C BURKE 1977 A biological sciences fort hays kansas state college riparian case history the colorado river pages hays 128 appp 35 47 in R R johnson and D A jones technical UNITED STATES environmental protection AGENCY coordinators importance preservation and man- 1983 methods for chemical analysis of water and agement of riparian habitat a symposium USDA wastes publication identification EPA 600 4790204 79 020 forest service general technical report RM 43 environmental monitoring and support laboratory fort collins CO office of research and development cincinnati ROBINSON T W 1965 introduction spread and areal OH extent of saltcedarsaltcedar Tatamarixmanx in the western states UNITED STATES geological SURVEY 1991 water re- united states geological survey professional paper sources data new mexico water year 1990 USU S geo- 491 A 13 appp logical survey water data report NM 90 SAS INSTITUTE INC 1990 SASSTAT user s guide ver- 1992 water resources data new mexico water sion 6 ath4th edition SAS institute inc carygary NC year 1991 USU S geological survey water data leportreport segelquist C A M L SCOTT AND G T AUBLE 1993 NM 91 establishment of populus deltoidesdeltoides under simulated WAISEL Y 1991 the glands of tanwrixtamanatamanx aphyllyaphylla a system for alluvial groundwater declines american midland salt secretion or for carbon concentration physiologiaphysiologic naturalist 130 274 285 Planpianplantarumplantariumplantarum83tarum 83 506 510 SIEGEL R S AND J H BROCK 1990 germination require- WARREN D K AND R M TURNER 1975 saltcedarSaltcedar tama- ments of key southwestern woody riparian species axnxrix chinenchinensissis seed production seedling establishment desert plants 10 3 8 34 and response to inundation journal of the arizona SNEDECOR G W AND W G COCHRAN 1980 statistical academy of science 10 135 144 methods iowa state university press ames 507 appp ZASADA J C AND R DENSMORE 1977 changes in sali- STROMBERGSTROMBEKG J C D T PATTEN AND B D RICHTER 1991 caceae seed viability during storage seed science flood flows and dynamics of sonoran riparian forests and technology 5 509 517 rivers 2 221 235 TOMANEK G W AND R L ZIEGLER 1962 ecological received 14 march 1994 studies of salt cedar unpublished report division of accepted 12 august 1994 great basin naturalist ssi551 0 1995 appp 66 73

NAMES AND TYPES OF HEDYSARUM L FABACEAE IN NORTH AMERICA

stanley L welshlwelsh1welshel

ABSTRACT the names and types of hedysarum L sensugensu strictostnctostripto for north america are included along with biblio- graphic citations type information and place of deposit of types and all synonyms lectotypes are designated for hedysarum auriculaaunculatwnauriculatumtum eastweasaw H carnucarnulosumlosum greene H marginatummargimangimargmatumnatum greene H tabularepabulare A nels and H truncatumtruncatum eastweasaw

key words hedysarum types nomenclature

the following list of names and types in ale glacial events during the pleistocene have hedysarum L was prepared preliminary to been suggested as having separated the sub- submittal of a summary revision to the flora sets allowing them to achieve the degree of north america project the genus hedysarum morphological and genetic integrity of the L as here interpreted for american taxa extends present populations the present juxtaposition from the bering strait to newfoundland and is suggested to have resulted by expansion of vermont and from the polar sea and the cana- the respective entities into areas previously dian arctic archipelago south through the occupied by glaciers mountains and plains of western north america the rather large number of names involved to oklahoma new mexico arizona and in the genus is indicative of variation inherent nevada excluded from this treatment are in the various taxa flower size plant size those taxa originally included in hedysarum leaflet size and pubescence are features vari- which are now interpreted as belonging to able in both complexes apparent correlation other genera ie to desmodium the genus in of two or more of these features has served as the restricted sense consists of two complexes justification for several names indeed when ie those with leaflets thickened and veins one observes dwarf large flowered plants in obscured the boreale complex and those with either complex there appears to be a compel- relatively thin leaflets in which the veins are ling need for their recognition however rather readily apparent the alpinum complex much if not all of the variation is haphazard the earliest taxon within hedysarum alpinum or the attempt at segregation devolves to use complex is that by michaux 1803 who estab- of a single characteristic such as presence or lished the trinomial hedysarum alpinum ameri absence of pubescence which fails also there canum michxmicha the boreale complex was initi- are few truly diagnostic characteristics once ated by nuttall 1818 with the publication of the two complexes are separated the taxono- H boreale mist ultimately must rely on a series of varying taxa in the two complexes demonstrate re- features to identify a particular specimen markable morphological and geographical par- fortunately the taxa are with some notable allelism each consists of additional taxa sepa- exceptions disjunct from each other if the rable generally into two geographical sub- disjunction is not apparent from examination groups juxtaposed at or near the soth parallel of a distribution map it is often apparent in of longitude somewhat north of the canada the field where the plants grow in different US boundary north of that parallel lies most habitats for example the range of yellow ofhofjofhH boreale sspasp mackenzii richards welsh flowered H sulphurescens apparently overlaps and most of H alpinum sensugensu strictostricko to the that of pink purple flowered H occidentale in south occurs H occidentale greene most ofofhjofhH large part yet they seldom occur together sulphurescens redbrydb limited extensions of H and only an exceptional intermediate is alpinum L and most of H boreale sspasp bore known

depditmentidepartment of botany and life science museum brigham young Univeruniversityuniversituniversiasit provo UT 84602

66 199511995 NORTH AMERICAN HEDYSARUM 67

there are in spite of gross similarities of still another trend resulting in the forma- the taxa within the respective complexes few tion of inconsequential names was the well recorded intermediates intentioned effort to provide epithets for spec- adding to the difficulties of interpretation imensamens differing in insubstantial ways iei e the of the north american materials is the inter- naming of white flowered or teratological speci- rupted circumboreal distribution of H mens as formaecormae alpinumalpinum a species with several close allies in the following list is thought to be exhaus- siberia the initial interpretation by michaux tive for hedysarum names in north america of north american H alpinum as being taxo- pertinent types have been received on loan nonomically different H alpinumalpinum ametiamerlameriameficanumamericanumcanum through the kindness of curators of herbaria from that of the old world has paraded alparappar cited with the specimens abbreviations for itionaition like through most subsequent treatments herbaria are those standard ones cited in information is of the genus unpublished work by northstrom index herbanorumherbariorumHerbanorum type is pre- sented below in dual format for some taxa with 1974 refuted the claim to difference between in type information type locality as recorded siberian and american phases of the species with the protoprotologuelogue cited first and label data at least as far as broad categories were con- of the type specimen type cited second where cerned the claim that north american mate- there is a substantial difference in the two is likewise rials constitute a separate entity re- accounts futed by comparison of specimens from siberia and north america in the present study other hedysarum albiflorum macoun fertschfedtsch acta hort workers have asserted that large flowered petrop 19 252 1902 Basi H boreale macoun low growing plants of the species are identical basionymonym var albiflorum H sulphurescens redbrydb with substantial asian taxa ie H hedysari oides L schinz & thellungThellung astragalus hedy hedysarum alpinum L sp PIpl 7501753750.1753750 1753 type locality in Sisiberia 1 claims investigatedd habitat in berisberlsberid linnaeus ic sarisarioidesoides L such were investigate typeTpypeype possible lectotype 92154921 54 LINN micromicrofichemicroficbefiche by northstrom 1974 who determined that BRY there is little basis for such assertions evidence hedysarum alpinumalpmum var ameriamerlamericanumamertcanum michamichx fl bor to support the conspecific nature of the sup- amer 2 74 1803 posed entities is apparent when localities of H alpinum L such supposed taxa in western alaska are synonyms H alpinum sspasp americanumamericanum michamichx examined and plants with larger flowers are fertschfedtsch acta hort petrop 19 255 1902 in part H americanumamericanum michamichx britt mem torrey bot found to occur within populations having club 5 201 1894 small flowers and that flower size within the type locality in boreaborealibuslibus canadaecanadas et in cataractiscataractis species in a broad sense forms a continuum montium allegianisalleghanisalleghanis and occasional tall specimens within the type hedysgedys J iin p 74 75 herb MX isotype NY there is a mounted half herbarium sheet at NY alpinum complex also bear large flowers torrelltorreyltorrey bearing a large portion of a stem with a leaf another factor leading to the creation of a and mature strigose fruit ofH boreale var boreale large number of synonyms was the early mis- this specimen is apparently superfluous probably interpretation of specimens ofhjofhofH alpinum under having been added later when additional material be- the name H boreale this switching of names came available to dr torrey from western american while not uniquely a problem in this genus collections but more pertinent to the present case also has an attached fragment envelope became of great importance to those workers the sheet which is the type information noted and on is written who encountered the genus piecemeal above the envelope contains a portion of an inflo- treated the variants as though they had not rescencerescence a flower and several immature loment already been named it was not helpful per- segments the segments are glabrous have a defi- haps that the most ardent authors of western nite winged margin and are identifiable as H american plant names should be involved with alpinum L it is probable that the specimen from the genus ie edward L greene per axel which the fragments were removed is with the michaux herbarium at P rydberg and aven nelson greene as the continued recognition of the american materi- record indicates was prone to name the same als of H alpinum at any infraintraspecificinfraspecificspecific rank is fraught species several times in this and other genera with difficulties there are no diagnostic features not recognizing or possibly not caring that he known that will allow segregation of the american was renaming the same taxon specimens from the asiatic ones 68 GREAT BASIN naturalist volume 55 hedysarum alpinum var americanumamericanum f albiflorum white flowered specimens occur sporadically standl fern rhodora 35 275 1933 through populations of taxa with generally pink basionymBasionym H ameriamericanumamertamerlcanum f albiflorum standl purple flowers their recognition at any taxonomic H alpinum L rank is probably moot and the publication of the publication by 1933 the fernald recognizes taxon by standley 1930 is therefore inconsequential white flowered plants from newfoundland hedysarum amenamericanumamencanumametiamerlamericanum var mackenzimackenzemackenzni richards britt hedysarum alpinum var grandiflorgrandiflorumum rollinsbollins rhodora Mmemem torreybottorrey bot club 5 202 1894 42233194042233.194042 233 1940 basionymBasionym H mackenzimackenziimackenziei richards type newfoundland pistolet bay mossy and turfy H boreale sspasp mackenzi i richards welsh trap cliffs and talus anse aux salvagesSasauvagesuvages M L fernald K M wiegand and bayard long 28625 hedysarum auriculaauriculatumaurtculatumtum eastweasaw bot gaz 33 205 1902 august 11 1925 holotype GH H alpinum L port port paratype new foundlandFoundland region of a bay type alaska cape nome blaisdell sns n summer 1900 no 10849 in humus or turf on the limestone table- lectotype selected here chiGHghi isolectotype usiUS land altitude 200 300 m table mountain M L specimens on which H auriculaaunculatumauriculatumtum is based fernald and H st john july 16 & 171914 CAN were distributed from the california academy of BM sciences herbarium with collection information is based on large flowered about 16 this name is recorded on labels of that institution only mm long low from newfound- the growing specimens known specimens in contemporary collections are land are other similar plants scattered there those at GH and US the two specimens consist of through much of the distribution of H alpinum in in almost identical branches of H alpinumalpinum with both north america but they are more consistently rep- flowers and fruit although that at GH is designated frigid inhospitable resented in or other arctic or on the label as a duplicate of the type which was subarctic even in the cited with sites in type serlesseriessetles presumably at CASGAS prior to the san francisco the protoprotologuelogue there is considerable variation is the earthquake early in this century fire resulting from paratype cited above differs significantly from the that devastating tragedy destroyed much of the holotype specimen it is much taller and has flowers early academy herbarium ofaof a size intermediate with those ofofspecimensspecimens more usual for var alpinum inm a more strict sense indeed hedysarum bakeri greene ex redbrydb bull agne exper the low growing larger flowered phase appears to sta colorado 100 215 1906 pro syn be a phenotypically recurring recombinant form H boreale nutt var boreale within a complex exhibiting much variation in intended type flora of colorado plants the gunnison flower size and other features however size of watershed cimarron june 28 stems in large clus- ters 8 to 1 ft on dry slopes in flower is not always correlated with plant height or in 11212 open collected in 1901 by C no flower number all possible combinations of flower F baker 274 NDG evidently the name was never published by E L size flower number and plant height are represent- greene but was cited as a synonym of H tabularepabulare ed in the species as a whole it is possible to write a A nelson by rydberg in his flora of colorado key that will separate these plants but it seems that in the intended type has three mounted stems showing such a key will not then be segregating natural taxa flowers and maturing fruit they are strigose both on herbage and on the lomentsfoments plants differ in hedysarum alpinum sspasp philosociaphilosocia A nels love & the in taxon 31 no material way from a great many specimens from lovetaxon31love 347 1982 greene basionymBasionym H philosociaphilosocia A nels colorado perhaps also realized as much H alpinum L hedysarum boreale nutt gen N amer PI 2 110 1818 hedysarum alpinum var philophilosociasocia A nels rollins type locality north dakota around fort mandan on rhodora 42 224 1940 the banks oftheofodthethe missouri nuttall 1 c basionymBasionym H philosociaphilosocia A nels type hedysarum boreale sources of the missouri H alpinum L nuttall probably late june 1811 boloholotypetype BM the name H boreale was early transferred to the hedysarum ameriamericanumcanum michxmicha britt mem torrey concept of H alpinumalpinum and part of the synonymy bot club 5 202 1894 reflects attempts by various authors to resolve the basionymBasionym H alpinum var americanumameriamerlcanum michxmicha apparent lack of a name for this wide ranging and H alpinum L highly variable species nuttall torrey and gray 1838 named the species a second time as H hedysarum americanum E field mus americanum f albiflorum standl canescentcanescenscanescens based on from along the pub bot 815.19308 15 1930 specimens 8151930 1834 H alpinum L snake river in present idaho taken in he synonym H albiflorum macoun fertschfedtsch was in the vicinity of fort hall idaho from 14 july type alaska davidson glacier july 4 1929 william to 6 august 1834 mckelvey 1955 602 whether he S cooper & frances E andrews 95 herb field noted the similarity between earlier and later mus no 598264 type holotype F named materials is not known it seems likely that 199511995 NORTH AMERICAN HEDYSARUMHEDYSARCM 69 he did not have authentic material of the earlier hedysarum boreale var leucanthumleucanthum greene M E jones named taxon at hand for comparison with his col- proc calif acad sci 5 677 1895 lections on the 1834 wyeth expedition the type at basionymBasi onym H mackenzii var leucanthumleucanthum greene H boreale sspasp mackenzi BM a solitary flowering stem is mounted with sev- i richards welsh eral flowering stems of H alpinum of unknown col- hedysarum boreale sspasp mackenzi lector i richards welsh great basin nat 28 152 1968 basionymBasionym H mackenzi i richards hedysarum boreale var albiflorum macoun cat canad PIpl 1 510 1884 nom nud sio5101884510.1884 hedysarum boreale var mackenzimackenziimackenziei richards C L H sulphurescens rydb redb hitchchitche vase pi pacific N W 3 275 19619611 syn H albiflorum macoun fertschfedtsch basionymBasi onym H mackenzii richards type locality this form is peculiar to the foothills and drier slopes and is abundant from the mountain is hedysarum boreale var mackenzii f niveum boivin Kananakananaskisskis through the rocky mountains to the boivin naturalisteNaturaliste canad 93 433 1966 columbia valley at donald 51 macoun lat basionymBasi onym H mackenzi var mackenzimackenziimackenzie f eastern summit of the north kootanieKoo pass rocky i i niveum tanie boivin mountains ie1 H boreale sspasp mackenzimackenziimackenzie richards welsh type geological and natural history of canada no i 111153901111 5390 hedysarum boreale var albiflorum iiiilili hedysarum boreale var obovatum rollins rhodora 42 dry soil east summit of north kootanieKootanie pass R 2351940235.1940235 1940 mts dawson july 29th 1883 and geological and H boreale nutt var boreale natural history survey of canada no 533 5389 vaival type county E hedysarum boreale nutt var alba macoun nevada eiloeikoelko thorpe creek of 1928 H price mountain slopes Kananakananaskisskis rocky mts macoun lamoile 25 july H 168 holotype photo RM june 24th 1885 syntypessyntypes CAN there is no description aside from the designation hedysarum boreale fE proliferum dore boivin naturalisteNaturaliste albiflorunalbiflorum proposed as an epithet only other the canad946301967canad 94630196794 630 1967 provided by macoun information aside from that basionymBasionym H mackenzii f proliferum dore related with the locality data is the statement this H boreale sspasp mackenzimackenziimackenznmackenziei richards welsh fine plant is closely related to H boreale but is cer- tainly distinct the name is regarded as a nomen hedysarum boreale var utautahensehense redbrydb rollinsbollins rhodora nudumdudum the collection by dawson is nevertheless 42235194042235.194042 235 1940 an excellent flowering example of H sulphurescens basionymBasionym H utautahensehense redbrydb and the macoun sheet consists of two plants with hedysarum boreale nutt var boreale both flowers and immature to mature fruits both hedysarum canescenscanescens in torr & gray N amer also H sulphurescens the indication by macoun of canescent nutt in fl 1 357 1838 not H canescentcanescenscanes cens L relationship of var albiforumalbi to H boreale reflects forum Basibasionymonym H rydb H boreale var cincm general by cinerascens redb the misapplication many american erascenserascens redbrydb rollins botanists of H boreale to the alpinum complex in H boreale nutt var boreale north america of which H sulphurescens is a por- type locality idaho plains of the rocky mountains tion macoun used the number 533 for several col- particularly near lewis s river nuttall ie1 lections of hedysarumofhedysarum taken from 1883 to 1885 type hedysarum canescentcanescenscanes cens H mackenzii hook R mts lewis snake R nuttall sns n probably in hedysarum boreale var cinerascenscineraseens redbrydb rollins july 1834 boloholotypetype PHPHI isoisotopesisotypestypes chiGHghi BM 2 rhodorarhodora4242 234 1940 sheets basionymBasionym H cinerascens redbrydb et H canescentcanescanescenscens nutt the specimen at PH which is mounted on a in seq sheet with two other superfluous collections bears H boreale nutt var boreale the date july 12 with the incorrect year date 1833 obviously added later nuttall was with the hedysarum boreale var f album boivin cinerascens wyeth expedition 1834 and on 12 july was a Naturnaturalistenaturahstealiste canad 87 34 1960 in H boreale nutt var boreale short distance east of where fort hall would be type canada saskatchewan maple creek district constructed subsequently despite the existence of eastendEastend hillside along river valley 19 july 1950 the earlier named H boreale with which H R C russell S 5075 holotype at DAO canescentcanescenscanescens is synonymous this name or its substi- tutes would be featured prominently in 19th cantucentu hedysarum boreale var flavescentflavescensflavescens coult & fisher ry accounts of the genus in the american west fertschfedtschFedtsch bull herb boiss 7 256 1899 there are two of nuttall s specimens on the sheet basionymBasi onym H flavesflacescensflavescentflavescenscens coult & fisher at GH each provided with a label both with flow- H sulphurescens redbrydb ers and both representing the same taxon the label hedysarum boreale var gregremialegrembialemiale rollins northstrom & information consists of the following hedysarum welsh great basin nat 30 125 1970 canescentcanescenscanescens H mackenzii R mts and hedysarum basionymBasi onym H gregremialegrembialemiale rollins canescentcanescanescenscens R mts since no additional locality 70 GREAT BASIN naturalist volume 55 information or date accompanies the labels the sta- june 1937 R C rollins 1733 holotype GH iso tus as exact duplicates is unknown it seems likely types RM US CAS UTC MONTU PH F that both were included within the concept of H this taxon stands on the feature of lateraloflateral spines canescentcanescenscanes cens by nuttall and both can be regarded as on the loment segments it is otherwise indistin- isoisotopesisotypestypes there is a second possible isotype of H guisguigulguishablebablehablehabieshableshabie from plants of var boreale by which it is canescentcanescenscanes cens at BM hedysarum mackenzimackenznmackenziimackenziei fort surrounded northstrom and welsh 1970 hall prairie common aug it lacks the usual for nuttall s labels and his name is not in evidence hedysarum lancifoliumlancifolium redbrydb mem new york bot but the handwriting appears to be his cardgard 1 256 1900 H occidentale greene type mountain woods near head waters of jocko hedysarum carnucarnulosumlosum greene Pitpitroniapittoniatonia 3 212 1897 montana pale M H boreale nutt var boreale river flowers purple W canby 93 july 15 1883 holotype NY type locality common in clayey soil about the mouth of the canon of the arkansas inm southern colorado the type specimen consists of a folded plant greene ie1 c some 75 cm tall bearing leaves and flowers and a type colorado fremont co plants of colorado stem fragment bearing immature fruit mature canon city 8 sept 1896 edw L greene lecto- flowers are about 16 mm long on the short side of type here designated NDG 2 isolectotypes also the variation in H occidentale but the fruit even NDG though immature is of the size and form of that while no specimens were cited with the original species description the three specimens so named in greene s handwriting at NDG are most certainly hedysarum leucanthumleuc anthum greene greene Pitpittoniapitroniatonia 3 213 type material all bear the same date and locality in-in 1897 formation the specimen bearing the Greenegreeneanumanum basionymBasionym H mackenzi i var leucleucanthumanthum greene herbarium number 35686 is here chosen as lecto- H boreale sspasp mackenzi i richards welsh type the others 35687 and 35688 are considered isolectotypes the lectotype has both flowers and hedysarum mackenzimackenziimackenziei richards in franklin ist journey bot append 745 1823 fruit the other two are in fruit and flower with in H boreale sspasp mackenzi i richardson welsh immature fruit respectively all have strigose type canada barren grounds point lake to the herbage and lomentsfoments usual flowering time for the arctic sea richardson s n holotype BM photo species is april to late july Is it possible that the casCANCAM isotype NY columbia 1 hedysarum macken species flowered again following late summer rainsrams zil franklinfranklinss journey dr hooker isotype PH at canon city in september 1896 astr crossed out hedysgedys Mackenzimackenziimackenziei richard N W amer fr eapexp dr ho and herb A gray H Mackenzii torr & gray N amer hedysarum cinerascens redbrydb mem N Y bot cardgard 1 fl richardson 257 1900 nom nov pro H canescentcanescenscanes cens nutt BDGB D G isoisotopesisotypestypes GH H boreale nutt vaivalvarvan boreale evidently dr william jackson hooker sent rep- basionymBasi onym H canescentcanescanescenscens nutt resentative material obtained by john richardson syn H boreale var cinerascens redbrydb rollins botanist with the franklin expedition to the this material was retained by northstrom 1974 philadelphia academy and to john torrey and asa at varietal rank the taxon stands on the sole char- gray collections from the franklin expedition acter of pubescence and a plotting of the distribu- demonstrate the variability represented in a rather tion of hairy versus glabrous plants demonstrates large set of specimens each matched by modern much overlap the specimens can be separated but collections of the taxon the second sheet cited at do they represent taxa GH is doubly mounted with a mere fragment pre- sumed to have come from the franklin expedition hedysarum flavescentflavescensflavescens coult & fisher bot gaz 18 300 in the lower portion and a second fragmentary col- 1893 non regel & schmalhschmald lection by burke apparently a phase of H boreale basionymBasionym of H sulphurescens redbrydb from the rocky mountains the latter material is type montana may 1892 near helena F D kelsey not a portion of the type of H mackenzii sns n holotype Ffl is unfortunate that epithet it is the flavescentflavescensflaves cens was hedysarum mackenzimackenziimackenziei var fraserifrasheri boivin canad field occupied it fits well the description of flower color nat 65 20 1951 in this taxon many of the kelsey collections are in H boreale nutt var boreale the USU S national museum eliseselisensElisens 1985 but the type locality canada saskatchewan W P fraser type of H flavescentflavescensflaves cens is at F where coulter s langham river valley june 12 and 26 1938 ae1e1 c herbarium is deposited type hedysarum Mackenzi i richards river valley Langlanghamhambam sask wpfraserWP Fraser june 12 & 26193826 1938 hedysarum gregremialegrembialemiale rollins rhodora 42 230 1940 holotype DAO H boreale var gregremialegrergrermalegrembialemalemaiemiale rollins northstrom & boivin lc1 c cites the revisionary treatment of welsh rollins 1940 as indicating that H mackenzimackenziimackenziei var type utah uintah county ca 14 mi W of vernal 16 mackenzimackenziimackenziei has flowers 18 21 mm long but with 199511995 NORTH AMERICAN HEDYSARUM 71

