Full Issue, Vol. 57 No. 2

Home , Ephydatia fluviatilis, Ephydatia muelleri

Great Basin Naturalist

Volume 57 Number 2 Article 14

5-7-1997

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

Recommended Citation (1997) "Full Issue, Vol. 57 No. 2," Great Basin Naturalist: Vol. 57 : No. 2 , Article 14. Available at: https://scholarsarchive.byu.edu/gbn/vol57/iss2/14

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

GREATG R E A T BASINBAS I1 naturalistnaturalist

VOLUME 57 NQ 2 APRIL 1997

br16hamBRIGHAM 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 801378soisol8013785053801 378 5053 8013786688801 378 6688 FAX 8013783733801 378 3733 emailE mail nmshbll1byuedu

associate editors J R CALLAHAN PAUL C MARSH museum of southwestern biology university of center for environmental studies arizona new mexico albuquerque NM state university tempe AZ 85287 mailing address box 3140 hemet CA 92546 STANLEY D SMITH BRUCE D ESHELMAN department of biology department of biological sciences university of university of nevada las vegas wisconsin whitewater whitewater wlWI 53190 las vegas NV 89154400489154 4004 JEFFREY J JOHANSEN PAUL T TUELLER department of biology john carroll university department of environmental resource sciences university heights OH 44118 university of nevada reno 1000 valley road reno NV 89512 BORIS C kondratieff department of entomology colorado state ROBERT C WHITMORE university fort collins CO 80523 division of forestry box 6125 west virginia university Morganmorgantowntown WV 26506612526506 6125

editorial board berranjerran T flinders chairman botany and range science duke S rogers zoology wilford M hess botany and range science richard R tolman zoology all are at brigham young university ex officio editorial board members include steven L taylor college of biology and agriculture H duane smith director monte L bean life science museum richard W baumann editor great basin naturalist the great basin naturalist founded in 1939 is published quarterly by brigham young university unpublished manuscripts that further our biological understanding of the great basin and surrounding areas in western north america are accepted for publication subscriptions annual subscriptions to the great basin naturalist for 1997 are 25 for individual sub- scriscribersbers 30 outside the united states and 50 for institutions the price of single issues is 12 all back issues are in print and available for sale all matters pertaining to subscriptions back issues or other busi- ness should be directed to the editor great basin naturalist 290 MLBM PO box 20200 brigham young university provo UT 84602020084602 0200 scholarly exchanges libraries or other organizations interested in obtaining the great basin naturalist through a continuing exchange of scholarly publications should contact the exchange librarian 6385 HBLL PO box 26889 brigham young university provo UT 84602688984602 6889

editorial production staff joanne abel technical editor

copyright 0 1997 by Brigbrighamhain young university ISSN 001736140017 3614 official publication date 7 may 1997 5975 97 750 22025 the great basin naturalist PUBLISHED AT PROVO UTAH BY BRIGHAM YOUNG university

ISSN 001736140017 3614

VOLUME 57 30 APRIL 1997 no 2

great basin naturalist 572 019971997 appp 93 103 freshwater SPONGES PORIFERA spongillidae OF WESTERN MONTANA

susan H bartoni and john S addis2addiseaddisl2

ABSTRACT between may 1992 and april 1996 freshwater sponges fonporiferaPonferahera spongillidae were collected at 24 sites distributed among 6 sub major drainage basins in western montana to determine the species present water samples also were analyzed from 16 of these sites and from 9 sites at which no sponges were detected to characterize sponge habitats chemically thieethree species of sponges were identified ephydatia muellerimuellen em eunapiusfragiliseunapiusEunaeunepiusprus frafragilisfragulispragilis Eef and spongilla lacus tris sl A ath4th type of specimen was present at 2 sites but could not be identified because of the absence ofgemoulesgemmulesofgemmulesgemmules and gemmoscleres at 46 of the sites containing sponges more than I1 specimen type was present sponges were most commonly found near outlets of lakes attached to sides or undersides of submerged rocks and logs they appeared as encrusting em ef sl lobate em and fingerlike sl growths varying in color from light tan to green dimensions of the spispiculesspicklescules varied greatly within each species and expanded previously recorded ranges no factors limiting sponge dis- tributiontribution were identified but ranges of conductivity em and of silica em sl calcium em and magnesium em con- centrationscentrations were expanded beyond those reported previously

key words freshwater sponges bonPonporiferaferapera spongillidae ephydatiaepbydatia muellenmuelleri eunapiusEunapius frafragilisfragulisgilis spongilla laculacustnslacustrisstris montana

although freshwater sponges form part of 1932 jewell 193519391935 1939 poirrier 1969 harrison the benthic community in many of the world s 1974 in addition some attempts have been lentic and lotic habitats they are among the made to identify factors that affect species dis- least understood of animal groups basic ques- tributiontribution old 1932 jewell 1935 1939 poirrier tions about their biogeography and ecology 1969 strekal and mcdiffett 1974 freshwater remain unanswered frost 1991 their distri- sponges of the west in contrast have received butions have not been completely determined much less attention this is unfortunate since and their habitat requirements have not been water quality is an important issue in the west- fully defined ern united states and since sponges are poten- freshwater sponges in the united states tially valuable indicatorsbioindicatorsbio of water quality have been studied most extensively in the east harrison 1974 francis and harrison 1988 and midwest species present in these regions richelle maurer et al 1994 have been described and their habitats charac- in the mountain west only freshwater terized in chemical and physical terms old sponges of colorado have been studied in

department of biology carroll college helena MT 59625 author to whom correspondence should be addressed

93 94 GREAT BASIN naturalist volume 57 detail williams 1977 1980 freshwater sponges tanes upper daridarkoarkclariciariclark fork lower clark fork have been collected in montana see for ex- kootenai and flathead water quality bureau ample young 1935 poirrier et Aal 1987 but no 1991 sites included lakes and ponds man studies focusing on the freshwater sponges of made as well as natural and in some cases the this state have been published in this paper first 50 m of rivers or streams draining lakes and we report results of a survey of freshwater ponds most sites were within 2 km of a road sponges in western montana the survey was in most cases we collected samples by wad- conducted to identify the species present and to ing near the shore and handhandpickingpicking likely determine chemical and physical ranges for substratasubstratalsubstrata submerged rocks or logs most each species habitat the latter is a necessary samples were obtained from substratasubstratalsubstrata sub- step in defining factors that influence sponge merged at depths of less than 1 m in I1 lake distribution blanchard lake samples were collected by diving substratum depth min this case was MATERIALS AND METHODS approximately 3 m we attempted to collect gemoulesgemgemmulesmules asexual propagules with the sampling samples by scraping the substratum with a knife between may 1992 and april 1996 we ob- or gathering a piece of the substratum itself tained sponge samples from 24 sites in western for transport to the laboratory samples were montana listed in table 1 these sites were placed inm small containers with lake water distributed among 6 sub major drainage basins between june and august 1994 surface missouri sun smith upper missouri tribu water grab samples were taken from 16 sites at

tableTABLL 1 surveyed lakes in westeinwestern montana containing freshwater sponges species lake county locality present

MISSOURI SUN SMIIIISMITH BASIN holter lake lewis and clarkglaik 49o59n149059n112000w12o00w 13 upper holteiholterholtelholtey lake lewis and clark 4650n11200w46050n112000w 1 UPPERUPPLR MISSOURI tributariesTRIBUTATRIBU parlesPARIESrlesRIES BASIN lowellower mineiminer lake beaverhead 4520n1i3o34w45020n113034w 13 rockroekbockboek island lake beaverhead 45018n113041w45118n113141w 23 upper mineiminelminer lake beaverhead 4516n11341w45016n113041w 2 hebgen lake madison 44052n1120w44152n1 I1 i20w 13 jerome rock lake madison 4523ni45023n1128w4523 NI I1 i28w 2 pond below blue paradise lake madison 4457n144057n1126wI1 i26w 2 quake lake madison 4450ni445044050n111026wNI I1 126w 23 willow creek reservoirreser volivoir madison 4543nmo42w4543n11142w 1 UPPER CLARK FORK BASIN blanchard lake missoula 47101n147001n113023w1323w 234 salmon lake missoula 4706n11324w47006n113024w 123 coopers lake powell 47005n112055w47105n112155w 34 pond nealnear west forifoifolforkk of bittelbitterrooti oot river ravalli 4606n11411w46006n114011w 1 lowerLOWLR CLARK FORK BASIN diamond lake mineral 4709n115111w47009n115011w 13 booiemooiemoore lake mineral 47011n5015w4711n11515w 3 kootenaiKOOTLNAI BASIN kilbikiikliKilbrennanennan lake lincoln 4835n115053w4835n115153w 13 loon lake lincoln 48105ni48005481054805n115011wNI 1511 IW 23 platheFLATHEFLAIIILADAD BASIN pond near bailey lake flathead 4828n11408w48028n114008w I1 spoon lake flathead 4829n148029n11409w1409w 2 cedargedal lake lake 4740n113157w47040n113057w 3 pond below cedar lake lake 4740n113156w47040n113056w 3 ducliarmeduchaime lake lake 4737n113157w47037n113057w 2 lowellower dueDucduchaimeducharmebalmehalmebarme lake lake 47037n113057w47137n113157w 2 spllicslspecies I1 eunapiusfiragilislunitptin fraffihs 2 spongilla laculitlii&tilacustrisstrisis 3 lphydatiaephydatia muelkrtinuelleri 4 unidentified sponge 199719971 freshwater SPONGES OF WESTERN MONTANA 95 the same time sponge specimens were collected piesples were collected total hardness as cac03 samples were taken at locations sponges were was obtained by multiplying values obtained collected with the exception of quake lake for calcium and magnesium by 2502.50 and 4124.12 where water and sponge samples were obtained respectively and adding the products silicon at sites separated by approximately 050.5 km was converted to sobslog by multiplying its val- during this period water samples were also ues by 2142.14 conductivity ph and tempera- obtained from 9 lakes that lacked sponges in ture were recorded on site using a horiba ulouioU 10 these cases samples were collected at conve- water checker nient sites close to shore water samples were collected in polyethylene bottles at a water RESULTS depth of approximately 020.2 m according to the field procedure manual water quality bureau from the specimens collected at sites dis- 1991 water to be analyzed for total organic tritributedbuted among 6 sub major drainage basins in carbon see below was preserved with h2so4HSO western montana we identified 3 species of freshwater eune fragilisfragulisgilis spongilla at a final concentration of 020.2 vv sponges eunapiusfragilisEunaeunapiuspiusplus fra lacustrislacustris and ephydatia muelleri locations of identification sites and types of sponges found at each are we identified sponges on the basis of de- indicated in figure I1 and table 1 morphologi- scriptscriptionsions provided by penney and racek 1968 cally distinctive specimens were also obtained appearances and dimensions of spispicklesspiculescules and at 2 sites located in the upper clark fork basin positions of gemoulesgemgemmulesmules within the sponge were but these could not be positively identified be- the principal criteria on which identifications cause of the absence of gemgemoulesgemmulesmules and gemmo were based scleresskleres spispiculesspicklescules associated with the gemmule we isolated spispicklesspiculescules from sponge tissue at 46 of the sites 2 or more types of sponges according to methods described in pennak were present sponge morphomorphologieslogies and habi- 1978 sponge samples were placed in test tats are described below tubes with nitric acid and boiled they were fragilisfragulisgilis 1851 allowed to stand for at least 24 h and then eunapiusfragiliseunapiusEunapiuspluspins fra leidy collection numbers 1107 1114 were rinsed repeatedly with water and ethyl 11071114ilot 1120 211128113321 1128 1133 35114235 1142 alcohol before drying and mounting on slides 114711641169117111471164 1169 1171 we did not boil gemgemoulesgemmulesmules but allowed them to figs 232 3 stand in nitric acid for 1 6 h before rinsing two permanent slides I1 containing intact gemgemoulesgemmulesmules eunapiusEunapius frafragilisfragulisgilis was collected at 10 of the and I1 containing megasmegasclerescleres needlelike spic 24 sites at 4 sites it was the only species col- ules of the main sponge skeleton and micro lected table 1 at the other 6 sites it was col- scleresskleres small dermal spispiculesspicklescules if present were lected with E muelleri or with both E muelleri made for most specimens collected slides and S lacustrislacustris no sites contained only E were deposited in the invertebrate collection frabrabrefragilisfragifisfragulisgilisgifis and S lacustrislacustris most often E frafragilisfragulisgilis of carroll college appeared as an encrusting growth on the undersides or sides of submerged logs less com- water analyses or monly rocks substratasubstratalSubstrata settled by E frafragilisfragulisgilis we performed the following analyses on were usually located at the outlets of lakes or each water sample methyl orange alkalinity ponds the sponge rarely exceeded 050.5os cm in calcium total hardness as cac03 magnesium height and its color was either tan or green silicon and total organic carbon analyses were due to the presence of symbiotic algae clif- performed by the montana state chemistry ford 1991 conspicuous asculaoscula were usually laboratory following procedures in the US present fig 2 environmental protection agency manuals specimens of E frafragilisfragulisgilis contained mega standard methods for chemical analysis of scleresskleres that were sharply pointed at both ends water and wastes 1983 and methods for the and were smooth and slightly curved micro determination of metals in environmental scleresskleres are absent in this species the gem samples 1991 24 96 h after the water sam mules which ranged in diameter from 250 96 GREAT BASIN naturalist volume 57

on5ona

I1 A

A A 0 n

scoleascole 4pap 8 ala4la 1

fig 1 distribution of sponge species in western montana Eunaeunapiuseunapwspiusplus frafragilisfragifisfragulisgilisgifisgibis EH spongilla laculacustrisstris A ephydatia muelleri 0 unidentified sponge s sub major drainage basins outlined by thick lines are as follows 1 missouri sun smith basin 2 upper missouri tributariesTributaries basin 3 upper clark fork basin 4 lower darkoarkclark fork basin 5 kootenai basin 6 flathead basin

to 550 gmlm were either arranged in a distinc- spongilla lacustrislacolacuiacolacustrzsstris linnaeus 1758 tive pavement layer or clustered in groups of collection numbers 1099 1100 1103 3 5 gemmoscleres oriented tangentially in 1108 9112211249 1122 1124 25113225 1132 the pneumatic layer of the gemmule had 1137 381140 41114911521154411149 1152 1154 blunt rounded ends were slightly to strongly 1156 57116157 ligi1161 631165 curved and were covered with thick spines fig 4 concentrated at the ends spicule dimensions are given in table 2 spongilla lacustnslacustrislacustris was collected at 11 sites one specimen of E frafragilisfragulisgilis from diamond it was the only species collected at 6 sites and lake an oligotrophic mountain lake located in at the 5 other sites it was collected with E the lower clark fork basin was unique in that muellenmuelleri or with E muellenmuelleri and either E frag almost every megasclere contained at least 1 ibs or the unidentified sponge table 1 more expanded bulb along its length fig 3 normal than the other species S lacustnslacustrislacustris varied in specimens of E frafragilisfragulisgilis and E muelleri were gross morphology while often encrusting it also collected from the same region of diamond occasionally extended long 10 cm cylindri- lake cal branches fig 4 it was usually found at 199711997 freshwater SPONGES OF WESTERN MONTANA 97 WK ses

A issiks773

fig 2 eunapiusEunapius frafragilisfragulisgilis on the underside of a log pond adjacent to bailey lake note conspicuous asculaoscula in upper specimen

TABLE 2 spicule dimensions of western montana sponges mean standard error range dimensions in tm species E frafragilisfragifisfragulisgilisgifis S lacustrislacustris E muelleri spicule dimendimensionsionaslona n 13 n 20 n 23 Megasmegasclerescleres length 275 27 90 3921392392.1 287 35 120 408408a 269 19 150 391 width 959.5gs 151.5 20 140b14 0bab 110 202.0 25252.5 19019.0190119.01igo 1 105los10.5 15 30303.0 180b microscleresMicros cleres length 67 7 35 120 width 353 5 050 5 isls151.5 757.5 gemmoscleres length 79 13513.5 30 148e148 89 24 25 1770 145 151.5lsis 95959.5 25025.0 width 606.0go 101.0loio 35 80 45 151.5isls 30303.0 959.5gs 404.0 05os050.5 15 757.5 rotulekotule diameter 17517.5 202.0 959 5 27527.5

lessaless and greater than previously reported see text blessless than previously lecordedrecorded see text greater than previously recorded see text

outlets of lakes or ponds but the branching curved megascleresMegas cleres however were smooth form of the sponge occurred in quiet water at while microsmicrosclerescleres were covered with small sites distant from the outlet or at sites near the spines spicule dimensions are given in table 2 lake outlet that were not noticeably affected Gemgemoulesgemmulesmules were small ranging in diameter by the current usually S lacustrislacustris was found from 250 to 500 imtm and were typically scat- growing on the sides or tops of rocks or logs tered throughout the sponge As others have the branching form also was found growing observed gilbert and simpson 1976 gemmo directly out of bottom sediments the sponge scleresskleres were not consistently present those was either light tan or green that were found had pointed or rounded ends both megasmegasclerescleres and microsmicrosclerescleres were were slightly to strongly curved and were cov- sharply pointed at both ends and slightly ered with prominent spines 98 GREAT BASIN naturalist volume 57

fig 3 abnormal megasclere ofeunapiusofeunapiusfragilisfragiasfragihs showing expanded bulbs diamond lake bar 50 tm

ephidatiaephydatia muellenmuelleri lieberkuhnLieberkuhn 1855 had I1 or more spines extending from their collection numbers 1106 1112 13 1115 shafts and misshapen rays were often present 1118 1126 27 1129301129 30 1139 1143461143 46 figs 7bcnbc 114811501148 liso1150 51115851 1158 60116760 1167 68117068 1170 figs 5 7 unidentified sponge collection numbers 1119 ephydatia muelleri was collected from 14 11551172 75 11 was collected with at least 1 sites at sites it fig 8 other sponge type E frafragthsfragilisfragulisgilis S lacustrislaculacustrzsstris or the unidentified sponge table 1 it was found some specimens collected from undersides most often on sides or undersides of logs or of rocks and logs in blanchard and coopers rocks the sponge varied in morphology from lakes lacked gemoulesgemgemmulesmules and gemmoscleres and thin 05 050.50 5 cm encrustations to lobate forms in could therefore not be positively identified which rounded masses approximately 2 cm tall although collected from 2 different sites these extended from basal mats fig 5 large asculaoscula specimens exhibited similar morphomorphologieslogies at were sometimes present its color varied from both locations sponges were light tan and light tan to green formed hard disc- or cushion shaped masses this species which like E frafragilisfragulisgilis lacks megascleresMegascleres were slightly curved sharply microsmicrosclerescleres contained megascleresmegascleres that were pointed at both ends and covered with short sharply pointed at their ends and slightly curved spines except at their tips many megasmegasclerescleres although they were often covered with small also had a slight midregionmidregion expansion fig 8 spines except at their tips many were smooth the megasmegasclerescleres bore an overall resemblance or had few spines fig 6 both smooth and to the spined megasmegasclerescleres of E muellenmuelleri but spined megasmegasclerescleres were frequently present in were substantially wider 2102101521021.0 15isls1.5 imtm versus the same specimen spicule dimensions are 10510.510losios 5 151.51 5 gmjmJm for E muellenmuelleri normal speci- given in table 2 mens of E muellenmuelleri also were present in both Gemgemoulesgemmulesmules langmalangmgranging in diameter from 350 to lakes specimens collected during return visits 450 gmum were scattered throughout the sponges to coopers lake in november 1995 and feb- or were concentrated at the base A distinct ruary 1996 did not contain gemoulesgemgemmulesmules in fact pavement layer was not present however the specimens of this sponge collected at these gemmoscleres were birotulatebirotulate fig 7aaa rotulesromules times resembled specimens collected during had similar diameters within each gemmo summer months in contrast E muellenmuelleri had scleresciere and had irregular and deeply incised rays died back and occurred only as mats of gem numbering no more than 12 shafts connect- mules held within skeletal frameworks the ing the rotulesromules were moderately thick their unidentified sponge might be a species ofephyof ephy lengths usually not exceeding the diameters of dotiadatla trochospongilla or anheteromeyeniaanheteromeyema fur- the rotulesromules gemmoscleres were arranged in a ther studies of this sponge are underway single layer perpendicular to the gemmule and chemical and physical factors were embedded in a well developed pneumatic layer chemical and physical ranges characteriz- slight malformations of gemmoscleres were ing habitats of the identified species are given not uncommon but those present in sponges in table 3 these langestangesranges were determined fromblombrom cedar lake and an adjoining pond flat- from data collected at 16 of the sites that con- head basin were extreme many birotulates tained sponges the table also includes ranges 19971 freshwater SPONGES OF WESTERN MONTANA 99

fig 4 spongillallaliaiia lacustrislacustris with fingerlike projections growing from beneath a rock salmon lake

0 NM ilkkis

ed N erwOHM

fig 5 ephydatia muelleri exhibiting encrusting and lobate growth salmonsaimon laielake

derived from data collected at 9 lakes that did were wider than those determined for E not contain sponges although there was over- muelleri or S laculacustrisstris in addition S laculacustrisstris lap among the ranges determined for the 3 had the narrowest ranges of alkalinity calcium species widths of the ranges varied chemical and magnesium concentrations and total hard- ranges determined for E frafragilisfragulisgilis except for ness differences between variances of data the ranges of total organic carbon and silica obtained from sites containing the different 100 GREAT BASIN naturalist volume 57

fig 6 smooth and sparsely spined megascleresmegas cleres ofephydatia muelleri bar 50 pmgm

Ffigig 7 gemmoscleres ephydatiaofofephydfitia muelleri a normal birotulatebirotulate gemmoscleres top and side views b and c malformed gemmoscleres top view b and side view c cedar lake bar 25 tmjm

fig 8 megasclere of unidentified sponge coopers lake note slight midregionmidregion swelling dark band bar 50 urnumm sponge species however were not statistically discussion significant at a 0050.05oos levene s test for equal- small ity of variances one tailed although at aX the number of sponge species found in 0100.10olo the variance of the data for magnesium western montana is consistent with results of concentration from lakes containing E frafragilisfragulisgilis other surveys of sponges of the mountain west was significantly greater than the variance and seems to reflect an overall paucity of sponge this region his listing of the determined from lakes containing S lacustrislacustris species in in sponges of clifford 1991 reports only differences among the means of the data were alberta 3 species the same 3 reported by us ephyraephyda also not statistically significant at X 0050.05 a tia muelleri eunapiusEunapius fragilisfragulisgilis and spongilla one way ANOVA fra lacustrislacustris williams 1980 reports 4 species some lacked values sites that sponges had fromhrom colorado the previously mentioned 3 plus for chemical and physical factors that lay out- ephydatia fluviatilisfluviatilis found at 1 location on side the ranges determined from sponge con the eastern plains that these 3 species of the tainingbaining lakes but others had chemical and 27 reported for the united states and canada physical profiles that were very similar to lakes frost 1991 were found in western montana is thatmat contained sponges the difference between not surprising when their distributions are con- means of the data obtained from lakes contain- sideredsi all 3 species are widespread eunapiusEunapius ing sponges and lakes lacking sponges was not frafragilisfragulisgilis is truly cosmopolitan being present significant at a 0050.05oos pooled t test on every continent whereas E muelleri and 199719971 freshwater SPONGES OF WESTERN MONTANA 101

TABLE 3 chemical and physical ranges of western montana sponges

lakes containing spongesponges S lakes without E frafragiasfragilisfragihsgilis S lacustnslacustrislacustris E muellenmuelleri sponges variable nann99 nan7n 7 n 10 nan9n 9 methyl orange alkalinity eglmglgl 11211 2 1390139.0139 0 iso15015 0 80080.080 0 11211.211 2 1060logo106log 0 000.0oo0 0 2440244.0244 0 calcium eglmgl 202 0 39439.439 4 434.34 3 214 202.02 0 30 laI 1 040.40 4 4794747.99 total hardness as cac03 eglmgl 6 151 11 81 6 112 2 231 total organic carbon TOC eglmgl 07 51 05 65 00 65 000.00oo 0 14414.414 4 magnesium eglmgl 080 8 12812.812 8 00 67 000.00oo 0 90bgob9 ohl015 020.20 2 27027.027 0 ph 7067.067 06 8838.838 83 7627.627 62 8398 39 7067.067 06 8268 26 611 826 sioa eglmgl 323.23 2 iti17117.117 1 212.1bl812 1 379a37 9 212.12 1 379a37 93 494.94 9 15415.415 4 conductivity nscmgscm 21 355 26 237 38 2372373 9 420 elevation m 884 2124 1006 2911 884 2537 640 2926 temperature C 15815.815 8 23423.423 4 13013.013 0 23423 4 15615.615 6 23423 4 13013.013 0 23423.423 4 ogreateragreatergi eater than previously recorded see text blessless than previously recorded see text