some 16 20 mm prairie plants from saskatchewan legumes native to alaska in certain of those in-in and alberta however have flowers 13 15 mm long stances the inflorescence typically elongates flow- these latter plants are the basis of his var fraprasenshriseri ers become erect on attenuated pedipedicelscels petals are the type of var brasen represents H boreale sspasp deformed and the ovary is typically exserted from boreale var boreale which is plesentpresent along the the flower in some specimens at least the ovary is plains and foot slope of the rocky mountains and filled with black spores the type specimen of f has flowers of the size indicated for the variety prohferumproliferum exhibits another variant than that typi- cally encountered the inflorescence is shortened hedysarum mackenzi i var leucanthumleucanthum greene Pitpittoniapitroniatonia and modified flower buds are in tight clusters 229418922294.18922 294 1892 whether elongate or compact specimens on which basionymBasionym of H leucanthumleuc anthum greene greene such aberrations are based are not taxa and the H boreale sspasp mackenzi i welsh richards need to name them is therefore moot type locality on the porcupine river northern alaska mr turner greene 1 c J J ie hedysarum macquenzii f canescenscanes cens nutt fertschfedtschFedt sch type plants of alaska collected on the porcupine canescent actaaetaactahorthort petrop 19 272 1902 river 1891 mr J turner holotype NDG greene basionymBasionym H canescentcanescenscanescens nutt 1892 notes that this is far more than H boreale nutt var boreale an albino state of H Mackenzi i perhaps identical with some asiatic species but the plants were just hedysarum marginatummarginatum greene Pitpittoniapitroniatonia 4 138 1900 coming into flower when gathered in that there is H occidentale greene no trace of the loment he later 1897 elevated it type locality mountains above cimarron southern to species rank however except for white flowers colorado collected by the writer 30 aug 1896 pagosa which occur with some frequency in the species also near springs colo 26 july 1899 C F baker greene ie1 c the plant differs in no respect from numerous other type colorado plants of colorado near cimarron plants from the range of the taxon generally arctic 30 aug 1896 edw L greene lectotype NDG here chosen plants of southern colorado pagosa hedysarum mackenzii var mackenzii f niveum boivin springs 26 july 1899 C F baker syntypessyntypes NDG canad field nat 65 20 19519511 NY RM GH F basionymBasi onym of H boreale var mackenzi f i niveum the lectotype collected by greene is in fruit boivin boivin synsyntypestypes at NDG and NY have both flowering and H boreale sspasp mackenzi richards welsh i fruiting branches both the lomentfomentmoment articles type yukon territory J W abbott 17a pine creek in are sandy land june 7 1946 holotype DAO markedly winged and strigose herbage is strigose greene the collection consists of five flowering stems of also see 1900 H boreale sspasp mackenzii all with white flowers hedysarum occidentale greene Pitpittoniapitroniatonia 3 19 1896 condition of white flowers is occaoceaoccasionalsionaslona j the type locality 1890 throughout the subspecies and hardly worthy of olympic mountains washington C V piper greene le1 c taxonomic consideration type washington olympic mts C V piper 905 flowers 11 august 1890 fruit 30 sept 1890 hohholo- hedysarum mackenzii var tabularepabulare A nels kearney & type NDGNDGI peebles S J wash acad seisciselci 29 485 1939 greene 1896 provides a description and a short pabulare Basibasionymonym H tabulare A nels note plant like H boreale when in flower though hedysarum boreale nutt var boreale in with broader leaflets and widely different fruit A hedysarum boreale var leucanthumleucanthum greene M E jones second sheet from the olympic mountains at NDGNDGI proc calif acad II11 5 677 piper 2227 august 1895 has the epithet occiden- H boreale sspasp mackenzii richards welsh tale in greene s hand but it was not cited by him syn H leucanthumleucanthum greene greene it is much better material than the type for a long time the name H boreale was included within the hedysarum mackenzii f proliferum dore canad field concept of H alpinum it is likely that greene was nat 73 151 1959 under a similar misconception the general aspect basionymBasionym of H boreale f proliferum dore boivinbolvin of H occidentale iei e conspicuously veined leaflets H boreale sspasp mackenzii richards welsh and large lomentsfoments with prominently reticulate type plants of alaska A single clump in shallow soil broken rock beside petalif- venation which occurs from vancouver island over growing common peninsula erous plant cf 4983 north side of tanana ririverriveniverlver british columbia and the olympic mile 277 richardson highway 6410n 1455214552wW washington disjunctdisjunctlyly eastward to northern and W J cody & T J M webster 4984 june 3 1951 eastern idaho western montana western wyoming holotype DAO I1 northeastern utah and montane southern colorado this name is based dore 1959 on a teratologi- is that of H alpinumalpinum and it differs generally in the cal specimen of H boreale sspasp mackenzimackenziimackenziei a recur- manner indicated by greene ring variant induced by a pathogen likely a smut the most distinctive feature separating most if fungus teratology occurs in several if not all boreal not all specimens of H occidentale from H 72 GREAT BASIN naturalist volume 55 alpinum is the much larger rather conspicuously material from the black hills of south dakota wing margined loment segments flowers are gen- and from southeastern wyoming is morphologically erally larger often much larger however plants similar and has been recognized as belonging to a from the absaroka range of northwestern taxon that survived south of the major glacial events wyoming approach H alpinum in occasionally hav- of the pleistocene the mainmaln diagnostic criterion is ing small flowers but when collected at maturity however loment pubescence that feature is incon- the fruit is that of H occidentale additional collec- stant within the southern material and often is pres- tions might demonstrate that H alpinum per se is ent in plants far beyond its supposed range which indeed in the absarokasAbsarokas large flowered plants of has been plotted to include plants as far north as H alpialpinumnum mainly of frigid sites in the arctic the soth parallel recognition of plants at any taxo- approach the size of flowers of some H occidentale nomicnomie rank is therefore problematical specimens but the fruit there is that ofhofjofhH alpinum H occidentale has lomentsfoments very similar to those of hedysarum roezlianumromlianum prandprantiprantl ind sem hort wircebwizceb the closely allied H sulphurescens 818738.18738 1873 H boreale nutt var boreale I1 have to find hedysarum occidentale greene var canone welsh great been unable any reference to this basin nat 38 314 1978 taxon aside from its citation by rollins 1940 type utah carbon county ca 14 mi due ENE of helper soldier creek 30 june 1977 welsh & hedysarum sulphurescens redbrydbbydb bull torrey bot club taylor 15256 holotype brylBRY isotype at NY 24251189724251.189724 251 1897 syndrome of characters associated with this basionymBasionym H flavescentflavescensflaves cens coult & fisher not regel & the schmalhsebmalhschmald taxon is shared individually elsewhere within the is yellow to yellowish flowers easily distinguish as a whole from species however specimens this entity which shares the peculiar loment fea- duchesne carbon and emery and counties utah tures ofjofh occidentale ranges from the gunnison county recognizable ofh the species cunnison colorado are by southern british columbia alberta rockies south thick to their large ovate ovate lanceolate yellow through north central washington northern idaho green leaflets and large plants pale flowers are western montana and northwestern wyoming known from rather xericxene sites in pinyonpryonpmyon juniperjumper and mountain brush communities whereas plants of the hedysarum truncatruncatumtum eastweasaw bot gaz 33 205 1902 type variety are mainly of more mesic sites H alpinum L although the taxon is segregated on weak diagnos- type alaska nome dr FE E blaisdell sns n summer tic features it seems to be at least a trend worthy of 1900 lectotype NYnyl here designated lectotypeisoisolectotype taxonomic recognition it has long been known in GH collections type material is low about 2 252.52bs85 5 dm tall has mature flowers about 12 mm long and has fruit the hedysarum tabularepabulare A nels proc biolabiol soc wash 15 size and conformation of H alpinum it is identical 1851902185.1902185 1902 for all practical purposes with material named by H boreale nutt var boreale eastwood simultaneously as H auriculatumauricula tumturntuyn and type wyoming wind river dubois A nelson 752 taken at the same place and time by the same col- 1894 lectotype here designated RM lector in 1900 this name is based on several wyoming proc colorado and utah syntypessyn types iei e M E jones hedysarum uintauintahensehense A nels biolabiol soc wash 5592 soldier summit utah in 1894 POM BM 15186190215186.190215 186 1902 greene F snake river wyoming A nelson 3496 19 H occidentale 1 type draws of the foothills evanston august 1899 RM wyoming natrona co bates wyoming in A nelson 7198 14 june 1900 holotype RM iso L N 201 5 july 1901 RM iso creek goodding F types RMRMI NY GH type sheets uniformly bear thick lanceolate to hedysarum pabulare var nvularerivulare L ann tabulare rivulare 0 williams lance ovate leafleafletsfets similar to var canone but with missouri bot cardgard 21 344 1934 flowers al H plants H boreale nutt var boreale of typictypical occidentale from south- type wyoming teton county along the snake river west wyoming are not uniformly of the uintauintahensehense 31 july 1932 L 0 williams 975 holotype RM type but vary from one population to another with isoisotopesisotypestypes CHGH caslCASCASI most being similar to traditional H occidentale hedysarum philphilosciaoscia A nels proc biolabiol soc wash 15 hedysarum utautahensehense redbrydb bull torrey bot club 34 1851902185.1902185 1902 4241907424.1907424 1907 basionymBasionym H alpinum var philphilosciaoscia A nels rollins H boreale nutt var boreale H alpinum sspasp philphilosciaoscia A neknelsneis love & love type utah salt lake county vicinity of salt lake H alpinum L city utah leonard 55 26 may 1883 holotype type wyoming albany county head of crow creek NY laramie mountains 1896 A nelson 2034 holotype the type consists of two complete stems and a rmlRMerml fragmentary branch it is typical of the material 199511995 NORTH AMERICAN HEDYSARUM 73 growing through much of utah and elsewhere in MCKELVEY S D 1955 botanical exploration of the trans the west mississippi west 1790 1850 arnold arboretum jamaica plains 1144 appp northstrom T E 1974 the genus hedysarum in north references america unpublished master s thesis brigham young university provo UT DORE W G 1959 some inflorescence forms in clovers northstrom T E AND S L WELSH 1970 revision of the and other legumes canadian field naturalist 73 hedysarum boreale complex great basin naturalist 147 154 3010930 109log 130 ELISENS W J 1985 the montana collections of francis nuNUTTALLriall T 1818 the genera of north american plants duncan kelsey Britbnttoniabrittaniabrittoniatonia 37 382 391 volume 2 D heartt philadelphia PA genus fedtschenkoFEDTSCHENKO B 1902 A revision of the hedy- MMICHAUXichauxmichaux A 1803 flora boreali americana 2 1 340 sarum acta hortus petropolitaniPetropolitani 19 185 325 ROLLINS R C 1940 studies in the genus hedysarum in FERNALD M L 1933 recent discoveries in the newfound- north america rhodorarhodora4242 217 239 land flora bnttoniabrittaniabrittoniaBrittonia 35 265 283 STANDLEY P C 1930 studies of american plants ililiiIII111 GREENE E L 1892 new or noteworthy species XIV field museum of natural history botanical series 8 Pitpittoniapitroniapittoma2tonia 2 293 298 3 73 1896 new or noteworthy species XV Pitpittoniapitroniatonia TORREY J AND A GRAY 1838 flora of north america 3123 12 28 volume 1 wiley & putnam new york NY 1897 new or noteworthy species XIX Pitpitroniapittoniatonia 32123 212 230 received 28 february 1994 fascicle of pittoniapitronia 4 1900 A new papilionaceae Pittonia accepted 3 june 1994 132 139 great basin naturalist 551 0 1995 appp 74 77

WHIPWORM TRICHURIS DIPODOMYS INFECTION IN KANGAROO RATS DIPODOMYS SPESPP EFFECTS ON DIGESTIVE efficiency

james C muegerlmungerlmunger1 and todd A shchterlshchter1

ABSTRACIABSTRACT to determine whether infections by whipwormswhip worms trichuris dipodomys nematoda Trichtnchuratatrichurataurata tnchundaetrichuridaetrichiuridae might affect digestive efficiency and therefore energy budgets of two species of kangaroo rats dipodomys micronsmicropsrnicrnicropstopsrops and dipodomys ordu rodentia heteromyidae we compared the apparent dry matter digestibility of three groups of hosts those naturally infected with whipwormswhipworms those naturally uninfected with whipwormswhipworms and those origi- nally naturally infected but later deinfected by treatment with the anthelminthic ivermectin prevalence of T dipodomys was higher in D micronsmicrops 53 than in D ordnordiiordia 14 apparent dry matter digestibility was reduced by whipworm infection in D micronsmicrops but not in D ordiiorduordia although a statistically significant effect was shown its small mag- nitude indicates that whipworm infection is unlikely to have a biologically significant impact on the energy budgets of host kangaroo rats

key words parasite digestive efficiency whipwormwhipwonnmonnmown kangaroo rat trichuris dipodomys energy budget

parasites inhabiting the gastrointestinal species were captured at the site ammospennoammospermo tract of a host may reduce the efficiency of the philus leucurusleucurus neotoma lepidacepida perognathus organs they inhabit either through direct com- flavus peromyscus maniculatus and two petition for nutrients or through damage to species of kangaroo rats dipodomys ordiiordia and absorptive surfaces because decreased diges- dipodomys micronsmimicropscrops dipodomys ordiiordia ranges tive efficiency may reduce the rate of energy from 42 to 72 g and consumes a diet consisting input into a host gastrointestinal parasites have primarily of seeds zeveloff 1988 dipodomys the potential to cause a change in host energy micronsmicrops is larger 72 91 g and is unique among allocation eg reduced activity or reduced kangaroo rats in that it relies heavily on leaves reproduction and thereby impact the ecology of atriplex conferticonfertifoliafolia for forage kenagy of the host munger and karasov 1989 1972 zeveloff 1988 both species are liable to tapeworm infections have a measurable infection by the whipworm trichuris dipodo- effect on digestive efficiency but a biologically mys a nematode that inhabits the cecumcaecum of unimportant effect on the energy budget of infected hosts Grundgrundmannmarm 1957 whitaker et host white footed mice peromyscus leuleucopuscopus al 1993 munger and karasov 1989 the present study on the study site we established a 13 X 13 was designed to determine if infection by a grid of 169 sherman live traps baited with nematode the whipworm trichuris dipodomys millet and placed at 15 in intervals during has a substantial effect on one aspect of the two trapping sessions 14 22 june and 15 18 energy budget digestive efficiency of host august 1990 kangaroo rats 30 individuals of kangaroo rats dipodomys micronsmicrops and D ordiiordia D micronsmicrops and 85 of D ordiiordia were captured and brought into the laboratory fecal speci- MATERIALS AND METHODS mens from each animal were analyzed for the presence of parasite eggs by standard centrifu- our study site located 2 km N of murphy gal flotation techniques using saturated sucrose owaheeowyhee county ID is in desertdesertscrubscrub habitat solution pritchard and kruse 1982 six in- with sandy loam substrate primary shrub fected but untreated animals from the june species of the study area are artemisia experiment were included in the pool of ani- spinescentspinescensspines cens artemisia dentatatritridentatatridentate atriplex mals used in the august experiment the few canescentcanescenscanescens atriplex conferticonfertifoliafolia atriplex spin- animals that failed to thrive in the lab were osa and chrysothamnus nauseosus six rodent removed from the experiment data from a

departmentidepartment of biology boise state university 1910 university drive boise ID 83725

74 199519951 WHIPWORMS IN KANGAROO RATS 75 total of 29 D micronsmicrops individuals and 56 D 02 131111311 1 d ordiiordia were analyzed infected each month s set of captures was subjected 0 uninfected to the following protocol 01 1 kangaroo rats were acclimated to a diet 11 of millet seed for 3 11 d r sm

2 A pretreatment feeding trial was per- 0 formed animals were placed in wire bottomed 0 3 cages with a measured amount of whole millet 5 seed at the end of 5 d fecal pellets were sep- 01 arated from spilled food and dried 24 h at 10

50500soo C initial digestive efficiency of each ani- 11 mal was measured as apparent dry matter 02 digestibility ie the proportion of mass con- D micronsmicropsmicromicrg D ordlinordli sumed but not lost as waste which was calcu- lated as MFO MFE MFO where MFO aidaldand fig 1 effects ofvariationof variation in parasite load on propor- tional change in dry matter digestibility means SE migMFE are the mass of food consumed and feces in t produced respectively numbers represent sample sizes 3 half of the infected animals were then injected subcutaneously with a solution of some minor morphological differences from ivermectin a systemic anthelminthic ivomec the original species description read 1956 brand from MSD AGVET rahwayradway NJ do exist perhaps as a result of geographical figure I1 gives sample sizes of treatment groups variation the specimens most closely match june captures received on each of two con- read s description of T dipodomys A shostak secutive days a 02ogee cc injection of ivermectin personal communication measurements of in 40 glycerol formal and 60 propylene several key morphological characters are as glycol each injection delivered ca 350 agg follows X SD total length 6 25625.625956gsg 6 080.80 8 ivermectin kg body mass controls received mm Y 41341.341 3 292.92 9 mm hindhindbodybody length dS equal volume injections of the glycerol for- 12712.712 7 040.40 4 mm Y 23723.723 7 191.9iglg1 9 mm spicule mal propylene glycol carrier this dosage had length 850 85185.185 1 lumamjom egg length 64864.864 8 505.0so5 0 little effect on the presence of whipworm eggs amjim egg width 33533.533 5 10 amjumju m voucher speci- in feces of injected animals therefore animals mens were deposited with the university of received 8 d later a second set of two injec- alberta parasite collection s UAPCuapc1146411464 tions each of 0150.15 cc and delivering ca 2 mg and UAPCuapc1146511465 although we did not identi- ivermectin kg body mass control animals fy whipwhipwormsworms from D ordiiordliordia we are confident received the carrier august captures received they are T dipodomys the type host for T on each of two consecutive days an injection dipodomys is D ordnordiiordia and T dipodomys is of 0015olsois0.1515 cc volume delivering ca 2 mg known only from D ordiiordliordia and D micronsmicrops ivermectin kg body mass control animals whitaker et al 1993 received the carrier to control for possible prevalence in host species side effects of ivermectin half of the uninfect- in ed animals captured in august were also trichuris dipodomys occurred at substan- injected with a solution of ivermectin tially higher prevalence in D micronsmicrops than in 4 two days after each set of injections a D ordnordiiordia table 1 a result similar to that of posttreatmentposttreatment feeding trial was conducted grundmann 1957 we can speculate as to using techniques in 2 above only results of three possible explanations for this pattern the pretreatment feeding trials and feeding the first is that eggs produced by adult worms trials following the 2 mg ivermectin kg body in D micronsmicrops may become embryonated more mass injection will be presented below easily than those in D ordnordiiordia freshly produced fecal pellets of D micronsmicrops appear moister than RESULTS AND discussion those of D ordliordiiordia munger personal observa- tion probably because of the higher amount adult worms seven of each gender taken of green or leafy vegetation in the diet of D from a dipodomys micronsmicrops at our site were micronsmicrops if moisture is necessary for embryonaembryoma identified as trichuris dipodomys although tion of the eggs as is implied by parry 1968 76 GREAT BASIN naturalist volume 55

TABLETABLL 1 infection of two species of kangaroo rat with the TABLE 2 effects of whipworm infection on apparent nematode trichuris dipodomys dry matter digestibility ADMD standard errors are in parentheses figures on change between initial and final D microps D ordu microns feeding trials as well as sample sizes are in figure 1 see infected uninfected infected uninfected text for a description of treatments june trapping 10 5 5 39 treatment deinfectedDe infected uninfected august trapping 6 9 7 34 infected dipodomys micronsmicrops initial ADMD 956 965 955 0051 0029 0103 moister feces may lead to higher embryonationembryo nation final ADMD 961 950 953 rates and therefore higher prevalence among D 0039 0026 0052 micronsmicrops the second explanation is that social dipodomys ordnordiiordia and burrow use behavior may differ between initial ADMD 967 957 961 0107 0076 0022 these species for example perhaps D micronsmicrops final ADMD 955 958 957 individuals visit one another s burrows and 0034 0037 0014 thereby become exposed to parasite eggs at a substantially higher frequency than do D ordiiarchiorchiordia also D micronsmicrops inhabits a mound up to 2 m species experimental period july vs august in diameter while D ordnordiiordia inhabits less sub- was included as a blocking factor the depen- stantial individual holes studies of another dent variable in the analysis was proportional system of two species of kangaroo rats has change between pretreatment and posttreatposttreaty shown that the larger mound inhabiting D ment ADMD post preprepreprefrepre this measure spectabihsspectabilis uses its burrow system for pro- should be more sensitive than posttreatmentposttreatment longed periods while the smaller D mernardmerriamimerrimerrlmernarmami ADMD in expressing treatment effects because rotates among several burrows jones 1989 it takes account of initial differences in ADMD this latter behavior would tend to reduce among hosts reinfection of individuals it would be interest- although there were no statistically signifi- ing to see if behaviors differ similarly between cant main effects of treatment or species on D micronsmicropsrnicropsrnictopsrops and D ordu the third explanation ADMD there was a significant interaction is that resistance to infection may differ between these factors table 3 indicating that between these two host species the two host species differ in their response to effects on digestive efficiency treatment this difference between species was explored using a separate ANOVA for each apparent dry matter digestibility ADMD species which revealed that treatment with of millet seed was quite high 95 on aver- ivermectin had a significant effect on change age table 2 a figure comparable to that in ADMD in D micronsmimicropscrops but not in D ordiiordia found by schneberschrieber 1979 for granivorous table 4 fig 1 A tukeylukey s a posteriori multiple rodents injection of ivermectin did not sample test revealed that within D micronsmicropsmicrops appear to affect ADMD of animals uninfected the change in ADMD of the infecteddeinfectedde group by whipwormswhip worms an effect that might occur differed significantly from the change in through the removal of other symbionsymbiontsts or ADMD of both the infected group and the through some direct effect proportional uninfected group these results can be inter- change in ADMD X SE untreated preted as showing that the deinfected group 0 0043000430.0043 t 0 0035000350.0035 treated 0 0058000580.0058 0 0037000370.0037 had 191.9ig higher ADMD than the other two therefore in the following analyses all natu- groups rally uninfected animals are combined into one of interest is the lack of effect trichuris class causes in D ordiiordia this may be due to what the effect of whipworm removal on ADMD appears to be a higher intensity of infection was analyzed with a two way analysis of vari- more parasites per infected host in D micronsmimicropscrops ance ANOVA one factor analyzed was the fecal floats of D micronsmicrops in general contained treatment deinfected naturally infected but more eggs than did floats of D ordiiordia D treated with ivermectin vs infected naturally micronsmicrops XTC 254 SE 1152115.2 D ordiiordia X infected but not treated with ivermectin vs 63563.5 SE 21021.0 mann whitney U test U naturally uninfected the other factor was 79 FP 1.1 if fewer worms were present in D 199519951 WHIPWORMS IN KANGAROO RATS 77