S lacustrislacustris are present throughout the cold of the measured chemical factors in diamond temperate region of the northern hemisphere lake were not unusual although values for penny and racek 1968 hardness calcium and magnesium were near although only 3 species were identified each the upper ends of the ranges we determined showed considerable morphological variation from E frafragilisfragulisgilis coneoncontainingcontamingtabingtaming sites the variation encompassed spicule size in all 3 another type of malformation occurred in identified species spicule shape in E frafragilisfragulisgilis gemmoscleres of the specimens of E muelleri and E muelleri and spine development in E collected from cedar lake and a nearby pond muelleri variations of spicule morphology are here gemmoscleres had misshapen rays and of concern since taxonomic classification rests at least I1 spine extended from their shafts largely on this feature poirrier 1974 described similar malforma- spicule size ranges for each species identi- tions in a related species ephydatiafluviatilis fied were expanded beyond those reported pre- and observed that they can be induced by viously penny and racek 1968 ricciardi and altering environmental conditions suggesting reiswig 1993 the broader range of mega- that the presence of malformed gemmoscleres sclere lengths in E frafragilisfragulisgilis was especially pro- in cedar lake might be due to the environ- nounnouncedced some megasmegasclerescleres observed were ment since no chemical or physical data are 120 gm longer and others were 70 gmim available for cedar lake we do not know shorter than those previously recorded short whether unusual conditions exist in this lake thin megasmegasclerescleres and long megasmegasclerescleres were finally smooth megasmegasclerescleres or megasmegasclerescleres sometimes found in the same specimen having few spines were common in E muelleri atypical spispicklesspiculescules with bulbs along their throughout western montana both penny and lengths were present in I1 specimen of E frag racek 1968 in their taxonomic revision of the ilis collected from diamond lake simpson spongillidae and ricciardi and reiswig 1993 and vaccaro s 1974 report that germanium in their description of the sponges of eastern inhibits silica deposition during spicule forma- canada state that E muelleri only rarely has tion and can cause spispiculesspicklescules with bulbs to form smooth megasmegasclerescleres williams 1977 however suggests that these malformations might have reported that in colorado E muelleri frequent- an environmental basis since other specimens ly has smooth megasmegasclerescleres A high frequency of of E frafragilisfragulisgilis in the same area of the lake had smooth megasmegasclerescleres therefore might be char- normal megasmegasclerescleres however the malforma- acteristicacteristic of this species in the mountain west tions must either be due to highly localized most chemical and physical ranges deter- conditions or have a genetic basis the levels mined from sponge containing lakes fell within 102 GREAT BASIN naturalist volume 57 limits established in previous reports jewell technical assistance and dr jean smith for 1939 harrisonharrlsonhaggson 1974 williams 1977 excep- her encouragement and many suggestions tions were the ranges of conductivity and of calcium magnesium and silica concentrations literature CITED determined for E muellenmuelleri and the range of silica concentration for S lacustnslacustrislacustris CLIFFORD H FE 1991 porifera sponges pages 17 33 in aquatic invertebrates of albertaofalberta university ofalbertaalbernaalbennaofalberta in her study of colorado sponges williams press edmonton alberta canada 1977 observed that broadest chemical ranges FRANCIS J C AND FE W HARRISON 1988 copper and were associated with E frasfaafafragtlisfragifisfragulisgifis whereas the zinezinc toxicity in ephydatia fluviatilisfluviatilis transactions of ranges of alkalinity calcium and magnesium the american microscopical society 107 67 78 FROST T M 1991 porifera pages 95 124 in J H thorp concentrations and total hardness determined and A P covich editors ecology and classification for S lacustnslacustrislacustris were significantly narrower than of north american freshwater invertebrates acade- those for E fraataarafragifisfragilzsgifis or E muemuellenlleTi even though micmie press new york the chemical ranges determined for these GILBERT J J AND T L SIMPSON 1976 gemmule polymor- phism in the freshwater sponge spongilla lacustnslacustrislacustris species were in several cases broader in west- archibarchivarebiv fur hydrobiologie 78 268 277 ern montana than in colorado similar rela- HARRISON FE W 1974 sponges PonPoriponfieraporifierafieraflera spongillidae tiontionshipsships among the widths of their ranges were pages 29 66 in C W hart jr and S L H fuller edi- pollution fragilisfragulisgilis tors ecology of freshwater invertebrates observed E fra in western montana as in academic press new york colorado had the widest chemical ranges JEWELL M E 1935 an ecological study of the fresh- overall in western montana total organic car- water sponges of northeastern wisconsin ecological bon and silica were exceptions and S lacus monographs 5 463 504 1939 an ecological study of the freshwaterfresh ans had the narrowest ranges of alkalinity cal- water tnstris sponges of wisconsin II11 the influence of calcium clumcium and magnesium concentrations and total ecology 33 11 28 hardness from the 2 studies it appears as if OLD M C 1932 environmental selection of the freshwater sponges spongillidae of michigan transac- E fragilisfragulisgilis in the mountain west is less con- fra tions of the american microscopical society 51 strained by chemical factors than the other 2 129 136 species are and that S lacustnslacustrislacustris is more lim- PENNAK R W 1978 porifera sponges pages 80 98 in ited by such factors these conclusions need to freshwater invertebrates of the united states 2ndand be substantiated by analyses involving larger edition john wiley and sons inc new york PENNEYPENNEYJJ TANDAT AND A A RACEK 1968 comprehensive revi- sample sizes sion of a worldwide collection of freshwater sponges the fact that sponges were absent fromhrombrom some pooporiferaPonferarera spongillidae united states natural his- lakes having measured chemical and physical tory museum bulletin 272 1 184 profilesfilesflies almost identical POIRRIERFOIRRIER M A 1969 louisiana freshwaterfresh water sponges tax- pioplo to those of lakes con- onomy ecology and distribution unpublished dis taining sponges suggests that sponge distribu- sertionbertionsertion louisiana state university baton rouge tion is influenced by more factors than those university microfilms inc ann arbor MI 110 appp we examined in fact as williams 1977 pointed 1974 Ecoecomorphiccecomorphicmorphic variation in gernmoscleresgemmoscleres of ephydatiafluviatilisephydatia tilis poriferaferahera distribution fluviafluviatilis linnaeus bonPon spongillidae out is probablypioplobablybabiy controlled by chem- with comments upon its systematics and ecology ical physical and biological factors acting syn- hydrobiologia 44 337 347 ergistically the identification of species of POIRRIER M A P S MARTIN AND R J BAERWALD 1987 morphology freshwater sponges in western montana and comparative of microsclere structure in spongilla alba S cecotacenota and S lacustnslacustrislacustris fonporiferaPonferarera the initial chemical and physical characteriza- spongillidaetransactionsspongillidae transactions of the american micro- tion of their habitats lay the foundation for scopicalscopical society 106 302 310 investigations of the effects of additional fac- RICCIARDI A AND H M REISWIG 1993 freshwater porifera tors on sponge distribution sponges fonPonferahera spongillidae of eastern canada taxonomy distribution and ecology canadian journal of zoology 71 665 683 acknowledgments RICHELLE MAURERMAURFR E Y dfcoudfnnedegoudenne G VAN DE VYVER AND L DEJONGHE 1994 some aspects of heavy metal pages we thank bahls of the montana tolerance in freshwater sponges 341 349 in dr loren R W van soest et al editors sponges in time and department of environmental quality for his space proceedings of the ath4th international porifera advice and for securing funding for the water congress A balkema rotterdam analyses bob bukantis also of the department SIMPSON T L AND C A VACCARO 1974 an ultrastructuralultrastructural study of silica deposition in the freshwater sponge of environmental quality and the staff of the spongilla lacustnslacustrislacustris journal of ultrastructuralUltra structural research montana state chemistry laboratory for their 4729647 296 309 199719971 freshwater SPONGES OF WESTERN MONTANA 103

STREKAL T A AND W FE mcdiffettmcdlffett 1974 factors affect- WILLIAMS R E 1977 distribution ecology and reproduc- ing germination growth and distribution ofthe fresh- tive cycles of coloradoofcolorado freshwater sponges fonPonporiferaferahera water sponge spongillafragilisspongilla fragiasfragihs leidy poriferaPonfera bio- spongilllidae unpublished dissertation university of logical bulletin 146 267 278 colorado boulder university microfilms inc ann UNITED STATES environmental protection AGENCY 1983 arbor MIM 1 155 appp standard methods for chemical analysis of water and 1980 the freshwater sponges fonPonporiferaferahera spongilli- wastes cincinnati OH 275 appp dae of colorado natural history inventory of colo- 199iggi19911 methods for the determination of metals in rado 4 1 12 environmental samples office of research and devel- YOUNG R T 1935 the life of flathead lake montana opment washington DC 339 appp ecological monographs 5 93 163 WATER QUALITY BUREAU 1991 field procedure manual montana department of health and environmental received 5 august 1996 sciences helena pages 313 1 3213 21 accepted 31 january 1997 gleatgreat basin naturalist 572 0 1997 appp 104 115

BOGGY MEADOWS LIVESTOCK GRAZING AND interspecific interactions influences ON THE INSULAR distribution OF MONTANE LINCOLNS SPARROWS MELOSPIZA LINCOLNII ALTICOLA

carla cicerolciceroacicero1

ABSTRACT I1 surveyed 34 meadows in california and oregon to count lineolLincollincolnrss sparrows melospiza lincolniilincolmilinclincollincolniaolmigimigimonii alaltialtoaitocolfcoldcoidticola and to identify habitat features that might influence their local insular occurrence lineolLincollincolnrss sparrows were found at 72 of the sites surveyed counts of singing males were low and uncorrelated with meadow size lineolLincollincolnrss sparrows were most common in wet meadows with little damage by grazing singing males were concentrated in flooded or boggy areas near meadow edges where pines pinus sp provided elevated perches for singing and vigilance patches of willows salix sp were often present nearby numbers of male lineolLincollincolnnss sparrows were strongly and negatively correlated with abundance of sympatric song sparrows M melodiaftsherellamelodia fisherellafisgisggsherella lineolLincollincolnrss sparrows breeding inm montane meadows are potentially vulnerable to local extirpation because of their insular distribution low population density and fluctuating habitat conditions heavy damage from livestock grazing drastically increases the probability of local extirpation

key words melospiza lincolniilincolnnlincollincolynniinilnn lincoln s sparrow montane meadow insular populations habitat association livestock grazing conservation biology melospiza melodia song sparrow

meadows form ecological islands through- fragments eg forman et al 1976 however out montane forests of the western united detailed studies of specific taxa occupying nat- states although these systems generally sup- ural insular habitats such as meadows are port a rich avifauna they are highly variable in scarce terms of physiographic hydrologic edaphic the montane form of lineolLincollincolnrss sparrow vegetative and floristicflon stic characteristics ege g melospiza lincolniilincollincolnianii alticola miller and mccabe see kuramoto and bliss 1970 benedict and 1935 american ornithologists union 1957 is major 1982 ratliff 1982 1985 alienallenailen 1987 well suited to such a study this taxon breeds grazing and fire history also shape meadow insularly in particular kinds of meadows from environments kuramoto and bliss 1970 oregon to california and from idaho to new debenedetti and parsons 1979 1984 parsons mexico such sites are separated from other 1981 ratliff 1985 changes in grazing and suitable islands of habitat by unoccupied coni- fire management practices combined with cli- ferous forest two other subspecies M I1 lin mate further influence the ecology and stability colniicolnai and M I1 gracilis occur more broadly of meadows by promoting invasion of lodge- in brushy bogs from central alaska through pole pine and other conifers franklin et al canada to the northern contiguous united 1971 dunwiddie 1977 vale 1981a 1981b states despite the widespread distribution of ratliff 1985 Linclincolnolisolyss sparrows the species has been poorly because of their mobility birds respond studied compared with either of its convenerscongenerscongeners quickly to habitat change and thus are model the song sparrow M melodia or swamp organisms for illustrating the effect of habitat sparrow M georgiana information on the dis- on the distribution and abundance of insular tributiontribution and natural history ofmof M I1 alticola is as well as continental populations cody 1981 especially lacking in this study I1 provide base- wiens 1989 numerous researchers have exam- line data on occurrence abundance habitat ined total avifaunal distribution and abundance association and other factors potentially related on montane islands ege g johnson 1975 kratter to their local distribution in western montane 1992 lentz 1993 in natural habitat patches meadows secondarily I1 evaluate the potential ege g aspen flack 1976 or in disturbed forest impact livestock grazing has on this taxon

I1museum of vertebrate zoology university of california berkeley CA 94720

104 199719971 LINCOLN S SPARROWS IN MEADOWS 105 range ecologists have shown unequivocally dance may reveal degradation of meadows by that grazing occurs unevenly across montane livestock vegetation types and that meadows and other riparian areas receive disproportionately heavy STUDY AREAS AND METHODS use relative to their total acreage cook 1966 roath and krueger 1982a 1982b gillen et al I1 surveyed meadows for lineolLincollincolnrss sparrows 1984 platts and nelson 1985 numerous stud- from mid may to early july 1987 1989 A total ies have assessed the impact of such use on of 34 meadows belonging to 29 systems were riparian habitats and associated wildlife eg visited including I1 in northern oregon and 28 leege et al 1981 kauffman et al 1983 kauff- in california from the southern cascade moun- man and krueger 1984 taylor 1986 ohmart tains lassen county through the sierra nevada 1994 montane lineolLincollincolnnss sparrows are poten- to the san bernardino mountains fig 1 ele- tially vulnerable to disturbance by heavy graz- vations ranged from 1365 to 2470 m with ing because of their tendency to nest on or near lodgepole pine pinus contorconcontortdcontortacontortedtottatortatd forest dominat- swampy ground in wet meadows grinnell ing the surrounding vegetation size land own- and miller 1944 austin 1968 consequently ership and type and intensity of livestock graz- changes in their local occurrence or abanabun ing varied among meadows although several

120

rxarx r 0 400 r x lp 4 2 676 7 8128 12 13 140 015 016olg 171817 18 21 192019 20 wiemwmem 23 24 0u25 26 soon