TABLE 3 F values and probability values FP from three way analyses of variance on effects of species month and treatment deinfected infected or uninfected on apparent dry matter digestibility ADMD dependent variable proportion change initial ADMDADMD final A DMDADMD inm ADMD source df F P F P F FP treatment 2 15 86 72 49 47 63 species 1 46 50 82 37 33 57 treatment species 2 133 27 178 18 474 012 block month 1 51 48 911 003 00 95 error 77

in the lab sara murray and aaron munger helped in the field discussion with mary price TABLE 4 results from one way analyses of variance on the effect of treatment deinfected infected and uninfect- was helpful as were comments from anony- ed on change in dry matter digestibility in D micronsmicrops mous reviewers this research was supported and D ordu by an intramural faculty research grant from species source df MS F P boise state university D micronsmicrops CITED treatment 2 00106 464 019 literature error 27 000229 GRUNDMANN A W 1957 nematode parasites of mam- mals great D ordnordiiordia of the salt lake desert of utah journal ofofparasitology43parasitology 43 105 112 treatment 2 00034 121 31 W 1989 error 52 01442 JONES T dispersal distance and the range of nightly movements in merriam s kangaroo rats journal of mammalogy 70 27 34 KENAGY G J 1972 saltbush leaves excision ofhypersalineofhypersalinehypersaline tissue by a kangaroo rat science 178 1094 1096 ordiiordia the effect of eradicating those worms METTRICK D F 1980 the intestine as an environment would have been less apparent for hymenolepis diminutediminutadiminuta pages 281 356 in H P one might question the biological impor- arai ed biology of the tapeworm hymenolepis press tance of the slight albeit statistically significant diminutediminutadiminuta academic new york NY MUNGER J C AND W H KARASOV 1989 sublethal para- decrease by infec- in ADMD caused trichuris sites and host energy budgets tapeworm infection in tion munger and karasov 1989 showed an white footed micemlee ecology 70 904 921 effect of similar magnitude resulting hombomfromhrombrom tape- PARRY J E 1968 transmission studies of nematodes with worm infection hymenolepis citelli in white direct life histories in selected utah mammals peromyscus unpublished doctoral dissertation university of footed mice leucopusleucopus they utah salt lake city argued that hosts can easily compensate for PRITCHARD M AND G KRUSE 1982 the collection and such small effects by slight increases in food preservation of animal parasites university of consumption or decreases in expenditures or nebraska press lincoln 141 appp by changes in gut morphology mettrick 1980 READ C P 1956 trichuris dipodomys n sp from ord s kangaroo rat proceedings of the helminthological and concluded that such effects on ADMD are society of washington 2311923 ilg119 unlikely to affect host energy budgets or to SCHRIEBER R K 1979 coefficients of digestibility and translate through to population level effects caloric diet of rodents in the northern great basin the same conclusion is likely to apply to the desert journal of mammalogy 6041660 416 420 kangaroo rat whipworm system WHITAKER J 0 JR W J WRENN AND R E LEWIS 1993 parasites in H H genoways and J H brown eds biology of the heteromyidae american acknowledgments society of marnmalogistsmammalogists special publication 10 ZEVELOFF S I1 1988 mammals of the intermountain we thank alienallenailen shostak of the university of west university of utah press salt lake city 365 Albertalbertadss parasitology museum for measuring PP specimens of the parasite and for its identifica- received 27july27 lulyjuly 1993 tion kay kesling helped both in the field and accepted 20 june 1994 great basin naturalist ssi551 0 1995 appp 78 83 LOCAL distribution AND FORAGING BEHAVIOR OF THE SPOTTED BAT EUDERMA MACULATUM IN northwestern COLORADO AND ADJACENT UTAH

jay F storzistorz1storze

ABSTRACTABSFRACT this study investigated local distribution and foraging behavior of the spotted bat eudeemaeudermaEuderma maculatummaculatum in dinosaur national monument colorado utah by monitoring audible echolocation calls the occurrence of this species was verified in a variety of habitat types in canyon bottoms and other relatively low elevation sites indicating that the animals are widely distributed and locally common in the area foraging spotted bats concentrated flight activi- ty in the open air space above meadows and occasionally exploited near canopy habitat within 8 m of foliage bats began to forage shortly after dark and activity levels were relatively constant throughout the night foraging spotted bats attacked airborne prey every gis2159152 15 minmm on average consistent with published observations spotted bats maintained exclusive foraging areas distinct vocalizations indicating agonistic encounters occurred when a bat encroached on the foraging area of a conspecific

key words spotted bat eudermaeudeema maculatummaculatum colorado utah dinosaur national monument foraging habitat use attack rates echolocation

the spotted bat eudermaeudeemaEuderma maculatummaculatum is foraging habitat throughout the remainder of widely distributed across western north the geographic range of E maculatummaculatum america and apparently exists in low popula- the purpose of this study was 1 to investi- tion numbers throughout its range fenton et al gate local distribution of E maculatummaculatum by 1987 the species is rare in collections and monitoring echolocation calls 2 to identify viable populations have been documented in and describe foraging habitat and 3 to make only a few widely separated localities watkins a preliminary examination of spatial and tem- 1977 ofarrell 1981 findings presented here poral patterns of habitat use by spotted bats in and those of navo et al 1992 indicate that E the study area maculatummaculatum is locally common in canyon bot- toms and other low elevation sites in dinosaur METHODS national monument colorado utah and study was conducted in the canyon occurs throughout a diverse range of habitat this bottoms and other relatively low elevation types sites in dinosaur national monument log109 W population studies eg and fenton leonard 403 northwestern colorado and north- 1983 in south central british columbia have 403rn4031nrN eastern utah from 17 may to 9 june 1993 demonstrated that foraging spotted bats exhibit navo et al 1992 provided a description of considerable habitat specificity radiotrackingradiotracking the physiography and vegetation of dinosaur in this same area wai ping and fenton 1989 national monument in each study site where has demonstrated that individuals are faithful spotted bats occurred I1 monitored movement to specific sites over several consecutive nights patterns and foraging behavior by listening to however no clear association between forag- the low frequency 15 9 khz leonard and ing activity and any specific habitat conditions fenton 1984 echolocationecho location calls of this is apparent in british columbia spotted bats species which are clearly audible to the unaid- forage over clearings in ponderosa pine pinus ed human ear woodsworth et al 1981 ponderosa forests open fields and marshes As reported previously navo et al 1992 leonard and fenton 1983 wai ping and E maculatummaculatum is readily identifiable because it fenton 1989 there is little information about has the lowest frequency echolocation calls of

ienvironmentalnv nonmental population and organismic biology university of colorado boulder CO 80309033480309 0334 present address department of biology boston university boston MA 02215

78 199519951 SPOTTED BATS IN COLORADO UTAH 79

any bat species in the study area nyctinomops wood form a dense continuous canopy over macrotismamacrotincrotis and idionycterisIdionycteris phyllitisphylphyllotislotis also pro- much of the adjacent creek duce orientation sounds that are partly audible remaining locations consist of a moist mead- to humans with frequencies of 25 17 khz for ow hog canyon 1635 in open sagebrush N mamacrotismacrotincrotis fenton and bell 1981 and 24 12 shrublandsshrublands rainbow park 1488 in island khz for I1 phyllitisphylphyllotislotis simmons and ofarrell park 1512 in and a narrow canyon with thick 1977 these two species inhabit southern riparian vegetation bonesjonesuones hole 1585 in parts of the colorado plateau and the great sampling basin milner et al 1990 tumlison 1993 but methods neither is known to occur as far north as at all locations I1 remained at a single site dinosaur national monument extraextralimitallimital during each night of sampling by pacing from records ofofnoanN mamacrotismacrotincrotis notwithstanding milner a boxelderboxelder which served as a focal point of bat et al 1990 to further ensure correct call identi- foraging activity in echo park meadow I1 esti- ficationfication I1 referred to recordings of known E mated that calls of E maculatummaculatum were detect- maculatummaculatum calls I1 also visually identified free able at a distance of roughly 100 in therefore flying individuals based on conspicuous white the area sampled at each site is here defined venter and large ears at close range in the as the air space within a hemisphere of radius beam of a high intensity flashlight after locat- 100 in on several nights periods of high wind ing the animals by listening to orientation andor rain reduced this range of detectability sounds it should be noted that I1 phyllitisphylphyllotislotis is with an attendant underestimation of bat activ- often buff colored ventrally and therefore could ity furthermore sites differed slightly in levels be visually misidentified as E maculatummaculatum in of background noise from nearby streams the areas of sympatry amount of obstructive vegetation and various atmospheric conditions as sampling locations such relative humidity all of which affect the propagation of to investigate the ecological distribution of sound lawrence and simmons 1982 E maculatummaculatum I1 sampled 15 sites at 12 loca- study sites were situated either at the tions representative of common low elevation mouths of canyons or draws or in the middle habitat types in the area fig 1 riparian sites of open areas where movement patterns of bats jenny lind rock 1603 in echo park 1553 could best be assessed and the range of detect- in split mountain gorge riverbank and sand ability was maximized in locations character- bar 1439 in are characterized by wide chan- ized by expansive terrain open meadows or nels and reaches of calm water bounded by shrublandsshrublands I1 monitored two different sites steep sandstone cliffs isolated stands of box separated by 300 in on consecutive nights to elder acer negundo and cottonwood populus assess uniformity of activity levels over large fremonfremontiifremontiatii line the riverbanks along with thick- areas all sites were monitored from 2000 to ets of tamarisk tamarix sp 0200 h with the exception of echo park orchid draw 1484 in and red wash 1537 meadow which was monitored from 2000 to in are dry desert washes characterized by rab 0400 h for seven consecutive nights 19 26 bitbrushbitbrush chrysothamnus nauseosus sage- may to assess temporal patterns of foraging brush seriphidium dentatatritridentatatridentate greasewood activity sarcobatus venniculatusvermiculatus and shadscaleshad scale atri- at locations where I1 observed high levels of plex confertifoliaconfertifolia with tamarisk dominating foraging activity eg echo park meadow and drainage bottoms pool creek bat activity was quantified by echo park meadow 1548 in and pool creek timing the duration of individual foraging ses- 1635 in are both open meadows with domi- sions and recording the number of feeding nant ground cover of cheatcheatgrassgrass anisanthaAnisantha buzzes the increased rate of echolocation tectoriumtectorumtectorum various bunchbunchgrassesgrasses and isolated pulse repetition associated with attacks on air- clumps of boxelderboxelder echo park meadow en- borne prey griffin et al 1960 following compasses an area of ca 18 ha bounded by the leonard and fenton 1983 the occurrence of green river to the west and high 150 230 in feeding buzzes indicates foraging activity and sandstone cliffs on remaining sides the mead- a foraging session is defined as the time during ow at pool creek ca 8 ha is situated at the which a single spotted bat hunted continuous- mouth of a narrow canyon boxelderboxelder and cotton ly within the study site to permit comparison 80 GREAT BASIN naturalist volume 55

N A

0 5 km

400 35 N

jones hole echo I1 park jenny lind rock island I1 park 9 1 1 echo park rainbow poolpooi creek A meadow park

orchid red draw wash yampa river splitspilt mtnman gorge sandbarsandsandbagbar splitspilt mtnman gorge riverbank

U hog canyon UTAH COLORADO 1090logo W green L river I1 I1

fig 1 map showing sampling locations for monitoring activity of eudeemaeuderma maculatummaculatum in dinosaur national monument in late spring 1993 circles locations at which transient occurrences of commuting or foraging bats were recorded triangles foraging areas see text for details of relative levels of activity throughout the sessions of duration 3 min during which all night the time spent by spotted bats in the feeding buzzes produced by a single individ- study site was totaled for every 15 min period ual within the study site could be counted sampled sampling periods during which accurately heavy rain occurred were not considered at other locations where I1 observed only to assess spatial patterns of habitat use I1 transient occurrences of foraging or commut- described the foraging flights of spotted bats ing spotted bats activity was quantified by into a minicassetteminicassette recorder and noted flight recording the number of bat passes gensusensu patterns and use of available foraging space fenton 1970 per 15 min sampling period relative to a near canopy habitat zone within 8 m of tree canopiescanopies and an open area zone RESULTS AND discussion the clutter free air space over the open mead- ow these habitats correspond to habitat zones I1 observed spotted bats in 13 of 15 sites I1 and 4 respectively proposed by aldridge sampled table 1 at 8 of these locations I1 and rautenbach 1987 I1 recorded the dura- observed only commuting bats passes of com- tion of foraging activity occurring within each muting spotted bats occurred sporadically zone as well as the number of bats simultane- throughout the night at locations where two ously present within the study site and inter- separate sites were monitored on consecutive actions between them nights the number of passes remained fairly I1 recorded the number of feeding buzzes constant passesnightpassesnigbtpasse snight echo park 5 4 hog heard during each foraging session for each canyon 5 6 island park 6 10 and direc- night of observation at echo park meadow tions of travel appeared similar for bats on and pool creek for the purpose of calculating both nights attack rates feeding buzzesminbuzzesmin of foraging availability of cliff roosting sites has been spotted bats I1 considered only those foraging suggested as a limiting factor in the distribu 199519951 SPOTTED BATS IN COLORADO UTAH 81