350 a

27go28

too100 mi

2 00 km

fig 1 breeding distribution of lineolLincollincolnrss sparrows in california the general locations of 28 meadow systems surveyed in california are indicated an additional meadow in oregon is not shown closed circles denote meadows where Linclincolnolyss sparrows were present open circles meadows where lineolLincollincolnnss sparrows were absent closed squares show other known breeding localities based on specimens deposited in the museum of vertebrate zoology berkeley california published records grinnell and miller 1944 lentz 1993 records obtained during a survey of meadows for willow flycatchers empidonax trailletrailtrailhitraillnlnhi M A flett and J harris unpublished data 106 GREAT BASIN naturalist volume 57 groups of meadows were clustered geographi- grazing surveys took longer in large meadows cally differences in habitat characteristics ele- or at sites with high numbers of lineolLincollincolnnss vation andor grazing regime occurred between sparrows because the amount of time spent even the most proximal sites consequently at each site varied count values were stan- each meadow was treated as an independent dardizeddardized by dividing the number of singing sample point twentyfivetwenty five sites consisted of sin- males observed per visit by the length of the gle meadows without any connection to other survey As with all count methods some silent sites another meadow beasore meadow site males may have been overlooked thus these 22 was divided by a fence into 2 parts with counts represent minimal estimates of total strikingly different grazing regimes because abundance the 2 sides also contrasted dramatically in the geographic scope of this study pre- abundance of Lincollincolnrss sparrows they were cluded surveying all meadows simultaneously separated for purposes of analysis and discus- to verify the reliability of counts conducted at sion three meadow systems lacey 818 hay different times I1 surveyed 17 meadows 50 press creek 9 and sagehen creek 12 con- twice or more during the same or subsequent tained multiple meadows within I1 drainage or seasons counts of singing males in the same basin that were separated from each other by meadow at different stages of the breeding a distance of at least 080.8 km because these cycle were identical because annual climatic were visited only during the breeding season differences might also influence counts taken when males were singing and thus territorial in different years I1 obtained data on mean presumably there was no movement of lineolLincollincolnnss temperature and precipitation during may and sparrows between meadows this was con- june 1987 1989 national oceanic and atmos- firmed by multiple visits to the same meadow pheric administration 1987 1988 1989 from system eg harpresshaypressHaypress creek during a single weather stations located near 3 main clusters breeding season when individual singing males of meadows 1 northern sierra nevada sage could be identified repeatedly by their loca- hen creek nevada county california 1932 m tion in each meadow 2 central southern sierra nevada hunting- to ensure breeding status counts were lim- ton lake fresno county california 2140 m ited to singing males Lincollincolnnss sparrows sing 3 san bernardino mountains big bear lake vigorously from elevated perches during the san bernardino county california 2070 m I1 breeding season particularly near watercourseswatercourses analyzed these data by analysis of variance or along the meadow edge and thus are easily using statviewStatview for the macintosh abacus con- detectable numbers of singing males were cepts 1988 with the exception of mean june counted by traversing each meadow and record- temperature which was slightly different among ing their presence and location because my years P 003870.0387 there was no annual effect goal was to survey a broad range of sites in on temperature or precipitation P 0050.05oos I1 order to evaluate the kinds of meadows inhab- counted similar numbers of lineolLincollincolnrss spar- ited by montane lineolLincollincolnnss sparrows lengthy rows at the same meadow in different years and repeated visits to individual meadows in a daily field journal I1 recorded numbers were not possible however the size and dis- of other singing birds at each meadow and crete linear configuration of most meadows provided detailed descriptions and sketches of enabled complete surveys of all singing males the meadows I1 also took notes on the charac- during I1 or at most 2 consecutive mornings teristicste of meadows with and without lineolLincollincolnrCss 2 5 hmhmorningorning thus standard census tech- sparrows and on the location of singing males niques appropriate for expansive areas of non relative to the meadow edge and to habitat insular habitat were unnecessary features such as extent of flooding presence or complete counts of singing males were ob- absence of willows salix sp presence or ab- tained at 26 76 of the 34 meadows partial sence of corn lily veratrum sp and presence surveys were conducted at the remaining 8 sites or absence of pines pinus sp the first 3 because of their large size andor because habitat variables flooding willows corn lily fencing restricted access nonetheless these are presumably important for breeding grin- estimates still provide valuable information re- nell and miller 1944 austin 1968 speirs and garding the occurrence of lineolLincollincolnrss sparrows speirs 1968 although published accounts do six of the 8 sites were visibly impacted by not provide information on the use of edges 199719971 LINCOLN S SPARROWS IN MEADOWS 107

TABLE 1 rating scheme used for characterizing meadows based on wetness and extent of grazing damage score characterization WETNESS I1 meadow very dry no standing water or boggy ground single well defined stream channel 2 less than 25 of meadow wet few areas of standing water or boggy ground 3 25 50 of meadow wet some flooded or boggy areas other areas dry 4 50 75 of meadow wet many areas of standing water or boggy ground some rivulets of running water 5 75 100 of meadow wet most or all of meadow covered with standing water andor rivulets of running water GRAZING I1 meadow essentially pristine no bare ground exposed grassy undercut streamstreambanksbanks no evidence of gullying andor bank erosion few to no signs of livestock 2 slight grazing damage 25 of meadow with bare ground exposed slight gullying andor streamstreambankbank erosion low density of livestock droppings and trails 3 moderate grazing damage 25 50 of meadow with bare ground exposed gullying andor streamstreambankbank erosion clearly evident low to moderate density of livestock droppings and trails 4 heavy grazing damage 50 75 of meadow with bare ground exposed pronounced gullying andor streamstreambankbank erosion moderate to high density of livestock droppings and trails 5 meadow severely damaged 75 of meadow with bare ground exposed extreme gullying andor streamstreambankbank erosion very high density of livestock droppings and trails andor pines observations of singing males sug- meadows caused by livestock grazing see fig gest that these features might be equally im- 2 was assessed subjectively and also rated on porportanttant because of the broad geographic sam- a scale of 1 to 5 taking into account the amount pling and concomitant variability in meadow of bare ground exposed extent of gullying and type data on the herbaceous composition of or streamstreambankbank erosion presence of old or recent each meadow were beyond the scope of this livestock droppings and network of livestock study differences in habitat features associated trails table 1 As with wetness scores were with individual males n 75 were tested sta- assigned based on overall damage observed tistically using a parametricnonparametricnon binomial test during the period of study although gradient with the normal approximation for N 35 see andor soil type also may contribute to differ- siegel and castellan 198838 44 ences in erosion seen between meadows im- A complex classification system has been pacts from grazing clearly had a major effect developed for meadows in the sierra nevada evaluation of the effects of grazing on plant incorporating similarities in physiographic species composition and diversity was beyond hydrologic edaphic vegetative and floristic the scope of this study both wetness and graz- characteristics ratliff 1985 however only in ing damage may vary with changes in grazing a few situations have enough sites been studied practices to adequately define the classification series areas of meadows were estimated from ratliff 19859 because of the close associa- USGS 75 minute topographical maps using a tion between breeding lineolLincollincolnnss sparrows and point grid system modified for a scale of boggy or flooded ground I1 used a simpler 124000 these data were supplemented by approach to rate meadows on a scale of I1 to 5 US forest service data where available for according to wetness table 1 scores assigned large meadows where counts are incomplete to meadows reflect the wetness characteristics both the total area and the area surveyed were observed during the period of study higher estimated or lower ratings may be more appropriate at other times depending on seasonal and annual RESULTS variability in hydrologic regimes distribution and abundance quantitative information on livestock use in of s sparrows each meadow was difficult to obtain because of lineolLincollincolnrs variability in landownershipland ownership patterns and be- I1 found lineolLincollincolnnss sparrows in 26 76576576.5 of cause stocking rates determined for entire the 34 sites surveyed table 2 absolute num- grazing allotments do not accurately reflect the bers of breeding males varied from I1 to 16 concentration of livestock on meadows and which translates to standardized counts of 020.2 4 other riparian areas consequently damage to malesh counts were low in most meadows 108 GREAT BASIN naturalist volume 57

1 1 for TABLE 2 characteristics of meadows surveyed for M I1 alticolaficola inm california 28 and oregon 29 see figure I locations in california number of total area time per number of singing males grazinggrazing area surveyed number of survey singing per survey wetness damagee type of gd1 meadoomeadowmeatlow ha ha surveystsurveysurveysst h males hour schrecscorecscore score grazingrazinggrazing1grazingd

1 battle creek meadows 3 4 1460 in P 511 447 1 10 0 00 cattle 2 grass lake 1 grazed 1980 m NF 5 5 1 2 3 15 5 not 3 church meadows 2040 m P 6 6 2 5 11 22 3 2 sheep 4 french meadows 2 3 2035 in P 16 16 1 3 6 20 cattle 5 lincoln valley 6 2220 m P 26 26 3 4 16 40 4 1 CattcattieCattlecattleecattle6cataleeleeiee 6 cottonwood creek 1770 m NF 52 52 2 4 0 00 4 2 sheep 7 perazzo meadows 2010 m P 369 59 1 5 0 00 3 4 cattle 8aaa lacey valley 2070 m P 164 120 1 5 1 02 2 3 sheep ab8b little lacey valley 2085 m P 42 42 1 2 3 15 3 4 sheep 9aaa upper harpresshaypressHaypress creek grazed 2195 m NF 20 20 2 2 4 20 4 2 not ab9b middle harpresshaypressHaypress creek 1 not grazed 21102110mnfm NF 8 8 3 2 3 15 3 9cac lower harpresshaypressHaypress creek grazed 2035 m NF 26 26 2 3 2 07 2 2 not 10 coppins meadow 2070 m P 164 60 2 4 8 20 3 3 cattle 11 W of coppins meadow 2070 m P 9 9 2 2 4 20 4 1 cattle 12a upper sagehen creek 2 2 sheep 2050 in NF 14 14 2 3 2 07 12b lower sagehen creek 2010 m NF 54 54 2 4 1 03 4 1 not glazedgrazed 13 austin meadow 2070 m NF 6 6 1 3 2 07 2 3 cattle 14 leek springspong valley 2 Cattle 6 2255 m P 56 49 1 2 4 20 3 cattieCattleeiee 15 pleasant valley 1850 m P 207 51 1 2 0 00 1 4 cattle 16 swauger canyon 2390 m NF ft 23 1 2 0 00 4 1 sheep

2 malesh were recorded in 16 61561.5 of the species was absent from all meadows with 26 meadows 2 3 malesh in 8 meadows and heavy grazing pressure score 4 the pres- 3 4 malesh in only 2 meadows numbers of ence of Linlincolncohss sparrows at these 2 sites is singing males were highest at church mead- accounted for by the method of analysis in ows 3 and lincoln valley 5 in the northern which scores were assigned based on overall sierra nevada bluff lake 27 and metcalf appearance of the meadow thus all 3 males creek 28 in the san bernardino mountains at little lacey valley were concentrated in the and hood river meadows 29 in northern lower wetter portion of the meadow where oregon counts were not correlated with impacts from grazing were slight none was meadow size r 02400.240 FP 0050.05oos observed in the more heavily damaged upper distribution and abundance were clearly reaches likewise the single male in upper associated with scores for wetness andor extent beasore meadow occurred at the lower edge of grazing damage fig 3 lineolLincollincolnrss sparrows of the site which was in good condition rela- were most common in moderately wet to very tive to the meadow as a whole As expected wet ie flooded meadows with low levels of lineolLincollincolnnss sparrows were absent from the single grazing damage except for little lacey valley site that lacked standing water and showed pleasant 15151 ab8b and upper beasore meadow 22a the signs of heavy grazing valley 199719971 LINCOLN S SPARROWS IN MEADOWS 109

TABLE 2 continuedContiDued number of total area time per number of singing males grazingerazmgrazmg area surveyed number of survey singing per survey wetness damagee type of meadoomeadowmeaelow1lowa ha ha surveys h males hour scoieschieschrecscorec score grazinggrazgrazingligrazinglyingli 17 white wolf 2380 rn NP 6 6 1 2 0 00 3 1 not grazed 18 ackerson meadow 1400 m P 139 61 1 3 0 00 2 4 cattle 19 hogdon meadow 1400 m NP 30 30 1 4 3 08 4 2 not gigrazedazedabed 20 crane flat 1890 m NP 22 22 1 3 2 07 3 1 not glazedgrazed 21 porcupine flat 2470 m NP 6 6 1 2 0 00 2 1 not grazed 22a upper beasore meadow 2088 m P 16 16 2 3 1 03 2 5 cattle 2213 lower beasore meadow 2057 m P 15 15 2 3 5 17 3 2 not grazedgrazed91 23 poison meadow 1740 in NF 10 10 2 2 2 10 3 2 cattle 24 markwood meadow 1800 in NF 45 45 1 3 4 13 4 3 cattle 25 dinkey meadow 1680 m P 37 37 2 4 6 15 4 2 cattle 26 long meadow 2100 mnfm NF 15 15 2 4 3 08 3 3 cattle 27 blufflakebluff inflake 2315 m NF 7 7 2 3 8 27 3 1 not grazed 28 metcalfcreekmetcalf creek 2225 m NF 7 7 2 4 11 28 4 1 not grazed 29 hood riverbivelbiver meadows 1365 m NF 12 12 1 3 11 37 5 1 not grazed

multiple meadows within a single system aieare designated alphaalphanumencallyalphanumericallynumerically letters min parentheses indicate dominant ownership P private NF national forest NP yosemite national paikpark specific localities and dates of surveys are available from the author purveys were conducted on 1 2 consecutive days this was sufficient to count every singing male min all of the areas surveyed see methods rated anon a scale of I1 to 5 see table 1 determined flomfrom fieldfieldworkwoik and USU S foiestforest service data applies only to this study period ungrazed meadows may have been grazed in prior years dataodata unclear in lincoln valley I1 found no evidence ofrecentof recent grazing although the tahoe national forestfoiest lists this meadow as partpaitoftheoftheodthe lincoln valley allotment information was not available foiforbol little lacey valley leek springs valley or01 lowerlowet beasore meadow little lacey valley was obviously grazed presumably by sheep because of sheep grazing in adjacent lacey valley leek springs valley did not appear to have lecentrecent grazing activity althoualthoughgligil thetilctile general area has numerousnume iouslous glazinggrazing allotments and this meadow probably receives some impactit from cattle lower beasore meadow also did not appear to be grazedglazed in contrast to the heavily glazedgrazed upper beasore meadow see fig 2 fnotenotnot estimated because numerous small patchy openings occur along the canyon bottom

despite this general tendency the distribu- habitat features associated with tion of Linlincolncobiss sparrows showed a more com- individual male lineolLincollincolnrss sparrows plex example I1 failed to find the pattern for singing males were strongly associated with in several meadows with fairly high species particular habitat features fig 4 the most scores for wetness and low scores for grazing important attribute was the presence of nearby damage cottonwood creek 6 swauger can- surface water only 3 of the males were ob- yon igl16116 white wolf 17117 although abun- served in areas of dry ground while 93 were dance was highest in the wettest most pristine boggy 542 flooded 38938.9 meadows eg lincoln valley 5 metcalf creek seen in either 54254.2 or 389 a difference significant at FP 00010.001 28 hood river meadows 29291 other equally sites of small wet sites had notably fewer males eg upper numerous locations had networks coursed harpresshaypressHaypress creek aa9a west of coppins meadow narrow channels with runing water that through tussockstussockysocks of sedges grasses or other 11II11 lower sagehen creek 12b hogdon tus meadow 19 markwood meadow 24 dinkey herbaceous plants the presence of willows meadow 25 meadows with lower wetness corn lily and pines especially P contortscontorta also scores also supported relatively high numbers appeared to be important attributes of Lincollincolnrss of lineolLincollincolnrss sparrows as long as grazing dam- sparrow habitat approximately 84 of all males age was fairly low eg church meadows 331 occurred near clumps of willows P 00010.001 coppins meadow iollol10110 bluff lake 27197127 and 59 were in areas with at least scattered llolio110 GREAT BASIN naturalist volume 57

P W

E Y Q

u 4

4 14 t V

F J r j ace-

v 4kavf

571 ip j

X rv

figpigiigaig 2 views of the lower top and tipperlipperupper bottom portions of beasore meadow madera county california site 22 photostos illustrate 2 extremes in the condition of meadows surveyed in this study lower beasore meadow which was notno grazedzedged had an uneroded creekcreekbankbank swampy ground rich herbaceous cover and scattered patches of willow sahersabersalixsafer sp upperper beasore meadow showed severe soil erosion compaction and dessicationdessication due to cattle grazing with 3 7 in gully upper ing lincoln s sparrows were fairly common in lower beasore meadow but essentially absent from beasore meadow song sparrows were abundant in the willows in upper beasore meadow 199719971 LINCOLN S SPARROWS IN MEADOWS lillii111

4 floodedboggyfloodedtboggyFlooded Boggy dy

assmrs mss comcorncormcomm lily inono cornconncormcomm lily 2 isam MM willows ismmiSno willows B r pines 2aaa siI1 no pines 9 J

21 r ecotonal I1 non ecotonal B 17 inon 12 9 15 16 A 6 0 25 50 75 100 7 2 of singing males associated with 2 3 different habitat characteristics raza raz7g fig 4 proportion of singing male lincoln s sparrowsspan awsows s6csaciqrnclge associated with different habitat attributes ecotonal males were those singing along the edge between the meadow and adjacent coniferous forest all pairwisepahpan wise com- fig 3 three dimensional graph illustrating the relative parisonspari sons except for the presencepresenceabsenceabsence of corn lily abundance vertical axis counts per hour of singing male were significant at FP 0010 01 lincoln s sparrows in 29 meadows systems scored according to wetness and grazing damage see table 1 meadows are numbered as in table 2 circles indicate sites with lineolLincollincolnrss sparrows squares sites where lincolnLincohss tufts of grass andor other plants approxi- sparrows were absent mately 35 of the nests in the WFVZ records were situated under a patch of willows negativeassociationnegative association between patches of lily although this proportion was distribution and abundance of Lincollincolnnss nonsignificant P lineol 008230.0823 however unlike sparrows and song sparrows song sparrows which were observed only in areas with willow lineolLincollincolnrss sparrows were not counts of singing male lineolLincollincolnrss sparrows limited to willow patches the concentration of and song sparrows showed a strong negative male linelincolnLincolisolyss sparrows was greatest near the correlation r 07010.701 P oolooi0010.01 fig 5 edges of meadows 67 of males P ooi0010.01 although some meadows had approximately where they were often seen perched or singing equal numbers of the 2 convenerscongenerscongeners eg dinkey in pines 68 of males FP ooi0010.01 although meadow little lacey valley lower beasore most lineolLincollincolnrss sparrows were observed within meadow most sites appeared to be dominated meadows singing males or pairs were occa- by one or the other species song sparrows siosionallynally seen in small nearby openings in sur- were common at several meadows where lin- rounding forest as long as suitable habitat was coln s sparrows were either absent battle present creek meadows cottonwood creek perazzo data from egg sets at the western founda- meadows pleasant valley swauger canyon tion ofvertebrateofvertebrate zoology WFVZ also revealed and ackerson meadow or rare lacey valley the importance of wet ground and clumps of sagehen creek upper beasore meadow these herbaceous vegetation or shrubs for breeding sites included dry heavily grazed meadows as of 65 records from california 56 contained well as wet fairly pristine areas that otherwise information on moisture characteristics at the looked suitable likewise song sparrows were nest site and 54 96 of these indicated damp absent or rare from several meadows with rel- to very wet conditions over 90 of the nests atively large numbers of lineolLincollincolnrss sparrows were placed on the ground or slightly above- eg lincoln valley coppins meadow metcalf ground where they were well concealed by creek hood river meadows as well as from 112 GREAT BASIN naturalist volume 57

20 of singing males combined with descriptions r 07010.701 of nest sites from WFVZ egg data slips indicate that combinations of the following attrib- 15 utes are important for breeding boggy or cz 2 flooded ground thick groundcoverground cover of herba- ca ceous vegetation often with raised tustussockstussockysocks of CL 0030ouy live or dead grasses or sedges patches of corn L 10 SD w lily willow thickets or other low shrubs and 0.0 c E 75 some conifers raised clumps of herbaceous 0 vegetation are probably critical for breeding ZE11 5- II1 under such wet conditions as suggested by the nest site descriptions likewise dense herbaceous plant material in conjunction with 0 J 00 0 willows and corn lily may provide important 1 1 1 1100 1155 2200 concealment the association between male lineolLincollincolnnss sparrows and pines undoubtedly number of male reflects the importance of elevated perches for song sparrows singing and vigilance although Lincollincolnrss sparrows were present fig 5 negative correlation between number of singinsinging9 in the majority of wetmeadowswet meadows studied their male lineolLincollincolnrss sparrows and song sparrows absence at certain sites that otherwise looked suitable deserves discussion one example is white wolf 17 in yosemite national park other sites where lineLinclincolnolisolyss sparrows were less granholm reported common west of coppins meadow austin where beedy and 1985190 meadow leek spring valley the species in summer but did not present any grinnell and negative association between these 2 dates of nesting although storer the sparrows species was also evident when comparing 1924471 noted that lineolLincollincolnnss arrive yosemite may counts between meadows within a single sys- in the region by mid this date may apply to lower elevation meadows in tem for example whereas lineolLincollincolnrss sparrows outnumbered song sparrows at upper hay yosemite valley white wolf is one of the high- press creek 4 versus 2 males respectively est 2380 inm meadows surveyed in this study song sparrows were slightly more numerous and it is possible that the timing of my visit in the arrival of this than lincoln s sparrows at middle harpresshaypressHaypress early june preceded species creek 6 versus 3 males and noticeably more for breeding however examination of museum abundant at lower harpresshaypressHaypress creek 8 versus records MVZ WFVZ showed that lineolLincollincolnnss 2 males similar patterns were observed within sparrows already have nests with eggs by early single sites although approximately equal num- to mid june at sites of similar or higher eleva- bers 3 males of the 2 species were observed tion both in the central sierra nevada and at little lacey valley for example lincoln s elsewhere furthermore I1 observed lineolLincollincolnrss sparrows occurred in the wettest portion of sparrows singing in mid may at other high the meadow while song sparrows were found elevation meadows such as bluff lake and only in the drier more heavily grazed areas metcalf creek when temperatures were cold similarly although both Lincollincolnrss sparrows and snow was still present on the ground and song sparrows were fairly common in although these 2 meadows were visited in a lower beasore meadow 5 and 3 males re- different year than white wolf the lack of a spectspectivelyively the former species dropped out in significant annual difference in climate during upper beasore meadow while song sparrows the period of study suggests that timing alone increased significantly in abundance to 14 males cannot account for the disparity additional surveys are needed to determine the popula- discussion AND conclusions tion status of lineolLincollincolnrss sparrows breeding at white wolf of the 34 meadows surveyed lincoln s another wet meadow where I1 failed to find sparrows occurred only in sites with certain Lincollincolnrss sparrows was swauger canyon 16 habitat features data on the specific locations northeast of yosemite national park in the 199711997 LINCOLN S SPARROWS IN MEADOWS 113 sweetwater mountains according to johnson and in all cases song sparrows instigated the 1975 the species has never been found to chase displacing male Lincollincolnrss sparrows sing- nest in that range despite extensive fieldwork ing from elevated posts although there is no there by parties from the museum of verte- evidence that lincoln s sparrows and song brate zoology more puzzling was the absence sparrows are interinterspecificallyspecifically territorial addi- of lineolLincollincolnnss sparrows along cottonwood creek tional behavioral and ecological studies are 6 and lower sagehen creek 12b in the north- needed to understand the underlying factors ern sierra nevada especially since the species responsible for the negative association observed breeds regularly at other comparable meadows between these 2 convenerscongenerscongeners on both a local and in the same region subsequent visits to these regional scale removal experiments in which 2 sites have confirmed the results of earlier song sparrows are excluded from boggy mead- counts both meadows had large areas that ows within the range of lineolLincollincolnnss sparrows were flooded by beaver castor canadensis would especially shed light on the role of inter- activity during the period of study unlike lower specific competition if any in controlling lin- sagehen creek however the meadow along coln s sparrow distribution andor abundance cottonwood creek is grazed by sheep during spatial or temporal fluctuations in the dis- the summer with the season of use occurring tributiontribution and abundance of certain bird species from mid june through september S FE bishop may indicate short term or longtermlong term trends in personal communication although damage climate resource availability and habitat qual- caused by sheep eg trampling of herbaceous ity such effects are probably most pronounced vegetation browsing of willows may be suffi- in populations occupying ecological islands cient to disrupt breeding of linelincolnLincolif s sparrows which may be in nonequilibrium dynamics along cottonwood creek it does not explain johnson 1975 1995 species with narrow their rarity along lower sagehen creek habitat requirements are especially useful as in contrast to lineolLincollincolnnss sparrows song spar- indicators of trends because of their greater rows were among the most common birds seen vulnerability to natural or human induced at both of these meadows with abundance changes of the 4 species of sparrows occupy- higher than at most other sites surveyed diff- ing meadows in the sierra nevada cascade ferenerencesces in habitat choice and tolerance for mountains savannah sparrow Passerpasserculusculus disturbance may account at least partially for sandwichensis song sparrow lineolLincollincolnnss spar- the unexpected negative association between row and white crowned sparrow zonotrichia the 2 species at these and other meadows sur- leucophrysleucophrys lineolLincollincolnrss sparrows may be most veyed for example song sparrows were abun- susceptible to local extirpation because of their dant at several dry severely grazed sites that generally low population size and their restric- were unsuitable for lineolLincollincolnrss sparrows like- tion to wet or flooded areas although different wise heavily flooded areas such as hood river lines of evidence suggest that mountain mead- meadows may be shunned by song sparrows ows may be as temporally stable as the sur- only 1 singing male was observed of greater rounding environment benedict 1982 mois- interest than these extremes however are the ture characteristics of meadows are highly vari- patterns observed at intermediate sites which able depending on annual precipitation short were often dominated by one or the other spe- term fluctuations in precipitation may affect cies in fact lincolnLineoliscoliss sparrows were common habitat quality directly through snowbeltsnowsnowmeltmelt and at a number of wet meadows that appeared groundwater recharge wood 1975 andor similar in habitat to both cottonwood creek indirectly through availability of food eg and lower sagehen creek and where song insect resources cody 1981 johnson 1995 sparrows were surprisingly scarce one hypoth- in addition beaver activity can profoundly in- esis is interspecific competition acting in con- fluence the extent of flooding in meadows cert with differences in habitat use andor livestock grazing can alter natural hydro- tolerance for disturbance caused by grazing logic regimes by increasing runoff and exacer- speirs and speirs 19681440 noted that song bating erosion and gullying thereby lowering sparrows often utilized the same perches and the groundwater table eg upper beasore were able to compete strongly and very suc- meadow fig bli213 also see rauzi and hanson cessfullycessfully against lineolLincollincolnrss sparrows I1 observed 1966 lusby 1970 platts 1981 kauffman et al interspecific interactions on at least 5 occasions 1983 ratliffl985ratliff 1985 consequently grazing may 114 GREAT BASIN naturalist volume 57 eliminate potential nesting habitat for lineolLincollincolnrss and an anonymous reviewer for reading drafts sparrows in addition hydrologic and vegeta- of this manuscript and offering many useful tive changes associated with grazing can alter suggestions fieldwork was supported partially the distribution and abundance of more toler- by 2 frank M chapman grants from the amer- ant species such as song sparrows which may ican museum of natural history compete with lineolLincollincolnrss sparrows for territories and other resources these indirect effects of literature CITED livestock grazing combined with direct impacts such as reduction of cover and trampling of ABACAUS CONCEPTS INC 1988 statviewStatview version 103 berkeley CA nests undoubtedly have resulted in the in extir- AALLENN B H 1987 forest and meadow ecosystems in cali- pation of lincolnLincohss sparrows from some mead- fornia rangelands 9 125 128 ows because populations of this species are AMERICAN ornithologists UNION 1957 checklistcheck list of already vulnerable to natural fluctuations in north american birds and2nd edition lord baltimore in press moisture any further changes caused by graz- baltimore MD AUSTIN 0 L JR 1968 melospiza lincolniiimcolnulincolnialincolniinil alticola miller ing may exacerbate their probability of local and mccabe montane lineolLincollincolnrss sparrow pages extirpation 1467 1472 in 0 L austin jr editor life histories of careful range management practices can north american cardinals grosgrosbeaksbeaks bunbuntingstings tow significantly reduce the impacts of grazing on hees finfinchescinchesches sparrows and allies US national museum bulletin 237 part 3 washington DC plant and animal communities in riparian or BEEDY E C AND S L GRANHOLM 1985 discovering meadow ecosystems leege et al 1981 gillen sierra birds western slope yosemite natural history et al 1985 taylor 1986 schulz and lemmlerleiningerlemmger association and sequoia natural history associa- 1990 popolizio et al 1994 bich et al 1995 tion although range condition will vary with land- BENEDICT N B 1982 mountain meadows stability and change Madromadronofiobnonno 29 148 153 ownership patterns loring and workman 1987 BENEDICT N B AND J MAJOR 1982 A physiographic strict control of grazing intensity and season of classification of subalpine meadows of the sierra use will result in higher abundances of breed- nevada california Madromadronofio 29 1 12 ing birds primarily through increased shrub BICH B S J L BUTLER AND C A SCHMIDT 1995 effects volume and height taylor 1986 longtermlong term ex- of differential livestock use on key plant species and rodent populations within clusion of livestock on meadows combined selected oryzopsis hymen with oideshilariaoideslhilariaoldesoides Hilaria jamesiijamesie communities of glen canyon erosion control measures will especially bene- national recreation area southwestern naturalist fit lineolLincollincolnrss sparrows and other similar species 4028140 281 287 because of the combined vegetative and hydro- CODY M L 1981 habitat selection in birds the roles of logic effects baseline data on abundance and vegetation structure competitors and productivity biosciencebioscience3131 107 113 distribution such as those provided in this COOK C W 1966 factors affecting utilization of moun- study are essential for monitoring population tain slopes by cattle journal of range management trends resulting from disturbance or restoration 1920019 200900goo 204 of sensitive ephemeral meadow systems debenedettiDEBENE DETri S H AND D J PARSONS 1979 natural fire in subalpine meadows a case description from the sierra nevada journal of forestry 77 477 479 acknowledgments 1984 Postpostfireportfirefire succession in a sierran subalpine meadow american midland naturalist illiliiiilii111 118 125 Numenumerousiouslous people from the USU S forest ser- DUNWIDDIE P W 1977 recent tree invasion of subalpine vicevlee and national park service assisted with meadows in the wind river mountains wyoming general information on meadows within their arctic alpine research 9 393 399 FLACK J A D 1976 bird populations of aspen forests in jurisdiction marianne flett and john harris western north america ornithological monographs kindly provided unpublished data on avian 191 97 species composition of meadows surveyed in FORMAN R T T A E GALLI AND C E LECK 1976 for- 1986 for willow flycatchers empidonax trail est size and avian diversity in new jersey woodwoodlotslots with some land use implications oecologia 26 1 8 hilillii including presence or absence of s lineolLincollincolnrs FRANKLIN J E W H MOIR G W DOUGLAS AND sparrows lloyd kiff supplied photocopies of C WIBERGWIBFRG 1971 invasion of subalpine meadows by data slips for egg sets housed at the western trees in the cascade range washington and ore- foundation of vertebrateofvertebrate zoology karen klitz gon arctic alpine research 3 215 224 GILLEN R L W C kreegerKRUECERKRUEGER AND R E MILLER 1984 sketched the Lincollincolnnss sparrow in figure I1 lineol in cattle distribution on mountain rangeland in north- I1 am grateful to ned K johnson bossross lein eastern oregon journal of range management 37 martin morton robert ohmart james rising 549 553 199719971 LINCOLN S SPARROWS IN MEADOWS 115