TABLE 1 number ofpassesof passes ofeudennaofeiiderma maculatummaculatum per 15 minmm sampling period between 2000 and 0200 h at dinosaur national monument 16 may 8 june 1993 see text for general description of habitat types number of number sampling sampling location of nights periods n X range jenny lind rock 1 24 0 0 echo park 2 48 9 19 0 2 hog canyon 2 48 11 23 0 3 orchid draw 1 24 18 75 0 6 red wash 1 24 32 133 0 1 split mountain gorge sandbarsandsandbagbar 1 24 1 04 0 5 split mountain gorge riverbanknverbank 1 24 8 33 0 8 rainbow park 1 16 4 25 0 1 island park 2 48 16 33 0 2 jones hole 1 24 0 0 all sites 13 280 99 35 0 8 tion of E maculatummaculatum easterla 1973 the 2 and foraging sessions lasted 5485.48 t 2742.74 abundance of high cliffs in dinosaur national min n 187 at pool creek spotted bats monument as well as transient occurencesoccurrences of hunted within the study site for 6826.82 t 5035.03 commuting bats throughout a variety of wide- min out of every 15 min sampling period ly separated low elevation sites navo et al between 2100 and 0200 h n 525 min and 1992 this study suggests that suitable roost foraging sessions lasted 8978.97 t 8788.78 min n ing habitat is widespread throughout the area 30 these activity levels offer strong evidence however information about microclimate re- that open meadows represent important forag- quirementsquirements of this species is needed to fully ing habitat for E maculatummaculatum in this area assess actual availability of suitable roost sites comparatively low levels of activity were I1 observed foraging spotted bats by sight and recorded at riparian sites adjacent to echo sound at five locations three of which echo park meadow echo park jenny lind rock park orchid draw red wash involved only because no physiographic barriers are present transient occurrences of bats that were that might restrict accessibility to the bats it observed executing steep dives and other appears that open water courses do not repre- abrupt flight maneuvers coincident with feed- sent foraging areas of choice these observa- ing buzzes as they passed through the area I1 tions agree with those of leonard and fenton observed a single spotted bat foraging over a 1983 who reported that in british columbia sand and gravel bar at echo park but activity spotted bats foraged in forest clearings and levels at this location were lower than those open fields to the exclusion of a nearby river reported by navo et al 1992 who sampled the temporal pattern of foraging activity in this same site previously fairly high levels of dinosaur national monument is similar to activity occurred at orchid draw and red wash that reported from british columbia leonard table 1 and I1 heard three feeding buzzes at and fenton 1983 where spotted bats were each site however because spotted bats active throughout the night because radio apparently capture prey opportunistically tracking wai ping and fenton 1989 has while commuting to specific foraging sites demonstrated that individual spotted bats wai ping and fenton 1989 observations of hunt on the wing 300 min per night reports foraging bats passing through an area cannot of apparent peaks in nightly activity which be considered indicative of habitat preferences have been especially pronounced in mistnet at echo park meadow spotted bats first ting studies eg easterla 1973 are likely arti- arrived at the study site at 2123 h t 11 min facts related to the proximity of sampling sites mountain daylight time n 6 rain free to diurnal boostsroosts andor drinking sites evenings always after dark and remained at echo park meadow and pool creek for- active throughout the night fig 2 spotted aging spotted bats typically flew in large circu- bats foraged within the study site for 6226.22 t lar or elliptical orbits at heights of 10 30 in 2402.40 min out of every 15 min sampling period above the ground in 108881088.8 min of observa- between 2100 and 0400 h n 2490 min fig tion of foraging spotted bats at echo park 82 GREAT BASIN naturalistNATu ramstRAUST volume 55

foliage 1501150- 1 observed these bats gleaning prey from during this study observations of near canopy foraging contrast with those of some other workers eg wai ping and fenton 1989 who have reported that this species never attacked 10010 0 insects near foliage or any other type of sur- face information about individual variability Z in foraging behavior is needed before drawing 5.5 conclusions about variability between popula- 16 tions related to different ecological conditions both park pool 50 at echo meadow and creek there were 118 instances in which two or three E maculatummaculatum were present within the study site simultaneously leonard and fenton 1983 1984 estimated that spotted bats in british columbia maintain a distance of at least 00 0 0 0 0 0 L 50 in between adjoining foraging areas and 0 0 0 0 0 0 0 0 04 CO 0 N CJ cli CJ 0 0 suggested that this spacing is accomplished time h through a combination of mutual avoidance and active monitoring of encroachmentsencroachments by con fig 2 foifolforagingaging activity patterns of eudermaeudeema macula specifics this same system appears to be oper- park 19 may tumturnturm at echo meadow 26 1993 bars repre- ating at foraging areas in dinosaur national sent mean time SD spent by bats in the study site per 15 min sampling period from 2000 to 0400 h n 6 for monument consistent with observations of each 15 min period in the interval 2000 0345 h n 4 for leonard and fenton 1983 foraging spotted 0345 0400 h bats often produced agonistic vocalizations when the 50som m buffer zone was breached by an intruding bat such vocalizations sounded meadow 81581.5 of activity occurred over the qualitatively from open meadow which constituted roughly 85 different feeding buzzes and occurred only during close range of the site while 185iss18.5 of activity occurred encounters conspecificscon specifics about within 8 in of the foliage of fully leafed box between information elders at mid to upper canopy level such known individuals and resource availability is activity consisted of bats circling closely above needed to elucidate the role of agonistic inter- and around individual trees or isolated clumps actions in the foraging ecology of E maculamaculatumtum this study I1 heard a total of 247 feed- of trees I1 rarely observed bats within 050.5os in of during the canopy and I1 never observed hovering ing buzzes and never more than one per min flight or other evidence of foliage gleaning in from the same individual in a sample of 37 2908290.8 min of observation of foraging spotted foraging sessions spotted bats attacked an bats at pool creek all activity occurred over insect every 2152.15gis915 min on average 046604660.466 02940.294 the open meadow although a much larger attacksminattacks min range 0160.16olg 0940.94 n 152 feeding percentage of the study site area comprised buzzes these rates generally agree with val- canopiescanopies of boxelderboxelder and cottonwood than at ues reported in previous studies leonard and echo park meadow fenton 1983 wai ping and fenton 1989 fur- the predilection of E maculatummaculatum for forag- ther confirmation that this species attacks prey ing over open terrain in dinosaur national at a rate much lower than is typical of bats that monument agrees with the pattern observed forage from continuous flight barclay 1985 in previous studies eg woodsworth et al hickey and fenton 1990 1981 leonard and fenton 1983 low fre density of clutter in an environment im- quency echolocation calls and long intercall poses differential constraints on the maneuver- intervals suggest that spotted bats use a forag- ability and perceptual capacities of bats there- ing strategy based on long range prey detec- by determining the accessibility of different tion and high level flight simmons and stein habitats by influencing foraging efficiency neu- 1980 woodsworth et al 1981 barclay 1986 weiler 1984 aldridge and rautenbach 1987 this strategy likely is best suited to open areas fenton 1990 spotted bats appear to forage neuweiler 1984 although I1 never directly preferentially in open areas which may be 199519951 SPOTTED BATS IN COLORADO UTAH 83

related to the use of a long range foraging strat- 1990 the foraging behaviour and ecology of ani egy barclay 1986 and the ability to exploit mal eating bats canadian journal of zoology 68 edge situations may a 411 422 reflect measure of FENTON M B G behavioral flexibility AND P BELL 1981 recognition of in this regard because species of insectivorous bats by their echolocation spotted bats are obviously not greatly restrict- calls journal of mammalogy 62 233 243 ed in foraging habitat with regard to vegeta- FENTON M B D C TENNANT AND J WYSZECKI 1987 tion associations wai ping and 1989 using echolocation calls to measure the distribution fenton of bats navo et al 1992 the case of eudennaeddennaeuderma maculatummaculatum journal of structural features of the mammalogy 6814268 142 148 environment related to density of clutter may GRIFFINGBIFFIN D R F A WEBSTER AND C R MICHAEL 1960 be more predictive of habitat suitability and the echolocation of flying insects by bats animal the use of available foraging space however behavior 18 55 61 HICKEY M B information on individual variability is C AND M B FENTON 1990 foraging by needed red bats Lasi hs before drawing lasiurusurus boreaborealis do intraspecific chases conclusions about the foraging mean territoriality canadian journal of zoology 68 strategy ofthisorthisof this species 2477 24824822 LAWRENCE B D AND J A SIMMONS 1982 measurements acknowledgments of atmospheric attenuation at ultrasonic frequencies and the significance for echolocation by bats journal oftheodtheof the acoustical society ofamericaof america 71 585 590 I1 am grateful to D M K armstrong W LEONARD M L AND M B FENTON 1983 habitat use by navo M B fenton M L leonard M A spotted bats eudermaeudeemaEuderma maculamaculatumtum chiroptera bogan C E bock J A gore and G T skiba vespertilionidae roosting and foraging behavior for advice regarding study site locations and canadian journal of zoology 61 1487 14914911 1984 echolocation calls of eudeemaeudermaofeudenna maculamaculatumtum sampling methods I1 thank the personnel of chiroptera vespertilionidae use in orientation and dinosaur national monument and especially communication journal of mammalogyMamm alogy 65 122 126 S J petersburg for cooperation and for shar- MILNER J C JONES AND J K JONES JR 1990 nystinycti ing knowledge of the area critical comment nomops macrotismacrotinmacrotis mammalian species 351 1 4 NAVO K W A GORE SKIBA on the manuscript from D M armstrong M B J AND G T 1992 observa- tions on the the spotted bat eddennaeudermaeudenna maculatummaculatum in R M M in fenton timm J ofarrell and two northwestern colorado journal of mammalogy 73 anonymous reviewers was much appreciated 547 551 funding was provided by the undergraduate NEUWEILER G 1984 foraging echolocation and audi- research opportunities program university tion in bats naturwissenschaften 71 446 455 of colorado at boulder 0parrellFARRELLOFARRELL M J 1981 status report eudennaeudermaeddenna maculatummaculatum JAJ A allenalienailen united states fish and wildlife service office of endangered species 29 appp literature CITED SIMMONS J A AND M J OTARRELLOFARRELL 1977 echolocation by the long eared bat plecotus phyllitisphylphyllotislotis journal of ALDRIDCEALDRIDGE H D J N AND I1 L rautenbach 1987 comparative physiology 122 201 214 morphology echolocation and resource partitioning SIMMONS J A AND R A STEIN 1980 acoustic imaging in insectivorous bats journal of animal ecology 56 in bat sonar echolocation signals and the evolution 763 778 of echolocation journal of comparative physiology BARCLAY R M R 1985 long versus short range forag- 13561135 61 84 ing strategies ofboaryof hoary lasiurusLasiurus cinereus and silver TUMLISON R 1993 geographic variation in the lappet haired lasionyctenslasionycteris noctivagantnoctivagans bats and the con- eared bat Idioidionycterisidionyctensnycteris phyllitisphylphyllotislotis with descriptions of sequences for prey selection canadian journal of subspecies journal of mammalogy 74 412 424211 zoology 63 2507 2515 WAIPINGWAI PING V AND M B FENTON 1989 ecology of spot- 1986 the echolocation calls of hoary lasiurusLasiurus ted bats EudereuderrnaeudermaeudeemaEudermarna maculatummaculatum roosting and forag- cicinereuscinereousnereus and silver haired lasionycterislasionyctens noctiva ing behavior journal of mammalogy 70 617 622 gans bats as adaptations for long versus short range WATKINS L C 1977 eudermaeudeema maculatummaculatum mammalian foraging strategies and the consequences for prey species 77 1 4 selection canadian journal of zoology 64 woodsworth G C G P BELL AND M B FENTON 1981 2700 2705 observations of the echolocation feeding behavior EASTERLA D A 1973 ecology of the 18 species of and habitat use of eudertnaeudermaeudeema maculatummaculatum chiroptera chiroptera at big bend national park texas north- Vespertilionidvespertiliomdaevespertilionidacvespertilionidaeac in southcentralsouthccntralsouth central british columbia west missouri state university studies 349 1 165 canadian journal of zoology 59 1099 1102 FENTON M B 1970 A technique for monitoring bat activity with results obtained from different environ- received 7 february 1994 ments in southern ontario canadian journal of accepted 20 june 1994 zoology 48 47 51 gleatgreat basin naturalist ssi551 0 1995 appp 84 88 THE chrysothamnus ericameria connection asteraceae

loran C AnderAndersonsoni1

ABSTRACT the genus chrysothamnus asteraceae contains 16 species recently 4 species weiewelewere transferred to EricEncaericameriaencamertaameriaamerlameriameniamerta and the remaining 12 were left in chrysothamnus the remaining species are now transferred to EncericameriaEricencamenaameriaamerlaamena as E albida E deprescadepressadepressa E eremeremobiaezemobiaobia E gramineagramineaegraminea E filiaalifilifoliaftlifoliafolia formerly CG greeneigreenedgreenei E humilis E limfohalinifolialinifolia E molesta E pulpulchellachella E pulchelloides a fossil species E spatspathulataspathulatehulata E vaselivaseyivaseyt and E visciviscidifloradiflora section alignments are given and some intraspecificmfraspecificinfraspecificinfraspecific combinations are also made

key words chrysothamnus erleerieEncEricericameriaeneencamenaameriaamerlaamena rabbitbrushrabbitbrush nomenclature transfers

the asteraceae are a relatively young group thamousthamnus with woody elements of haplopappus and yet they have experienced rapid evolution and suggested that the asirisamiris ericameria into a great number of species one result is Macromacronemanema complex of haplopappus probably that many taxa appear more distant morpholog- should be included in chrysothamnus but ically phenotypically than they actually are given the state of knowledge at that time I1 de- genetically and conversely some taxa may ferred in 1990 nesom reorganized erlEriericamellaericameriaEricameriaamerlacameliacamella appear more closely related than they are as a genus to include asirisamiris and Macromacronemanema these situations have created havoc amongst recently based on occurrences of interintergenericgeneric taxonomists in their attempts to circumscribe hybrids anderson and reveal 1966 anderson genera this is particularly evident in the tribe 1970 and DNA data morgan and simpson Astereastereaeae in 1894 E L greene stated 1992 nesom and baird 1993 transferred four species of chrysothamnus into ericameriaEricameria in north america the Astereastereaeae are excessively numerous and no natural assemblage of plants has C nauseosus and C pattri of section nauseosi seemed to present such difficulties to the systema- and C paniculatus and C teretifoliusteretifolious of section tist and the widest conceivable diversities ofopinof opin- Punctpunctatipunctateati they continued to recognize chryso- ion as to the limits of genera have found expression thamnus as a distinct but smaller genus and undertaking to classify among botanists when gave for the two them arguments separating A problem in separating ericameriaEricameria and the situation continues a century later chrysothamnus gensusensu nesom and baird is the the genus haplopappus was thought to be occurrence of hybrids anderson 1970 1973 an unnatural polyphyletic assemblage by between C nauseosus their ericameriaEricameria and many eg shinners 1950 anderson 1966 C albieusalbidus their chrysothamnus after study- johnston 1970 turner and sanderson 1971 ing a specimen of only one of the three collec- clark 1977 urbatsch 1978 nevertheless tions involved nesom and baird 1993 deval- because there was no suitable taxonomic reor- ued the connection by stating that the plant ganization of the group I1 continued to describe in question is characteristic ofcofaof C nauseosus and new taxa in haplopappus anderson 1980a we identify it as C nauseosus finding no strong 1983b even though the species would probably reason to implicate C albieusalbidus in its parentage be placed in some other genera at a later date they stated that achenes of C albieusalbidus are linear recently additional data have contributed to a and consistently producing 10 slightly raised clearer understanding of the relationships in nerves whereas those of C nauseosus are nar- this and related groups morgan and simpson rowly obovate with 5 7 nerves actually ach 1992 and several genera have been recog- enes of both species can be characterized as nized for north american haplopappi being narrowly cylindrical the number of in a 1976 presentation at national meet- vascular bundles associated with the nerves ings I1 discussed the close affinity of chryso in the achenes averages approximately 7 and

idepartinent of biological science florida state university tallahassee FL 32306204332306 2043