1985 cattle use of riparian meadows in the blue PARSONS D J 1981 fire in a subalpine meadow fremon mountains of northeastern oregon journal of range tiatiactia99 16 18 management 38 205 209 PLATTS W S 19819811 effects of sheep grazing on a riparian GRINNELL J AND A H MILLER 1944 the distribution stream environment US forestfoiest service research of the budsbirds of california pacific coast avifauna 27 note INT 307 1 617 PLATTS W S AND R L NELSON 1985 streamside and GRINNELL J AND T I1 STORER 1924 animal life in the upland vegetation use by cattle rangelands 7 5 10 yosemite university of california press berkeley POPOLIZIO C A H GOETZ AND P L CHAPMAN 1994 JOHNSON N K 1975 controls of number of bird species short term response ofnpananofriparianof riparian vegetation to 4 graz- on montane islands in the great basin evolution 29 ing treatments journal of range management 47 545 567 48 53 1995 seven avifaunal censuses spanning one half RATLIFF R D 1982 A meadow site classification for the century on an island ofwhiteof white firs abies concolor in sierra nevada california USU S forest service gen- the mojave desert southwestern naturalist 40 eral technical report PSW 60 76 85 1985 meadows in the sienaslenasierra nevada of califor- KAUFFMAN J B AND W C KRUEGER 1984 livestock nia state of knowledge US forestfoiest service general impacts on riparian ecosystems and streamside man- technical report PSW 84 agement implications a review journal of range RAUZI F AND C L HANSON 1966 water intake and management 37 430 437 runoff as affected by intensity of grazing journal of KAUFFMAN J B W C KRUEGER AND M VAVRA 1983 range management 19 351 356 impacts ofeattleof cattleeattleeattiecattie on streamstreambanksbanks in northeastern ore- roathROATII L R AND W C KRUEGER 1982a cattle grazing gon journal of range management 36 683 696911 and behavior on a forested langeiangerange journal of range KRATTERKBATTER A W 1992 montane avianavlan biogeography in management 35 332 338 southernsouthernthein california and baja california journal of 1982b cattle grazing influence on a mountain biogeography 19 269 283 riparian zone journal of range management 35 KURAMOTO R T AND L C BLISS 1970 ecology ofofsubsub- 100 103 alpine meadows in the olympic mountains wash- SCHULTZ T T AND W C LEININCERLEININGCR 1990 differences ington ecological monographs 40 317 347 in riparian vegetation structure between grazed LEEGE T A D J HERMAN AND B ZAMORA 1981 effects areas and exclosures journal of range management of cattle grazing on mountain meadows in idaho 4329543 295 299 journal of range management 34 324 328 slegelSIEGELSILCEL S AND N J CASTELLAN JR 1988 nonparametricNonnonparametncparametric LENTZ J E 1993 breeding birds of four isolated moun- statistics for the behavioral sciences and2nd edition tains in southern california western birds 24 mcgraw hill new york 201 234 SPEIRS J M AND D H SPEIRS 1968 melospiza hncolmilincollincolniilincolnianiinil LORING M W AND J P WORKMAN 1987 the relation- lincolniilincollincolnianii audubon lincoln s sparrow pages ship between land ownership and range condition in 1434 1467 in 0 L austin jr editor life histories rich county utah journal of range management of noinorthth american caidmalscaidcaldealdcardinalsmalsmais giosglosgrosgrosbeaksgiosbeaksbeaks buntingsbuntings 4029040 290 293 towtowheeshees finchesfincinchesches sparrows and allies USU S national LUSBY G C 1970 hydrologic and biotic effects of grazing museum bulletin 237 part 3 washington DC vs non grazing near grand junction colorado jour- TAYLOR D M 1986 effects of cattle grazing on passelpasserineme nal of range management 23 256 260 birds nesting in riparian habitat journal of range MILLER A H AND T T MCCABE 1935 racial differenti- management 3925439 254954 257 ation in passerella melospiza lincolhncolmilincolniilincolnianiinil condor 37 VALE T R 1981a ages ofofinvasiveinvasive trees in dana meadows 144 160 yosemite national park california madronoMadro fionionno 28 NATIONAL OCEANIC AND atmospheric administration 45 47 1987 climatological data annual summary california 1981b198 ib tree invasion of montane meadows in volume 91 oregon american midland naturalist 105 61 69 NATIONAL OCEANIC AND atmospheric administration WIENS J A 1989 the ecology of bird communities vol- 1988 climatological data annual summary california umes I1 & 2 cambridge university press cambridge volume 92 UK NATIONAL OCEANIC AND atmospheric administration WOOD S H 1975 holocene stratigraphy and chronologychlonchionology 1989 climatological data annual summary california of mountain meadows sienaslenasierra nevada california un- volume 93 published dissertation california institute of tech- OHMART R D 1994 the effects of human induced nology pasadena changes on the avifauna of western riparian habitats pages 273 285 in J E jehl jijr and N K johnson received 16 september 1996 editors A century of avifaunal change in western accepted 24 february 1997 north america studies in avian biology no 15 cleatcreatgleatgreat basin naturalist 572 0 1997 appp 116 123

DENSITY distribution AND HABITAT OF flammulated OWLS IN IDAHO

craig grovesl2gioves12 terry frederiekFrederickfrederickiFrederickl 133 glenngienn FrederickfrederickiFredericklfrederickl3frederick13133 eric atkinsonl4atkmson14 melonie atkinsonl4atkmson14 jay Shepherdshepherds 5 and gregg Servheen 1

ABSTRACT flomfrom 1990 to 1992 we surveyed for flammulated owls otusflammeolusotus flamflammeolusglammeolus in 3 areas in idaho salmon national foiestforestbolest SNFSNIT payette national forest and adjacent hells canyon national recreation area PNFHCNRAPNF HCNRA and nez peiceperce national forestfoiest NPNF we also collected and summarized information on all historic and modern records of flammulated owls in idaho flammulated owls were detected on 65 of 68 routes 2 16 km in length surveyed at densities ranging from 0040.040 04 to 1251.251 25 singing males40males 40 ha owls weiewelewere detected on survey routes as eailyeallyearly as 10 may and as late as 23 july mean percent canopy cover estimated at owl locations on the PNFHCNRAPNF HCNRA and NPNF study sites ranged hornhormhomm 52 to 64 while shrub covelcover ranged from 16 to 21 and ground cover was 39 to 49 our surveys and sum malymary of distributional records indicate that flammulated owls occur throughout the montane forests of idaho in old or matinmature e stands of open ponderosa pine pinus ponderosa douglas fir pseudotsuga menziemenziesiamenziesiimenziesnsnsilsiisll and stands co dominated by those 2 species fire suppression and timber harvest activity in ponderosa pine forests represent 2 mainmaln threats to the species future security in idaho more searchresearchle on the effects of various silvicultuialsilvicultural treatments on flammulated owl populations is wanwarrantedanted

key words flammulated owl otus flarnmeolusflammeolusflam meolus population densities distribution broadcast tape recordings habitat idaho ponderosa pine pinus ponderosaponde iosarosa douglas fir pseudotsuga menziemenziesiimenziesiamenziesnsnsiislisll

the flammulated owl otusflammeolusotus flamflammeolusglammeolus is a river mountains on the salmon national for- small insectivorous owl that nests in montane est in east central idaho atkinson and atkin- forests of western north america and appar- son 1990 2 several small mountain ranges ently migrates to the neotropicsNeotropics mccallum located primarily on the payette national for- 1994d1994a on its breeding grounds it is primarily est in west central idaho moore and freder- associated with forests containing yellow pine ick 1991 and 3 salmon river and clear- either ponderosa pine pinus ponderosa or water mountains on the nez perce national jeffleyjeffreyjeferey pine P jeffrejeffreyijeffreyjyi reynolds and linkhart forest in north central idaho shepherd and 1992 servheen 1992 herein we report the distri- knowledge of the biology of the flammu- butbutionsions densities and habitats used by flam- lated owl comes principally from studies con- mulated owls in these areas and summarize the ducted in colorado ege g reynolds and linkhart statewide distribution based on a compila- 1987a 1987b new mexico mccallum and tion of historic and modern records of flammulated owls idaho gehlback 1988 mccallum et al 1995 oregon in goggans 1986 and british columbia howie STUDY AREA METHODS and ritcey 1987 although the flammulated AND owl has been documented as a nesting species we surveyed for flammulated owls on por- inm idaho burleigh 1972 hayward 1986 hay- tions of the salmon national forest SNF ward garton and carton 1988 little published infor- leahilemhi county 4515n45015n IMW in 1990 mation exists on its breeding status distribu- the payette national forest 4445n 116 tion habitats or population density stephens 30w and adjacent hells canyon national rec- and sturts 1991 reation area 4535n4535x 11625w in adams we conducted surveys for flammulated owls washington and idaho counties PNFHCNRAPNF HCNRA from 1990 to 1992 at 3 study sites 1 salmon in 1991 and the nez perce national forest

dailoI1 dalioidaho C onsoivationconservation data centelcenter idaho department of fishrish and cainecalnegainecamegame box 25 boisedoisedolse ID 83707 2peseutK sent address the1 lielleile nature conservancy 2060 blobroadwaxBiobroadwayadway suite 230 boulderBouldeidel CO 80302 licscnt3pesent addiesaddresaddressaddi is alionaallonaarizona game and fish department 555 N greasewood dovedrive tucsonTULSOII AZ 8574585743 4preentaddresspicsciit addictsaddless hawk mountain Sandsanosinctuaiysandthuyinnythuy association rr2rrb box 191 kempton PA 19529 5prescntpiepit seitsent addlessaddress box 1153 orofino 0835441 83544

116 199719971 ECOLOGY OF flammulated OWLS IN IDAHO 117

NPNF idaho county 4540n 1155rw11550w1155 rW A in 1992 fig 1 survey routes 2 16 km in length were placed A in areas containing large stands of mature ponderosa pine and douglas fir pseudotsuga menziemenziesiimenziesiasii we broadcasted taped recordings AA 49 or vocal imitations of territorial songs for owls AA along trails or roads from dusk until 0200 h at stations established 500 m apart on the SNF and PNFHCNRAPNF HCNRA sites and 500 800 m apart on the NPNF study site distances between A A broadcast stations were based on the distance 50 0 50 km manp6monniiii that we determined singing owls could be NPN heard on our study sites howie and ritcey A 1987 estimated that flammulated owls are & usually heard within 050.5os km under most weather and habitat conditions at each broad- A A cast station we listened for 1 2 min for unso- licited male songs then alternated imin1 min A A broadcasts of the songs with i- to 2 min listen- A A s A ing intervals for a total of 10 min we esti- A A mated the azimuth and distance for each owl A heard and recorded time and weather on each survey route on the SNF and PNFHCNRAPNF HCNRA study sites 1990 91 we surveyed each route only once while on the NPNF study site A 1992 each route was surveyed 2 3 times surveys were conducted 11 may 10 july on fig 1 location of the salmon national forest SNF the NPNFNPNE 10 may 23 july on the SNF and 22 payette national forest hells canyon national recreation area PNFHCNRAPNF HCNRA and nez perce national may 11 july on the forest PNFHCNRAPNF HCNRA NPNF study sites triangles represent records for flam- broadcast stations were mapped on 124000 mulated owls in idaho located outside of these 3 study or 162000 USGS topographic maps and sites see appendix A 124000 or 115840 orthoorthophotosphotos in 1990 SNF study site only we estimated the densities of responding owls on each linear survey route 2 techniques for estimating density have the with the soundsoundscapescape formula modified from same assumptions and are mathematically howie and ritcey 1987 comparable on the PNFHCNRAPNF HCNRA and NPNF study sites area ha 100 nnr2 n 1 29851.298512985.2985 we characterized the stand level habitat at owl locations by measuring habitat characteristics where n equals the number of stops per route following methods described in noon 1981 and r equals the maximum distance at which on five 004 ha circular plots 1 plot centered owls can be heardbeard a distance assumed to be on the estimated or known owl location and 050.5 km howie and ritcey 1987 this formula the other 4 located 50 m from the center plot assumes that surveys are conducted along lin- in the 4 cardinal directions to minimize the ear transects a correct assumption for the effects of error in estimating azimuth and dis- SNF study site because we used roads and tance in the habitat analysis we used only trails for survey routes on the PNFHCNRAPNF HCNRA those owl detections estimated to be within and NPNF study sites that did not conform to 200 m of broadcast stations additionally we linear transects we measured survey areas on used only those owl locations that we did not these study sites with a planimeterplanometerplanimeter to estimate think were influenced by broadcasting tapes the density of responding owls for each survey because the owl called before we began broad- route all areas within 500 m of broadcastofbroadcast sta- casting 13 locations or the owl began calling tions were used in density calculations these immediately after the broadcast and had likely 118 GREAT BASIN naturalist volume 57 not yet moved in response to the taped call 4 the state s primary ornithological treatment locations burleigh 1972 records from a database de- tree density and diameter at breast height veloped by stephens and sturts 1991 for dahdbh were measured on each plot with the publication of their latilonglatilong bird book and point center quarter method cottam and curtis records from the idaho conservation data 1956 other measurements collected on each center idaho department of fish and game plot included elevation topographic position a comprehensive database on the status and aspect dominant tree cover and understory distribution of idaho s rare threatened and vegetation number of canopy layers ocular endangered flora and fauna appendix A pro- estimate past silvicultural treatment stand age vides a summary of these records and observa- percent canopy cover tensiometerdensiometerdensi ometer percent tions ground cover line intercept percent shrub cover line intercept and distance to nearest RESULTS snag with cavity moore and frederick 1991 owl densities and distribution stand age was defined as follows 1 immature trees not cone bearing 2 mature trees from 1990 to 1992 we conducted 85 surveys cone bearing and 3 old multiple canopy on 68 survey routes distributed over 3 national layers abundant large snags trees with dahdbh forests table 1 flammulated owls were de- 64 cm from noon 1981 data from the five tected on 44 of the 68 routes 65 on routes 004 ha plots were combined to calculate means where owls were detected mean owl densities for canopy cover ground cover shrub cover singing males40males 40 ha for the 3 study sites tree density and dahdbh ranged from 0280.28ogs to 0520.52 table 1 the earliest at the PNFHCNRAPNF HCNRA study site we also in- and latest dates that owls were detected were vestivestigatedgated macromacrohabitathabitat characteristics through 10 may 1990 and 23 july 1990 respectively an analysis of the type of habitat age of forest other owl species detected on these surveys and type of timber harvest that occurred in included great horned owl bubo virginianusvirginianus 400 m radius 503 ha circles approximately long eared owl aslaasioasiaasio otus northern saw the diameter of a flammulated owl territory whet owl aegoliusAegolius acadicus and barred owl reynolds and linkhart 1987a centered on strix eariaeatlaearldvariaearly on the SNF short eared owl the owl s estimated location we overlaid these asio flamflammeusflammeousmeus northern pygmy owl glau- 503 ha circles on 115840 aerial photos and cidium anomagnoma great gray owl strix cebunebu 124000 orthoorthophotosphotos on which forest service losaiosa barred great horned long eared and silviculturists had delineated polygons and northern saw whet owls on the PNFHCNRAPNF HCNRA assigned them to the following classes based and great horned northern saw whet and on air photo interpretation and stand exams barred owls on the NPNFNPNE moore and frederick 1991 nonforest canopy we compiled 74 additional distributional closure 10 clearcutsclearcuts seedlings not visible records of flammulated owls in idaho appen- on photo open woodlands noncommercial dix A fig 1 eighteen of these records came forests with low canopy closure poor accessi- from additional surveys conducted by payette bility and poor regeneration selective cuts national forest staff in adams county within immature poles and saplings visible on photo the PNFHCNRAPNF HCNRA study site for the 55 records trees generally 50 yr old mature trees that contained habitat information 43 78 50 100 yr old old trees 120 yr old and were from areas dominated by ponderosa pine other water unclassified lands the area of douglas fir or a combination of both species these classes within each 503 ha circular plot these records were distributed throughout was estimated with a planimeterplanometerplanimeter the montane forest portions of the state the in addition to the surveys conducted in this earliest record was dated 30 march and the study we collected information on all historic latest 17 october and modern observations or records of flam- habitat characteristics mulated owls in idaho we examined the collection records of all natural history museums in STAND LEVEL we measured stand level idaho college of idaho boise state university habitat characteristics at 12 owl locations on university of idaho idaho state university the PNFHCNRAPNF HCNRA and 5 locations on the records from american birds records from NPNF study sites table 2 forty percent of 199719971 ECOLOGY OF flammulated OWLS IN IDAHO 119

TABLE 1 flammulated owl survey results on the salmon SNF payette hells canyon NRA PNFHCNRAPNF HCNRA and nez perce NPNF national forests idaho 1990 1992

owl densitya1 number number owisowls40owls 40 haba survey routes with survey site survey dates routes owls Xy s range

SNF 9 may 23 july 1990 20 16 028 029 0040.040 04 1251.251 25 PNFHCNRAPNF HCNRA 22 may 11 july 1991 38 22 031 022 0090 09 0840.840 84 NPNF 11 may 10 july 1992 10 6 052 029 0250.25008525 0980.980 98 total 68 44

density calculated only forfoiborbbr survey routes on which owls were detected

owl locations were located on upper slopes discussion 25 on ridges 25 on mid slopes and 10 on valley bottoms elevations for the 17 loca- our surveys and compilation of distributions averaged 1561 m s 39839.8 on the tional records of flammulated owls show that PNFHCNRAPNF HCNRA study site and 1504 m s 27 this species inhabits montane forests through- on the NPNF site out idaho fig 1 in our surveys flammulated either ponderosa pine or ponderosa pine owls were not detected in only I1 area the mixed with douglas fir dominated the vegeta- south fork clearwater river of NPNFNPNE A lack tion at owl locations on the NPNF and PNF- of snags and large diameter ponderosa pine HCNRA sites table 2 cover estimates for trees due to timber harvest and firewood cut- canopy shrub and ground vegetation layers ting may have made this area less suitable to were similar in the 2 study sites tree density flammulated owls was approximately 500ha on both areas and densities of responding owls in idaho were mean dahdbh values for all trees was 32 cm for within the range of 0030.03 1091.09log owls40 ha re- the PNFHCNRAPNF HCNRA and 31 cm for the NPNFNPNE ported in northern california marcot and hill although we did not collect habitat data on 1980 during similar aural surveys although the SNF study site forest service timber crews we recorded high densities on some survey had previously inventinventoriesinventoriedoried 26 of the 67 stands routes average regional densities were less that contained owl locations in our surveys on than estimates of 070.7 males40males 40 ha recorded by the SNF atkinson and atkinson 1990 all but howie and ritcey 1987 in british columbia one of these stands were dominated by dou however caution should be exercised in com- glas fir ponderosa pine or a mix of both the paring these results because the actual areas exception was a stand of subalpine fir abies surveyed vary due to differences in wind top- lasiocarpalasiocampa I1 bearbeargrassgrass xerophyllum tenaxtanax ogographyraphy individual and seasonal variation in average dahdbh values were 28328.3 cm s 818.1 owl responses and observer bias furthermore for douglas fir and 38138.1 cm s 15015.0iso for pon- estimates of density calculated with a theoreti- derosa pine cal radius of coverage around broadcast sta- macrohabitatMACRO HABITAT we measured macrohabi tions may be biased upward because of owls tat features of flammulated owls at 9 owl loca- moving into survey areas in response to broad- tions on the PNFHCNRAPNF HCNRA study site within casted calls and owls following surveyors from the nine 503 ha macromacrohabitathabitat plots on the I1 broadcast station to another R reynolds per- PNFHCNRAPNF HCNRA mean percent cover of forest sonal communication 120 yr old was 31 xF 16 ha s 18 mean most detections of flammulated owls in cover of timber stands 50 100 yr old was 30 idaho were in mature to older stands of pon- xY 15 ha s 10 followed by natural open- derosa pine mixed with douglas fir however ings at 12 Tx 6 ha s 9 and selective several records of flammulated owls that we cuts at 12 Tx 6 ha s 5 clearcutsClearcuts im- compiled from areas outside our study sites mature forest open woodlands and other un- represented owls heard calling and nesting in classified areas were relatively low in cover I1 case from pure stands of douglas fir or 2 ha each aspen particularly in southeastern idaho where 120 GREAT BASIN naturalist volume 57

tableTABLL 2 habitat characteristics at flammulated owl locations on the payette national forest hells canyon national recreation area PNFHCNRAPNF HCNRA n 12 and the nez perce national forest NPNF n 5 idaho 1991 1992 percent cover dominant vegetation type lyasxsS I1 of plots location tree density xT dahdbh canopy shrub ground pipoanipoapipo1 hahass cmajcmsjcm s PSME PIPO other PNFHCNRAPNF HCNRA 498294 325 6412 1613 4910 25 50 25b NPNF 4941426 3127 5230 2120 3918 84 16ige pipovipoFIFO pimapimm ponderosa PSMEPSML pseudotsuga menziemenziesutesutegusii 1111isI1 ins catgorycatcategorytitorgory represetitsi epi lblnts plots dorninateddornin itedcited by populus trcinuloidestrcmuloides 01or abies grandis ribisL riiis1 ins categoryc itcoiiacoi representsi epiept cscnts plots doniiiiateddomin itceitcd by P menzilminenziesii 01or laralaritlarix occidentoccioccidentalsoccidentaleoccooccidentalisoccidentalistdentalsalis

ponderosa pine is absent in addition we fire suppression and timber harvest little is recorded several instances of owls calling from known about the effects of these management stands of grand fir abies grandis these pat- activities on flammulated owls at either the terns are consistent with other studies which stand level through changes in forest structure indicate that while open ponderosa pine forests or at the landscape level through habitat frag- represent the most common nesting habitat mentation mccallum 1994b 1994 biologists other forest types are used aspen is a frequent and land managers would greatly benefit from component of nesting habitat in colorado and more research aimed at a greater understand- nevada reynolds and linkhart 1987b mccal- ing of the habitat requirements of this forest lum 1994b and owls have nested successfully owl and the impacts that various silvicultural in selectively harvested douglas fir stands in treatments might have on these requirements british columbia howie and ritcey 1987 our habitat analyses indicated flammulated acknowledgments owls in western and north central idaho use forestfoiestbolesthorest stands with mature to old ponderosa pine owl surveys were funded primarily by chal- and douglas fir multiple canopy layers low lenge cost share grants from the salmon pay- tree densities moderate to low canopy closure ette wallowagallowa whitman and nez perce national and moderate ground cover these habitat fea- forests we particularly want to acknowledge tures are characteristic of old ponderosa pine the assistance of US forest service biologists forests and are similar to those reported for D wenger F gordon R anderson and S this species elsewhere in its bleedingbreeding range blair in organizing and funding these projects goggans 1986 mccallum and gehlbach 1988 C groves and G servheen were supported by bull et al 1990 reynolds and linkhart 1992 the nongame and endangered wildlife pro- however because habitat measurements were gram license revenues and pittman robertson taken only at occupied sites and survey routes funds from the idaho department of fish and were chosen to maximize the amount of old or game special thanks to J rohlman for his help mature ponderosa pine habitat selection can- on the PNFPNE numerous other forest service not be inferred from these data and fish and game employees assisted with marshall 1939 suggested that the flam- logistical or field support in conducting these mulated owl may be the most common raptor surveys G hayward R ryder F cassirer in pine forests of the western united states D A mccallum and R reynolds provided ouioulour surveys in idaho indicate flammulated helpful reviews of earlier drafts of this manu- owls are abundant in particular habitats but script D A mccallum kindly shared distribu- no comparative data exist for densities of other tional data on flammulated owls in idaho and forest raptorsraptores whether the populations we other western states assembled during a forest studied are increasing stable or declining is service technical assessment C trost E yen- not known nor is it known for the species sen and D johnson searched collections throughout its range mccallum 1994 of idaho museums for specimen records G there is however some concern for the stephens and P peterson provided summaries species future ponderosa pine forests in idaho of database records from the idaho depart- and elsewhere are declining due in part to ment of fish and game 199711997 ECOLOGY OF flammulated OWLS IN IDAHO 121