84 199519951 chrysothamnus ericameria connection 85 ranges from 5 to 10 but mostly 6 8 in ash clements 1923 used nerve number to distin- meadows for C albieusalbidus anderson 1970 guish C nauseosus sspasp gravegraveolentgraveolensolens from sppapp 1973 whereas achene bundle number in C conconsimilissimilis so the character is variable even nauseosus ranges from 5 to 12 but is restrict- within a species many of the latter group ed to 5 for those in ash meadows such as C albidus C greeneigreenedgreenei and some forms the interspecific hybrid examined by nesom of C viscidiflorusviscidiflorus appear to have 1 nerved and baird beatley 11894 KSQKSC was studied leaves actually all species of ericameriaEricameria and anatomically by anderson 1973 its hybridity chrysothamnus have trilacunar 3 trace nodal is indicated by low pollen fertility and by mor- anatomy personal observation thus the char- phphologicalological intermediacy between the two acter of I1 versus 3 nerves is a matter of per- species in its revoluterevolute leaves in vascular bundle cepceptiontion not of fact 2 leaf margins ciliate in number in the ovary wall in corolla lobe the former and never in the latter but C length and in anther appendage length it has albidus C ereeremobiusmobius and C viscidiflorusviscidiflorus secretory canals in the ovary wall and glandular subsp planifoliusplaniplanifoliousfolius of the former have entire chomestritrichomes on the corolla tube like C nauseanauseo leaf margins also in that group C pulpulchelluschellus sus unlike C albidus and ovary wall unlike subsp pulpulchelluschellus has entire leaf margins C nauseosus like C albidus further proge- whereas subsp baileyibaileyy has ciliate leaf margins ny from one of my C albidusalbieus garden plants and some populations of C graminousgramineusgramineus and C also has low pollen fertility and looks interme- vaselivaseyi have entire leaf margins but others do diate between its seed parent and C nauseanauseo not ericameriaEricameria gensusensu nesom 1990 has sev- sus anderson 1970 its flowers have secreto- eral species that have leaves with ciliate leaf ry canals in the ovary wall and glandular tri margins fairly prominent in E coopericooperia and chomes on the corolla tube but lack glandular less so in several other species eg E cervina chomestritrichomes on the ovary wall those three fea- E nana E ophitidis and E zizlonisonis 3 Cocorollasrollas tures are characteristic of C nauseosus but not more or less abruptly broadened from the of C albieusalbidus the seed parent clearly suggest- tube into the throat with long curvingrecursingrecurvingre or ing hybridity if existence of interspecific coiling lobes in the former and corollas tubu- hybrids is used to justify transferring C nau lar with short erect or spreading lobes in the selsusseosus to ericameriaEric ameria then this feature also latter but cocorollasrollas of C spathulatus of the argues for bringing the remainder of chryso- latter have relatively broad tubes that lack thamnus into ericameriaEricameria noticeably flaring throats C humilis of the the warranted position of chrysothamnus former has tubular cocorollasrollas with short erect teretifoliusteretifolious in ericameriaEricameria is taken by nesom lobes anderson 1964 226 and C nauseosus and baird 1993 80 because like many eri sspasp ceruminosus of the latter has cocorollasrollas caberiacameria species gensusensu striatumstrictumstrictum that species that are abruptly broadened from the tube has the tendency for the resiniferous ducts into the throat with long spreading lobes 4 that are almost always distinctly associated style appendage collecting hairs merely papil- with the phyllary midvein to expand near the late in the former whereas they are long and apex of the phyllary this characteristic also sweeping in the latter but C bidusalalbieusalbidus C occurs in many species of chrysothamnus molestusmomodestuslestus C pulpulchelluschellus and certain popula- gensusensu nesom and baird as illustrated for C tions of C viscidiflorusviscidiflorus all of the former have vaselivaseyi anderson 1963 660 and cannot be style appendages with moderately long sweep- used to distinguish the two groups I1 have ob- ing hairs diversity in collecting hairs is greater served adjacent populations of C viscidiflorusviscidiflorus in chrysothamnus gensusensu anderson 1986 than subsp puberuluspubpuberulouserulus in which plants of one had nesom and baird 1993 imply and does not prominently enlarged resin ducts at the phyl- fall into two groups collecting hair length lary tips and plants of the other did not may be correlated with other floral features with the transferral of four species from namely the cocorollasrollas style lengths and pollen chrysothamnus to ericameriaEricameria nesom and volumes of the former group anderson 1966 baird 1993 separate the two newly struc- are generally smaller than those of the latter tured genera with six criteria 1 leaves 3 5 involucral bractsbraats in vertical files in the for- nerved for chrysothamnus and 1 nerved for mer caveat noted and usually not in vertical ericameriaEricameria but many of their ericameriaEricameria files in the latter but perhaps the most have prominently 3 nerved leaves hall and strongly aligned bractsbraats occur in C nauseosus 86 GREAT BASIN naturalist volume 55 sspasp arenariusavenariusatharharenanusarenanusnariusnafius of the latter 6 achenes glan- 6 ericameriaEricameria humilis E greene L C anders dular with nonresinousnonnonresmousresinous nerves in the former comb nov basionymBasi onym chrysothamnus humilis E and eglandular with duplex hairs and greene Pitpittoniapitroniatonia 3 24 1896 greene resinous nerves in the latter but only five 7 ericameriaEricameria linifolialinifolia E L C anders species of the former have glandular achenes comb nov basionymBasionym chrysothamnus linilinifoliushnifohusfolius E greene Pitpittoniapitroniatonia 3 24 1896 in some they are hidden by duplex hairs and 8 ericameriaEricameria molesta blake C the other seven do not having either glabrous L anders comb nov basionymBasionym chrysothamnus viscidi achenes or achenes with duplex hairs exclu- viscidiflorusflorus var momolestusmodestuslestus blake J wash acad sci 30 368 sively 1970 1983a and many have anderson 1940 chrysothamnus molestusmodestusmo lestus blake L C resin canals associated with the bundles of the anders Madromadronofio 17 222 1964 achenes admittedly fewer than in those of the 9aaa ericameriaEricameria pulpulchellachella gray L C anders latter but well developed in C momolestusmodestuslestus of the comb nov basionymBasionym Linolinosynslinosyrissyris pulpulchellachella A gray former also C paniculatuspamculatus of the latter group- PIpl wrghtwnghtwright smiths contr knowl 35 96 1856 ing lacks resin canals in its achenes anderson bigeloviaBigelovia pulpulchellachella A gray A gray proc amer 1970 none of these six sets of characteristics acad arts 8 643 1873 chrysothamnus pulpulchelluschellus can be used to consistently separate the two A gray E greene erythea 3 107 1895 groups ab9b ericameriaEricameria pulpulchellachella subsp baileyibaileyy woot clearly chrysothamnus gensusensu anderson & standl L C anders comb nov basionymBasionym 1986 not nesom and baird 1993 is fairly chrysothamnus baileyibaileyy woot & standl contr USU S homogeneous and should not be dismem- nati heibhelbherb IS18 181 1913 ac bered if some are to go into ericameriaEncEricencamenaameriaamerlaamena and 9c ericameriaEricameria pulpulchellachella subsp pulpulchellachella var DNA data suggest they should then all should elatiorelation standl L C anders comb nov basionymBasionym chrysothamnus elatiorelation standl proc biolabiol soc wash go into ericameriaEncEricencamenaameriaamena therefore the remalremailemalremainilremainingnilnii g 26 118 1913 this variety with uniformly pubes- 12 of chrysothamnus are transferred to species cent leaves occurs sporadically in a few populations Ericericameriaertcarneriaameriaamerla and new combinations are made of the typically glabrous leaved subspecies pulchelpaulchel here lus and does not warrant a higher taxonomic status 1 ericameriaEricameria albida M E jones ex A gray than this quadnnomialquadrinomial affords L C anders comb nov basionymBasionym Bigbigeloviaelovia albi- 10 ericameriaEricameria spatspathulataspathulatehulata L C anders L C da M E jones ex A gray proc amer acad arts anders comb nov basionymBasionym chrysothamnus 17 209 1882 chrysothamnus albieusalbidus M E jones spathulatus L C anders madrono 17 226 1964 ex A giacegiayeglayegray E greene erythea 3 107 1895 chrysothamnus viscidiflorusviscidiflorus var ludens shinners 2 ericameriaEricameria depressadeprescadepressa nutt L C anders sida 1 374 1964 comb nov Basibasionymonym chrysothamnus depressesdepressusdepres sus 11 ericameriaEricameria vaselivaseyi A gray L C anders nutt proc acad nat sci philadelphia 4 19 comb nov basionymBasi onym bigeloviaBigelovia vaselivaseyi A gray proc 1948 Linolinosynslinosyrissyris depressadeprescadepressa nutt torr in stigreavesStigreaves amer acad arts 12 58 1876 chrysothamnus keptrept expedexpede zuni & colorado rivers 161 1853 vaselivaseyi A gray E greene erythea 3 96 1895 bigeloviaBigelovia depressadeprescadepressa nutt A gray proc amer chrysothamnus bakenbakeri E greene Pitpittoniapitroniatonia 4 152 acad arts 8 643 1873 1900 3 ericameriaEric ameria eremobiaeremezemobiaobia L C anders 12a ericameriaEric ameria viscidifloraviscidiflora hook L C L C anders comb nov basionymBasi onym chrysothamnus anders comb nov basionymBasionym crinttanacrinitariaCrinttana viscidiflo ereeremobiuseremohiusmobius L C anders Britbnttoniabrittoniabrittaniatonia 35 23 1983 ra hook fl bor am 2 24 1834 chrysothamnus 4 ericameriaEric ameria gramineagragramineaeminea H M hall L C viscidiflorusviscidiflorus hook nutt trans amer philos soc anders comb nov basionymBasionym chrysothamnus II11 7 324 1840 Bigbigeloviaelovia dougladougiadouglasndouglasbdouglasiisiisilsn A gray proc graminousgramineusgramineus H M hall muhlenbergia 2 342 1916 amer acad arts 8 645 1873 chrysothamnus dou radonaraconaPetpetradonapetradoriaPetr adoria discoidea L C anders trans kansas glasnglasiigladii A gray clements & clements rocky mtnman acad sci 66 676 1964 fis 226 1914 chrysothamnus pumilus nutt 5 ericameriaEricameria filifoliafilifolia redbrydb L C anders trans amer philos soc II11 7 323 1840 Linolmosynslinosyrissyris comb nov baslBasibasionymbcisionymonym chrysothamnus filiaallfihfoliusfilifoliusfolius serrserrulateserrulataulata torr stansbury rep 1 389 1851 chryso- rydh bull toireybottorrey bot club 28 503 1901 Bigbigeloviaelovia thamnus serruserrulatuslatus torr redbrydb bull torrey bot greeneigreened A gray proc amer acad arts 11 75 1876 club 33 152 1906 chrysothamnus tortitortifoliusfolius E not ericameriaEricameria greeneigreened A gray nesom greene fl fran 368 1897 chrysothamnus leuco chrysothamnus greeneigreened A clayglaygray E greene cladus E greene Pitpittoniapitroniatonia 5 59 1902 chrysotham- erythea 3 94 1895 chrysothamnus pumilus var nus stenostenolepislepis redbrydb bull torrey bot club 37 acuminatus A nels bot gaz 28 376 1899 1311910131 1910 chrysothamnus scopariusscopahuseanuspanus redbrydb bull torrey bot 12b ericameriaEric ameria viscidifloraviscidiflora subsp viscidifloraviscidiflora club 28 504 1901 chrysothamnus lanianlancinuslaricinuslaricinusficinuscinus E var latifolialatifolia D C eaton L C anders comb greene Pitpittorapittomapittoniatonia 5 110 1903 nov basionymBasi onym Linolinosyrislinosynssyris visciviscidifloradiflora var latifolialatifolia 199511995 chrmthamnuschrsothamnuschrysothamnus ericameria connection 87

D C eaton bot king eiplexpl 1571871157.1871 chrysotham- anders comb nov basionymBasi onym chrysothamnus nus latifoliuslatifolius D C eaton redbrydb bull torrey bot pulchelloides L C anders great basin club 3333152190633152.1906152 1906 naturalist 40 351 1980 12c ericameriaEricameria viscidifloraviscidiflora subsp viscidifloraviscidiflora nesom and baird 1993 suggest the chryso- stenostenophyllaphylla A gray L C comb nov varvan anders thamnus taxa that I1 have just transferred to basionymbasionyrnBasionym bigeloviaBigelovia douglasiisii var stenostenophyllaphylla A basl dougla ericameriaEricameria should be placed in a restructured gray proc amer acad arts 8 646 1873 chryso- odoria thamnus stenostenophyllusstenophyllousphyllus A gray E greene erythea genus to include elements of hesperodoriaHesper petradoria 3 94 1895 these quadrinoquadrinornialsquadrinomialsmials 12b and 12c Petradoria and vancleveaVanclevea they conclude that identify sporadic but rather distinctive morpho chloroplast DNA data suh 1989 show petra types that occur in the northern regions of this sub- doria to be integrally related to the solidago species for conceptual distinction between sub- lineage and far removed from ericameriaEricameria species and variety see anderson 1980b however they note that neither suh 1989 12d ericameriaEricameria viscidifloraviscidiflora subsp axillarisaxillaris nor morgan and simpson 1992 sampled any keck L C anders comb nov baslbasionymbasionyrnBasionym chryso- taxa of chrysothamnus sensugensu nesom and baird thamnus axillarisaxillaris keck aliso 4 104 1958 these taxa need DNA profiles determined subsp lanceolatalanceolatelanceolata 12e ericameriaEricameria viscidifloraviscidiflora because they certainly do not make a morpho- nutt L C anders comb nov basionymBasionym chryso- logically compatible grouping with Petrpetradoriaadoria thamnus lanceolalanceolatustus nutt trans amer philos soc Van example pettpetradoflapetradoria II11 7 324 1840 chrysothamnus elegansdelegans E greene or vancleveaclevea for bettPetradoria anderson erythea 3 94 1895 bigeloviaBigelovia dougladouglasiisii var spathespathu 1963 has radiate heads with disk flowers that lata jones proc calif acad sci II11 5 690 1895 lack stigmatic areas on the style branches and chrysothamnus glaucus A nels bull torrey bot have abortive ovaries and Vanvancleveaclevea anderson club 25 377 1898 chrysothamnus pumilus var and weberg 1974 has large turbinate heads latuslotusiotus A nels bot gaz 54 413 1912 with many phyllariesphyllaries many flowers and a 121 ericameriaEricameria viscidifloraviscidiflora subsp planiplanifoliafolia L tardily deciduous pappus of paleapaleacouspaleaceouscous awnsagns C anders L C anders comb nov basionymBasi onym none of these conditions are found in chryso- chrysothamnus viscidiflorusviscidiflorus subsp planiplanifoliusplanifoliousfolius L C thamnus sensugensu nesom and baird the cohe- anders madronofionionno 17 223 1964 Madro sivenesssiveness of chrysothamnus sensugensu anderson is 129 ericameriaEricameria viscidifloraviscidiflora subsp pupuberulaberula C spathulatus D C eaton L C anders comb nov baslbasionymbasionyrnBasionym further illustrated in that twigs linosyrisLino syris viscidifloraviscidiflora var pupubemlapuberulaberula D C eaton bot emit odor similar to that of C nauseosus king eiplexpl 158 1871 chrysothamnus pubpuberuluspuberulouserulus anderson 1964 227 D C eaton E greene erythea 3 93 1895 two alternate taxonomies are now available chrysothamnus marianus redbrydb bull torrey bot one for chrysothamnus as a genus anderson club 37 131 19191010 1986 or as a component of ericameriaEric ameria nesom and baird 1993 and here both are are the following sections in ericameriaEric ameria preferable to merging some elements of proposed to accommodate these species trans- chrysothamnus with Petrpetradoriaadoria or vancleveaVanclevea fers ericameriaEricameria section chrysothamnus A gray L C anders comb nov basionymBasionym bige acknowledgments lovia section chrysothamnus A gray proc amer acad arts 8 641 1873 this section in- james reveal and arnold tiehm offered cludes E albida E filifilifoliafolia E humilis E lini constructive comments on the manuscript folia E spatspathulatespathulatahulata and E viscidifloraviscidiflora ericadericam eria section craminigradinigramini L C anders L C literature CITED anders comb nov basionymBasionym chrysothamnus compos- proc ANDERSON L C 1963 studies on Petrpetradoriaadoria section craminigradinigramini L C anders lympsymp itae anatomy cytology and taxonomy transactions biology of artemisia and chrysothamnus 29 of the kansas academy of science 66 632 684 1986 this section includes E eremezemobiaeremobiaobia and E 1964 taxonomic notes on the chrysothamnus visdosuisdis gramineagragramineaeminea ericameriaEricameria section Pulpulchellichelli hall & cidiflorus complex Astereastereaeae compositae Madromadronofio 1722217 222 227 C comb Basi clements L anders nov basionymonym 1966 cytotaxonomic studies in chrysothamnus chrysothamnus section Pulpulchellichelli hall & Astereastereaeae compositae american journal botany 53 clements carnegie instdinst publ 326 175 193 204 211 1923 this section includes E depressadeprescadepressa E 1970 floral anatomy of chrysothamnus Astereastereaeae 3 466 503 molesta E pulpulchellachella and E vaseyi compositae sidasidaksida3 vaseli 1973 unique chrysothamnus hybridizations in ash additionally there is a fossil species eri meadows nevada bulletin of the torrey botanical caberiacameria pulchelloides L C anders L C club 100 171 177 88 GREAT BASIN naturalist volume 55

1980a haplopappus alpinusaepinus asteraceae a new JOHNSTON M C 1970 compositae pages 1523 1744 in species from nevada great basin naturalist 40 D S correll and M C johnston manual of the vas- 73 77 cular plants of texas texas research foundation 1980b identity of narrow leaved chrysothamnus rennertsrennertxrenner TX 1881 appp viscidiflorusviscidiflorus asteraceae great basin naturalist 40 MORGAN D R AND B B SIMPSON 1992 A systematic 117 120 study of machaeranthera asteracaee and related 1983a chrysothamnus eteeremobius asteraceae a groups using restriction site analysis of chloroplast dewnew species from nevada Britbnttoniabrittaniabrittoniatonia 35 23 27 DNA systematic botany 17 511 531 1983b haplopappus crispus and H zionts aster- NESOM G L 1990 nomenclatural summary of aceae new species from utah great basin naturalist ericameriaEncEricencamenaameilaamellaameriaamenaomena asteraceae Astereastereaeae with the inclusion 4335843 358 364 of haplopappushaplopapptis sects asins and Macromacronemanema phytol 1986 an overview of the genus chrysothamnus ogia68ogiaogla 68 144 155 asteraceae pages 29 45 in E D mcarthur and B L NESOM G L AND C I1 BAIRD 1993 completion of welsh eds proceedings symposium on the biol- Ericencaineriaericameriaamerlaameria asteraceae Astereastereaeae diminution of ogy Artemisiaofartemisiaof and chrysothamnus USDA forest chrysothamnus phytologiaphytologicPhytologia 75 74 93 service intermountain research station ogden SHINNERS L H 1950 notes on texas compositae IV V UT 398 appp field and laboratory 18 2542954225 42 ANDERSON L C AND J L REVEAL 1966 chrysothamnus SUH Y 1989 phylogenetic studies of north american bolanboianbolandenderidefidefl an intergenencintergenericintergenericgenerle hybrid Madromadronofio 18 Astereastereaea&tereaeae asteraceae based on chloroplast DNA 225 233 unpublished doctoral dissertation university of ANDERSON L C AND P S WEBERG 1974 the anatomy texas austin and taxonomy of vancleveaVanclevea asteraceae great basin TURNER B L AND S SANDERSON 1971 natural hybridi- naturalist 34 151 160 zation between the composite generdmachaerangenera machaeran CLARK W D 1977 Chemochemosystematicssystematics of the genus thera and haplopappushaplopappits sec Blepharablepharadondon american hazardiaHazardia compositae journal of the arizona journal of botany 58 467 academy of science 12 16 URBATSCH L E 1978 the chihuahuanchibuahuan desert species of GREENE E L 1894 observations on the compositae IV Ericencaineriaericameriaamerlaameria compositae Astereastereaeae sida 7 298 303 erytheaerythea22 53 60 HALL H M AND F E CLEMENTS 1923 the phylogenet- received 7 feburaryFeburary 1994 ic method in taxonomy the north american species accepted 2 june 1994 of artemisia chrysothamnus andatnplexand atriplex carnegie institute publication 326 1 355 great basin naturalist ssi551 0 1995 appp 89 91 reproductive BEHAVIOR IN MERRIAMS CHIPMUNK TAMIAS MERRIAMI

stephen B ComptocomptonlComptoncompton1nl1 and J R Callahan2

key words tamias eutamiaseutaimasEuEutatamiasimas neotamiasNeotamias chipmunk copulation offiolfactionaction

the literature contains little information the observation was made I1 april 1994 in a regarding mating chases and copulation in any wooded residential area in idyllwildIdyllwild riverside of the western chipmunks tamias subgenus county CA elevation 1590 m between 1000 neotamiasNeotamias callahan 1981 reported mating and 1130 h the habitat is mixed conifer forest chases for Merrmerriamlanisianiss T merrimerriamiami and dusky dominated by incense cedar calocedrusCalo cedrus chipmunks T obscuresobscurus but noted unpub- decurrentdecurrensdecurrens yellow pine pinus ponderosa live lished that both copulacopulatingpopulatingting pairs were partly oak quercus chrysolepischrysolepis and black oak Q concealed by foliage larson 1981 described kelloggkelloggiiii with a sparse understory of chapar- two copulationspopulations for merriamMerrianss chipmunk but ral shrubs when the observer arrived at 1000 a careful reading suggests that one of these h six to seven merriaasmerriamsMerrmerrlMerriaasiamslamsams chipmunks many of was a mounting attempt by an immature male them males were running over around and and the other was observed from a consider- through a large woodpile while performing able distance best and granai 1994 found no conspicuous leaping maneuvers no agonistic references on this subject other than callahan interaction was observed it was not possible at 1981 and larson 1981 this stage to identify the females or to tell in there has been some speculation and dif- which direction the chase was headed the ference of opinion regarding reproductive iso- overall effect was somewhere between a lating mechanisms in parapatricparapatric species of sciurus likeilke mating chase eg thompson western chipmunks blankenship and brand 1977 in which several males follow one 1987 reported differences in vocal behavior female and a lek involving male display the between tamias merrimerriamiami and T obscuresobscurus at chase covered an area 13 15 m in diameter black mountain riverside county CA and but centered on the woodpile and a nearby noted a possible role in reproductive isolation heap of smaller pine branches one of us JRCJRQ however had previously con- after about 20 min one chipmunk later ducted a more extensive study of vocal behavior identified as female ran up on one of the piled in these two cryptic species at black mountain branches A second chipmunk approached and from 1975 to 1980 vocalizing individuals were they ran around for a few seconds the female collected to confirm species identity and sono stopped on a branch and the other chipmunk grams were prepared and measured yet no a male ran up beside her his entire right side statistically significant vocal differences were was in contact with her left side for about I1 sec found callahan 1981 and in preparation during which he made a nuzzling motion with ecological olfactory and mechanical barriers the right side of his face on the rear left portion to hybridization also have been suggested of her face the expected nasalgenitalnasal genital contact callahan 1977 1981 patterson 1984 these was not observed but the pair had been out of hypotheses cannot be tested without more sight for a short time previously and this could data on chipmunk reproductive behavior have occurred the female then jumped to accordingly this note provides the first another branch which was 5 cm in diameter detailed description of western chipmunk cop- and 20 cm above the ground sloping at a 25 ulation that has been published to the best of angle so that the female was facing downhill our knowledge comparative data for other copulation then occurred only 2 m from the western chipmunk species would be of interest observer who was inside a parked vehicle

liendlsendend reprint requests to box 3140 hemet CA 92546 2museum2museurn of southwestern biology university of new mexico albuquerque NM 87131

89 90 GREAT BASIN naturalist volume 55

the female crouched on the branch as if breeding season scent marking is prevalent in resting the male jumped to that branch and sciundssciurids but usually this means marking the quickly grasped the female from the rear with ground or a branch not marking another ani- his forelimbsforelimbs around her pectoral region the mal the nuzzling and brushing behavior female s tail was deflected to the side and of the male Merberriansmerriansmernamrianss chipmunk before and slightly raised and the male s tail was extend- during copulation suggests that he was scent ed to the rear copulation consisted of four marking the female series of pelvic thrusts each series except the conspecific marking has been described for last lasted about 4 seesec and comprised an esti- various mammals such as rabbits mykytowyczmykytowyez mated 12 24 thrusts at a rate of 3 6 per sec 1965 but not for sciundssciurids gurnell 1987 each series of rapid thrusts was followed by a describes face wiping behavior by various short resting period during which the male tree squirrels but only in the context of sub- stopped thrusting and brushed his face strate marking and in Paraparaxerusxerus self groom mouth nose and chin from side to side 2 4 ing his description of copulation in sciurus times against the back of the female s neck and tamiasciurusTamiasciurus says nothing about the male the fourth and last series of thrusts was short- marking the female with reference to olfacto- er than the first three the male then released ry communication in ground squirrels halpin his grip on the female dismounted and ran 1984 wrote that there is no experimental off into the woodpile the female who had evidence that conspecific marking actually remained motionless during the act remained occurs among the sciundssciurids on the branch about 1 sec and then also ran to our observation indicates that conspecific the woodpile the entire copulation lasted marking does occur in merrlMerriaernammerriardsmernamaidsardss chipmunk as about 18 sec a component of reproductive behavior without although several male chipmunks partici- experimental data it is not possible to deter- pated in the chase none of them approached mine the significance of this marking pair the copulatingcopulapopulatingting pair no chipmunks were heard bonding comes to mind but there is no good vocalizing during the mating chase or copula- evidence of longtermlong term pair bonding in tion we did not note any pre mating vocal dis- merriam s or any other species of western play or lockrufelockhufeLockrufe by the estrous female chipmunk despite many years of field obser- callahan 1981 but we were not present on vation other possibilities include the follow- the days when the display if any would have ing 1 the marking induces some required taken place the tamias vocal display has physiological state in the female 2 the mark- been reported for a few species of chipmunks ing tells other males that the female has by callahan 1981 blake 1992 and others already mated before the copulation plug forms and the message becomes redundant it is not clear whether this vocal display is universal or occurs only at low population or 3 the marking reinforces a short term pair densities when the female benefits by attract- bond to ensure that subsequent copulationspopulations if ing more distant males any on the day of estrus will be with the same no further copulationspopulations were seen but as male larson 1981 indicated that the same female copulatespopulates more than many as seven male chipmunks continued to estrous sometimes copulates run around the same woodpile for another hour once mortality from all is higher for male the level of activity appeared to decrease and causes is there were none of the prodigious leaps seen than for female chipmunks smith 1978 per- haps due in to the dispersal and exposure earlier group then gradually dispersed as in part the associated breeding individuals headed for an adjacent area where with the season callahan 1981 the risk of predation other chipmunks were heard giving occasionoccasional after incurring 11 and expending considerable on the chipper vocalizations not the long series of energy mating chase it should be to the male s advan- chips that characterizes the LockrylockrufelockhufelockryfeLockrufetuperupefe tage to that his genes are passed to all the behavior described above suggests that ensure the female s offspring of the season scent glands play a key role in reproductive be- havior of this species 1981 and others larson CITED have noted that male chipmunks have scent literature glands near the chin and angle of the jaw oral BEST T L AND N J GRANAIGRANAL 1994 tamias mernamimerriamimetrimerriami glands that become enlarged during the mammalian species 476 1 9 199519951 NOTES 91

BLAKE B H 1992 estrous calls in captive asian chip- LARSON E A 1981 merrlMerrimerriaasaernammernamaass chipmunk on palo escrito munks tamiastarmas sibincussibsibiricussibisiblficusricusincus journal of mammalogy 73 in the santa lucia mountains of california part I1 597 603 regimen with recorded episodes of naturalistic be- blankenship D J AND L R BRAND 1987 geographic havior enid A larsonwacoba press big pine CA variation in vocalizations of california chipmunks mykytowyczmykyrowycz R 1965 further observations on the terri- tamias obscurusobscures and T mernamimerriamimerrimerrl ami bulletin of the torial function and histology of the mandibularsubmandibularsub southern california academy of sciences 86 cutaneous chin glands in the rabbit oryctolagus 126 135 cuniculus L animal behaviour 13 400 412 CALLAHAN J R 1977 diagnosis of eutamiasEutamias obscuresobscurus PATTERSON B D 1984 geographic variation and taxonomy rodentia sciuridae journal of mammalogy 58 of colorado and hopi chipmunks genus eutamiasEutamias 188 201 journal of mammalogy 65 442 456 CALLAHAN J R 1981 vocal solicitation and parental SMITH S FE 1978 alarm calls their origin and use in investment in female eutamiasEutamias american naturalist eutamiasEutamias sonomae journal of mammalogy 59 118872118 872 875 888 893 GURNELL J 1987 the natural history of squirrels facts THOMPSON D C 1977 reproductive behavior of the grey on file new york squirrel canadian journal of zoology 55 1176 1184 HALPIN Z T 1984 the role of olfactory communication in the social systems of ground dwelling sciuridssciunds received I111I1 april 1994 pages 201 225 in J 0 muriemurlemune and G R michener accepted 19 october 1994 eds the biology of ground dwelling squirrels university of nebraska press lincoln great basin naturalist ssi551 0 1995 appp 92 94 additional RECORDS OF FLEAS siphonaptera FROM UTAH

james R kuceralkucera1kucerak

key words siphonaptera fleas utah megabothnsmegabothris asio megamegacolpuscolpus euhoplopsyllus glaciaglacialisglacialistlis lynx

subsequent to the important work of stark nearctopsylla brooksi rothschild 1904 1959 few publications have given flea collec- utah co provo 21 august 1951 1516 1 Y tion records from utah these include jellison BYU 1366 & 1365 ex mustela fredatafrenatafrenata coll and senger 1976 and kucera and haas 1992 D brown provo canyon 16 june 1959 2s2asct 6 but most effort in this area has been that of 7 Y Y ex spilogale gracigracilisfishis coll D E beck egoscue 196619761977198819891966 1976 1977 1988 1989 this species was previously known in utah herein is presented information for 10 from a single collection in sevier county of siphonaptera for utah A number of species stark 1959 it is usually found on weasels records obtained from the flea important were mustela sppapp collection at the monte L bean MLB life science museum brigham young university nearctopsylla hyrtaci provo UT catalog numbers of host specimens rothschild 1904 deposited in the university of utah museum of in salt co wasatch mts big cotton- natural history UU mammal collection and lake wood canyon vicvie redman campground 2560 flea in the MLB museum BYU specimens m spruce fir 21 october 1990 1 Y ex sorex insect collection are given in parentheses when montimonticolusmonticuluscolus UU 29163 same locality 28 available otherwise unless indicated speci- october 1990 1 Y ex sorex montimonticolusmonticuluscolus UU mens were collected by me and are retained in 29164 my personal collection stark 1959 reported this species from cache county my collections extend the Cartcarterettecarterettaeretta alavataclavata good 1942 known range of this species further south in washington co west slope beaver dam utah along the wasatch cordillera it is found on shrews sorex sppapp and mustela sppapp mts vievic welcome spring 1220 m 20 march 1988 1 Y ex chaetodipus formosusformosus beaver delotelisDelotelis telefonitelegonitelegoni dam 23 february 1952 1 Y BYU 3462 ex rothschild 1905 perognathus chaetodipusformosuschaetodipus formosusformo sus coll C L hayward beaver dam wash 17 april salt lake co wasatch mts big cotton- wood canyon vicvie campground 2560 1952 16 BYU 3607 ex peromyscus truet redman spruce fir 21 1990 1 S ex coll grace grant et al in october 16 clethrionomys gasperigapperigapperi same date & locality few collections of this are known species 1 Y ex tamiasciurusTamiasciurus hudsonicushudsonicus nest from the utah tooelethoele county stark 1959 16 nest about 3 m above ground level also con- egoscue 1 1976 specimen sex unknown tained many red squirrel fleas orchopeasOrchopeas c washington county jellison and senger 1976 caedens the squirrel probably carried this 995assggs2ss2 6 6 2 Y Y 1 it has also been collected in vole flea to its nest same locality 15 clark county NV the type locality good 1942 september 1991 1 Y ex peromyscus caniculamanicula and mohave county AZ augustson and dur- tus same locality 5 october 1991 16lc ex ham 1961 it is likely a nest flea of C formoformosussus clethrionomys gasperigapperigapperi

lassociatcd regional and university pathologists incinelne saltsait like city UT 84108 address forhorbor correspondence 59303930 S sultan circlecircie murray UT 84107693084107 6930

92 199519951 NOTES 93

delotelisDelotelis telefonitelegonitelegoni has rarely been found in megabothris asio megamegacolpusmegaco1puscolpus utah single specimens have been collected in jordan 1929 sanpete county stark 1959 and in utah co Lake 22 county egoscue 1988 it is significant that rich laketowntown august 1952 11 & ct 26 Y Y BYU these collections were made in summer months lidd 509750995097 5099 510151035101 5103 august 1951 and july 1985 respectively 510551195105 5119 512151315121 5131 5135 5137 5138 many more collections will likely be made if 5737 5738 ex microtus montanus nests 3 ex- amined coll D E beck & L Lake this species is searched for during the cooler beck laketowntown 26 1953 4 Y Y BYU 7823 months of the year also it presumably would june id16 782378277827 ex microtus nests 3 examined coll al be profitable to search nests of microtus and beck et sevier co south end 5 august clethrionomys for this species fish lake 1952 1 Y BYU 5622 ex microtus sp coll coffey & killpack monroe mt 7 mi 11 deringismeringisMeringis shshannonishannonmiannoni jordan 1929 km W ofkoosharemofkoosharemKoosharem 30 july 1958 IS16 229Y Y eads et al 1987 listed two collections of ex microtus sp 3 examined coll unknown this species from utah the specimens are the range of this boreal vole flea extends present in the BYU collection however the deep into south central utah only two speci- records are erroneous because the collection mens are known from idaho including one locality douglas county does not exist in from bear lake county adjacent to rich utah all other cited records of this species county baird and saunders 1992 collections are from the states of washington and oregon from ravalli and beaverhead counties MT except a lone locality record in humboldt are documented holland 1950 and speci- county NV lewis et al 1988 mens from ravalli county are present in the natural history museum london T M stenistomeraStenistomera hubhubbardihubbarddbardi howard personal communication A point egoscue 1968 mapped record roughly on the utah wyoming border given in haddow et al 1983 is evi- this rare species was listed by tipton and dently meant to be laketownLake town because four saunders 1971 as occurring in utah although specimens with the same collection data as the no specific records were cited egoscue per- 22 august 1952 series are present in the sonal communication knows of no records from natural history museum london howard utah and no specimens were present in the personal communication MLB life science museum the main reposi- tory of Tiptiptontoistorss utah collections in addition to chaetopsylla stewartistewstewartearti johnson 1955 the type specimens from oregon egoscue utah co wasatch mts near summit of 1968 the only other published of S record alpine loop american fork or provo can- hubbardihubbarddhubbardi is that of et al 1988 also lewis yons 24 november 1965 6as6 ct 6 Y Y ex from oregon 6s it is unlikely that the species mustelafrenata coll D andrews summit co has been collected in utah uinta mts 12 mi 08080.8 kalkmlkm E bald mt 8 august 1957 id16 ex martes sp coll D allred megarthroglossus becki & M killpack tipton & allred 1951 these specimens are the only ones known salt lake co wasatch mts mouth of other than the type series from cache county little cottonwood canyon 1676 in scrub lewis and lewis 1994 weasels seem to be oak 3 december 1989 1516 1 Y ex neotoma the preferred host cinerea nest the species has been collected only in euhoplopsyllus glaciaglacialisglacialistlis lynx utah kane plute utah and wayne counties baker 1904 tipton et al 1979 and arizona augustson and salt lake co wasatch mts big cotton- durham 1961 this is the northernmost record wood canyon vicvie redman campground 2560 known some 37 km north of the type locality in 17 august 1988 3 Y Y 1 CT ex lepus ameri in the wasatch mountains megarthroglossus banuscanus UU 28674 big cottonwood canyon becki is a nest flea of woodratswoodrats principally the 2280 in 30 august 1988 191 Y ex lepus ameri bushy tailed woodratwoodral neotoma cinerea banuscanus big cottonwood canyon vicvie butler 94 GREAT BASIN naturalist volume 55 fork trailhead 2182 in 19 may 1991 2669669862sas S ex 1968 ariewarlewA new species of the genus stenistomerastemstomeraStenisStemstomera lepus americanosamericanusameric anus siphonaptera hystnchopsyllidaehystrichopsyllidae southernSouthernthein cali- fornia academy of sciences bulletin 67 138142138 142 previously unknown in utah the nearest 1976 flea exchange between deer mice and some published records are for ravalli county MT associated small mammals in western utah great kohls 1940 more than 570 km to the north basin naturalist 36 475480475 480 the type locality is moscow ID about 790 kinkm 1977 the sagebrush vole flea megabothnsmegabothris clan tonitorntormtomm prinprincelceicel in western utah with comments on the to the northwest baker 1904 flea con- in this is distribution of megabothrismegabothns in the bonneville basin sistently found on the snowshoe hare lepus great basin naturalist 37 757675 76 americanosamericanusamericanus and its predator the lynx lynx 1988 noteworthy flea records fromheom utah nevada canadensis the form E glaciaglacialisglacialistlis alfinisaffinis is com- and oregon great basin naturalist 48 530532530 532 mon in utah and surrounding states on rabbits 1989 A new species of the genus traubellaTraubella sipho- naptera ceratophyllidae southern california and jackrabbitsjackrabbits sylvilagusSylvilagus sppapp and lepus academy of sciences bulletin 88 131134131 134 sppapp other than L americanusamericanosameric anus questionable GOOD N E 1942 Cartcarterettecarterettaeretta cartericarten clalavataclavataavata a new sub- records of E g lynx from the states of species from nevada and notes on synonymy sipho- tamaulipas and veracruz mexico ex sylvilagusSylvilagus napnapteratera annals of the entomological society of america 35 110113110 113 florifloridanus1otiorioridanusdanus and unidentified sylvilagusSylvilagus sp are J HADDOW J R TRAUB AND M rothschild 1983 dis- listed by ayala et al 1988 tributiontrib ution of ceratophyllidceratophylhd fleas and notes on their hosts pages 4216342 163 in R traub M rothschild and acknowledgments J F haddow the rothschild collection of fleas the ceratophyllidae keys to the genera and host relation- ships with notes on their evolution zoogeography comments by glenn E haas B C kondra- and medical importance 288 appp privately published tieff and an anonymous reviewer improved HOLLAND G P 1950 notes on megabothnsmegabothris asio baker the manuscript richard W baumann curator and M caiecalecaiccalcanfercalearifercalcanariferferjer wagner with the description of a of insects at the M L bean life science new subspecies siphonaptera ceratophyllidae museum young canadian entomologist 82 126133126 133 brigham university kindly JELLISON W L AND C M SENGER 1976 fleas oftestofwestof west- allowed me to examine specimens kept there ern north america except montana in the rocky harold J egoscue confirmed identification of mountain laboratory collection pages 5513655 136 in the E g lynx theresa M howard of H C taylor jr and J clark eds papers in honor the of jerry flora western washington state college natural history museum london sent data bellingham on specimens in the rothschild collection KUCERA J R AND G E HAAS 1992 siphonaptera fleas collected from small mammals in montane southern great CITED utah basin naturalist 52 382384382 384 literature KOHLS G M 1940 siphonaptera a study of the species infesting wild hares and rabbits of north america AUGUSTSON G FE AND F E DURHAM 1961 records of north of mexico national institute of health bulletin fleas siphonaptera from northwestern arizona 175 southern california academy of sciences bulletin LEWIS R E AND J H LEWIS 1994 siphonaptera of 60 100 105 100105 north america north of mexico vermipsyllidaeVermipsyllidae and AYALA R J C MORALES N WILSON J E LLORENTELLORENTC AND rhopalopsyllidae journal of medical entomology PONCE 1988 catalognogatalogo las H E catalogoCatacatdlogo1090 de pulgasdulgas insecta 31 829882 98 siphonaptera en ele1ea museo dedc zoologiazoologicZoologia facultad de LEWIS R E H LEWIS AND C MASER 1988 fleas autonomy J the of cienciascienciakCiencias universidad nacional aut6nomaautonomaAutonoma de mexico the pacific northwest oregon state university press 1 gata Coleccicoleccionon alfredo barrera serieserlesene catalogsCatacatalogoscatdlogoslogos del corvallis 296 appp museo de Zozoologiazoologicologia alfonso L catalogocatdlogoCatacatalognologo herrera STARK H E 1959 the siphonaptera of utah USU S no 1 102 appp department of health education and welfare BAIRD C R AND R C SAUNDERS 1992 an annotated communicable disease center atlanta GA 239 appp checklist of the fleas of idaho siphonaptera idaho TIPTON V J AND R C SAUNDERS 1971 A list of arthro- agriculturalagi icultural experiment station bulletin 148 pods of medical importance which occur in utah BAKER C F 1904 A revision of the siphonap- revision american with a review of arthropod bomeborne diseases endemic in tera or fleas together with a complete list and bibli- the state brigham young university science bulletin ography of the group proceedings of the US national biological series 15131151lsiisi15 1311 31 museum 273652736546927 365469365 469 TIPTON V J H E STARK AND J A WILDIE 1979 EADS R B E G CAMPOS AND G 0 MAUPIN 1987 A anomiopsyllinae siphonaptera hystrichopsyllidaehystnchopsyllidae reviewi eviewaview of the genus Mederingismenngismeringisringis siphonaptera hystnhystrihysen II11 the genera callistopsyllus conorhinopsylla psyllidaechopsylhdaechopsyllidaechoeho journal of medical entomology 24 megarthroglossus and stenistomeraStenisStemsstemstomeratomera great basin 467476467 476 naturalist 39 351418351 418 EGOSCUEEGOSC UE H J 1966 new and additional host flea associ- ations and distributional records offleasof fleas from utah received 25 may 1994 great basin naturalist 26 717571 75 accepted 10 august 1994 information FOR AUTHORS the great basin naturalist welcomes previously VOUCHER SPECIMENS authors are encouraged to unpublished manuscripts pertaining to the biologi- designate properlproperlyy prepare label and deposit cal natural history of western north america pref- high quality voucher specimens and cultures docu- erence will be given to concise manuscripts of up to menting their research in an established permanent 12000 words simple species lists are discouraged collection and to cite the repository in publication SUBMIT manuscripts to richard W baumann references IN THE TEXT are cited by author and editor great basin naturalist 290 MLBM PO box date eg martin 1989 or martin 1989 multiple 20200 brigham young university provo UT citations should be separated by commas and listed 84602020084602 0200 A cover letter accompanying the man- in chrochronologicalnological order use et al after nameofnameonname of uscript must include phone numbers of the author first author for citations having more than two submitting the manuscript and FAX number and authors emailE mail address when applicable the letter must acknowledgments under a centered main also provide information describing the extent to heading include special publication numbers when which data text or illustrations have been used in appropriate other papers or books that are published in press literature CITED also under a centered main submitted or soon to be submitted elsewhere heading lists references alphabetically in the fol- authors should adhere to the following guidelines lowing formats manuscripts not so prepared may be returned for revision mack G D and L D flake 1980 habitat rela- manuscript preparation consult vol 51 no tiontionshipsships of waterfowl broods on south dakota 2 of this journal for specific instructions on style stock ponds journal of wildlife management and format these instructions guidelines FOR 4469544 695 700 manuscripts SUBMITTED TO tiietlleTHETEIE GREAT BASIN sousa W PE 1985 disturbance and patch dynamics naturalist supply greater detail than those pre- on rocky intertidal shores pages 101 124 in sented here in condensed form the guidelines are S T A pickett and PE S white eds the ecolo- printed at the back of the issue additional copies gy of natural disturbance and patch dynamics are available from the editor upon request also academic press new york check the most recent issue of the great basin nat- coulson R N and J A witter 1984 forest ento- uralist for changes and refer to the CBE style man- mology ecology and management john wiley ual ath5th edition council of biology editors one and sons inc new york 669 appp illinois center suite 200 111 east wacker drive chicago IL 60601429860601gogol 4298usa4298 USA PHONE 312 616- TABLES are double spaced on separate sheets and 0800 FAX 312 6160226616 0226 24 designed to fit the width of either a single column TYPE AND DOUBLE SPACE all materials including 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published the rate for individual subscribers will TITLE PAGE includes an informative title no longer be 35 per page however manuscripts with com- than 15 words names and addresses of authors a plex tables andor numerous half tones will be running head of fewer than 40 letters and spaces assessed an additional charge reprints may be pur- footnotes to indicate change of address and author chased at the time of publication an order form is to whom correspondence should be addressed if sent with the proofs other than the first author FINAL CHECK ABSTRACT states the purpose methods results cover letter explaining any duplication of and conclusions of the research it is followed by information and providing phone numbers 6 12 key words listed in order of decreasing FAX number and emailE mail address importance to be used for indexing 3 copies of the manuscript and WordwordperfectPerfect TEXT has centered main headings printed in all diskette capital letters second level headings are centered 0 conformity with instructions in upper and lowercase letters third level head- photocopies of illustrations ings begin paragraphs ISSN 001736140017ooi001 3614736147 GREAT BASIN naturalist voivol 55 no 1 january 1995 CONTENTS articles life histories of stonestonefliesflies plecoptera in the rio conejos of southern colorado R edward dewalt and kenneth W stewart 1 pollinator sharing by three sympatric milkvetches including the endangered species astragalus montiimontai S M geer V J tepedino T L griswold and W R bowlin 19 factors affecting selection of winter food and roosting resources by porcupines in utah dave stricklan berranjerran T flinders and rex G catesgates 29 historic expansion of juniperus occidentoccidentalisoccidentalistalis western juniper in southeastern oregon richard F miller and jeffery A rose 37 rangeland alpha diversities harvey valley lassen national forest california raymond D ratliff 46 effects of salinity on establishment of populus fremonfremontiifremontiatii cottonwood and tamarix ramosissima saltcedarsaltcedar in southwestern united states patrick B shafroth jonathan M friedman and lee S ischinger 58 names and types of hedysarum L fabaceae in north america

1 I stanley L welsh 66 whipworm trichuris dipodomys infection in kangaroo rats dipodomys sppapp effects on digestive efficiency james C munger and todd A slichter 74 local distribution and foraging behavior of the spotted bat eudermaEueudeemaderma maculatummaculatum in northwestern colorado and adjacent utah jay FE storz 78 the chrysothamnus ericameria connection asteraceae loran C anderson 84 notes reproductive behavior in merrlmerriammerriaasMerriaasamss chipmunk tamias merrimerriamiami stephen B compton and J R callahan 89 additional records of fleas siphonaptera from utah james R kucera 92