literature CITED MCCALLUM D A AND FE R GEHLBACH 1988 nest site preferences of flammulated owls in western new 90 653 ATKINSON E C AND M L ATKINSON 1990 distribution mexico condor 661 and status of flammulated owls otus flmmo1flamflammeolusglammeolus MCCALLUM D A F R GEHLBACH AND S W WEBB on the salmon national forest idaho department of 1995 life history and ecology of flammulated owls fish and game boise 25 appp in a marginal new mexico population wilson bul- 107 BULL E A WRIGHT AND M HENJUM 1990 nesting letin 530 537 MOORE T AND G P FREDERICK 1991 distribution habitat of flammulated owls in oregon journal of L raptor research 24 52 55 and habitat of flammulated owls otus flamflammeolusmeolus BURLEIGH T D 1972 birds of idaho caxton printers in west central idaho idaho department of fish game caldwell ID 467 appp and boise 28 appp NOON B R 1981 techniques foiroiforbol sampling avianavlan habitats COTTAM G AND J T CURTIS 1956 the use of distance avianavlan pages 42 52 D E capen editor use of mul- measures in phytosociological sampling ecology 37 in the 451 460 tivariatetitivanate statistics in studies of wildlife habitat USDA general GOGGANS R 1986 habitat use by flammulated owls in forest service technical report northeastern oregon unpublished thesis oregon RM 87 rocky mountain forest and range experi- state university corvallis 54 appp ment station fort collins CO REYNOLDS R T AND B D LINKHART 1987a fidelity to HAYWARD G 1986 activity pattern of a pair of nesting and in owls pages flammulated owls otus flammeolusflammeolus in idaho territory mate in flammulated northwest science 60 141 144 234 238 in R W nero R J clark R J knapton and R H hamre editors biology and conservation HAYWARD G AND E 0 GARTON 1988 resource partition- forest owls proceedings 3 7 ing among forest owls in the river of no return of northern symposium wilderness idaho oecologia 75 253 265 february 1987 winnipeg manitoba USDA forest general RM 142 HOWIE R R AND R RITCEY 1987 distribution habitat service technical report rocky and range selection and densities of flammulated owls in mountain forest experiment station british columbia pages 249 254 in R W nero R J fort collins CO 1987b biology clark R J knapton and R H hamre editors the nesting of flammulated owls pages nero R biology and conservation of northern forest owls in colorado 239 248 in R W J clark R and R editors biology symposium proceedings 3 7 feblfebruaryFebi uary 1987 win- J knapton H hamre nipeg manitoba USDA forest service general and conservation of northern forest owls symposium technical report RM 142 rocky mountain forest proceedings 3 7 february 1987 winnipeg manitoba general and range experiment station fort collins CO USDA forest service technical report RM 142 rocky and range MARCOT B G AND R HILL 1980 flammulated owls in mountain forest experi- northwestern california western birds 11 141 149 ment station fort collins CO 1992 owls ponderosa MARSHALL J T 1939 territorial behavior of the flammu- flammulated in pine evievlevi- pages lated screech owl condor 41 71 78 dence of preference for old growth 166 169 R MCCALLUM D A 1994a flammulated owl in A poole inm M R kaufmann W H moir and L bassett and FE gill editors birds of north america no 93 editors old growth forests in the southwest and academy of natural sciences philadelphia PA rocky mountain regions proceedings of a workshop 1994b review of technical knowledge flammu- USDA forest service general technical report and range lated owl pages 14 46 in G D hayward and J RM 213 rocky mountain forest experi- verner editors flammulated boreal and great ment station fort collins CO SHEPHERD AND G SERVHEEN 1992 gray owls in the united states a technical conser- J F flammulated sampling vation assessment USDA forest service general owl otusflammeolusotus flamflammeolusglammeolus surveys and habitat RM 253 rocky mountain on the clearwater red river and salmon river technical report forest perce range collins CO i angerranger districts nez national forest idaho and experiment station fort game 1994c conservation status of flammulated owls department of fish and boise 23 appp STEPHENS D A AND S H STURTS 1991 idaho bird dis- in the united states pages 74 79 in G D hayward tribtribution special publication 11 idaho museum of and J verner editors flammulated boreal and great ution pocatello 76 pp gray owls in the united states a technical conser- natural history ap vation assessment USDA forest service general technical report RM 253 rocky mountain forest received 26 february 1996 and range experiment station fort collins CO accepted 10 march 1997

appendix A begins on the following page 122 GREAT BASIN naturalist volume 57

APPENDIX A

summary of all flammulated owl records and observations in idaho not recorded during surveys conducted at 3 study sites SNFSNE PNFPNFHCNRAHCNRA NPNF and reported in this paper more detail on these records is provided by the idaho conservation data center IDCDC idaho department of fish and game box 25 boise ID 83707 date observer HabHabitatitata1 county source of data 22 september 1890 merriam blaine burleigh 1972 28 september 1914 rust kootenai burleigh 1972 summer 1973 trost urban bannock ISU Museum 15 22 september 1977 powers urban ada AB 32234 23 september 1977 jeppson urban bannock AB 32234 september 1978 jeppson urban bannock AB 33197 7 may 1980 humble shoshone IDCDC 5 october 1980 trost DF bannock ISU museum 1981 1985 hayward PIPP idaho hayward and garton 1988 june 1982 hayward DFPP boise hayward 1986 26 august 1982 jeppson urban bannock AB 37204 26 september 1986 jeppson urban bannock ISU museum 6 july 1987 ulmschneider DF elmore IDCDC 3 october 1987 dudley gem stephens and sturts 19911 5 july 1988 hansen kootenai AB 421319 september 1988 trost urban bannock ISU museum 17 october 1988 hansen kootenai AB 43138 july 1989 trost DF bannock IDCDC may 1990 trotter twin falls AB 44467 22 may 1990 trost DF bannock AB 44467 14 june 1991 atkinson DF fremont IDCDC 19 june 1991 patla aspen teton IDCDC may 1991 trotter twin falls AB 45475 31 may 1991 trochlell DF camas IDCDC I1 june 1991 trochlell twin falls IDCDC july 1991 trost DF bannock AB 451140 18 september 1991 urban bingham ISU museum 12 april 1992 leppert PP adams IDCDC 21 april 1992 mccammon PP boundary IDCDC 24 may 1992 gray DFPP idaho IDCDC 29 may 1992 Baerbaerlockerbaerlocherlocker idaho stephens and sturts 1991 30 may 1992 ulmschneider boise IDCDC june 1992 leppert DFPP adams IDCDC 16 june 1992 leppert adams IDCDC 18 june 1992 mccammon PIPP boundary IDCDC 18 june 1992 walker DFPP adams IDCDC 18 june 1992 leppert DFPP adams IDCDC 19 june 1992 leppert DFPP adams IDCDC 20 june 1992 leppert DFPP adams IDCDC 21 june 1992 leppert DFPP adams IDCDC 22 june 1992 riley PP bonner IDCDC 23 june 1992 atkinson aspen bonneville IDCDC 24 june 1992 richards PP adams IDCDC 7 july 1992 leppert DFPP washington IDCDC 14 july 1992 skinner DFPP washington IDCDC 17 july 1992 belt grand fir idaho IDCDC 17 july 1992 leppert idaho IDCDC 27 july 1992 naderman bonneville IDCDC 17 august 1992 feldham DF bear lake stephens and sturts 1991 5 october 1992 svingen benewahbenejah ab47122AB 47122 30 march 1993 leppert DFPP adams IDCDC 23 april 1993 wessman DFPP boise IDCDC 5 may 1993 trost DF bannock AB 46453 26 may 1993 evans adams IDCDC 16 june 1993 oneill PIPP adams IDCDC 17 june 1993 johnston DF idaho IDCDC 23 june 1993 holliday DFPP adams IDCDC 24 june 1993 holliday DFPP adams IDCDC 24 june 1993 holliday DFPP adams IDCDC 199719971 ECOLOGY OF flammulated OWLS IN IDAHO 123

APPENDIX A

continued date observer Habhabitatsitata county source of data 14 july 1993 johnston PP idaho IDCDC 14 july 1993 cassirer nez perce IDCDC 15 july 1993 robinson aspen bear lake IDCDC 3 may 1994 garwood DF camas IDCDC 24 may 1994 dhaenens PP idaho IDCDC 26 may 1994 garwood dfaspenDF aspen blameblainebiame IDCDC june 1994 stotts DFPP idaho clearwater NF 26 june 1994 holliday PP adams HCDCIDCDC 28 june 1994 leppert DFPP adams IDCDC 28 june 1994 holliday PP adams HCDCIDCDC 28 june 1994 belt grand fir idaho IDCDC october 1994 jefferson ISU museum 17 may 1995 wondolleck DFPP shoshone HCDCIDCDC 8 june 1995 faike blameblainebiame HCDCIDCDC 27 june 1995 mccammon PP valley IDCDC douglas fir DF ponderosaponderosiderosa pine PP douglas fir and ponderosa pine DFPP bisubasuISU idaho museum of natural history idaho state university pocatello cabeabB american budsbirds dStephensstephens and sturts 1991 a latilatilonglong database developed by D A stephens and maintained by the idaho department of fish and game great basin naturalist 572 0 1997 appp 124 130 DEN AND relocation SITE characteristics AND HOME RANGES OF peromyscus TRUEI IN THE WHITE MOUNTAINS OF california

linnea S haill and michaelmiehaelmichaei L MornsonMorrisonmorrison2morrisonnmornson22

ABSIKACIABSTRACT we used ladioradioradlo telemetry to describe nighttime movements and daytime den sites ofofpmyonpinyon mice bereerperper- omyscus trueltruettruer in the white mountains of california 1991 1993 characteristics of nighttime relocations and den sites loltoifoiemermeer mice eonconconcurredcuned with previously reported habitat use information for the species and supported the claim that pinyon mleemice use multiple daytime sites however males and females were associated with different habitat characteristics at den sites indicating differential micromicrohabitathabitat selection by the sexes perhaps related to reproductive constraints on females pinyon mleemice also exhibited high variability in den site habitat use in the summer but low variability in the fall and winter the dens of male micemlee were fartherharthelbarthel apart than those of females and home range areas averaged 292.92gg 9 ha s 42744.2727 ha for 8 males and 080 8 haba s 00760.7676 ha for 7 females overall xT 171 7 4- 2972 97 ha these areas weiwelwere e larger than those reportedi epol ted for other species of peromyscus the combined effects of drought and reduced food availability may have contributed to the larger areas used

kenkeifkey words den sites habitat use peromyscuspeipel omyscus truel pinyon mouse radio telemetry white mountains

several studies have described home range descriptions of habitat use than would live or sizes of peromyscus species especially as they snap trapping stations because the latter can are affected by food availability population bait animals into them and thus not necessarily densities iteimteiinterspecificspecific competition and habitat represent the habitat that is typically used use taitt 1981 wolff 1985 douglass 1989 cranford 1977 douglass 1989 researchers commonly used live trapping data because of these advantages we used tele- to determine home ranges and core areas ege g metry to describe den sites and home range douglas 1969 meserve 1977 merritt and sizes of pinyon mleemice P trueltruettruer in the white merritt 1978 ofarrellOTarrell 1978 ribbleribbiekibble and sam- mountains of california pinyon mleemice are com- son 1987 but recently have increased their mon inm the western USU S hofemeisterhoffmeisterHoffineisterlster 1981 but use of radio telemetry for this purpose madi- they have not figured prominently in habitat son 1977 mineau and madison 1977 douglass use studies relative to other peromyscus species 1989 many studies of home ranges and habi- but see douglas 1969 scheibe 1984a ribble tat use have been conducted on P maniculatusmamculatus and samson 1987 scheibe and ofarrell 1995 and P leuleucopuscopus wolff 19892851989 285 but only a few therefore we initiated this study to discern of these have detailed den or nest site charac- habitat use by P truet in the white mountains teristicsteristics or telemetry relocation characteristics madison 1977 kleinklemkiem and layne 1978 stah STUDY AREA 1980 wolff and hurlbutt 1982 wolff and durr 1986 douglass 1989 frank and layne 1992 the white mountains are located east of most information on mouse dens comes from bishop inyo county california and rise from older studies in which researchers located and 1515 to 4245 m elevation tst7s r35e sec 30 excavated sites they came upon by chance ege g 32 the dominant vegetation between 2090 and mccabe and blanchard 1950211950 21 douglas 1969 2725 m elevation is pinyon jumperjuniper binmpinmpmus mono telemetry allows the location of dens without phylla juniperuslumJumpetusperus osteosperma woodland this disturbance and the determination of the fre- woodland is xericxene with sparse vegetative cover quency with which mice return to den sites in and warm dry summers snow cover and rain-rain addition telemetry may permit more accurate fall usually last from november through may

dcpartnentidcpaitnientI1 of biological sciences caliloimacalifinniaCaligail loimafolmafinnia state university sacramentoSacia mento CA 95819 nviionmental2erivironinental scienceSLILIILL policy and management university of california berBeiberkeleykeleskeley CA 94720 plesentpresent address department of biological sciences cali-ca forniana state univeisityunivelsityuniversity saciaiiientosacranientoSacranientolento CA 95819

124 199719971 DEN SITES AND MOVEMENTS OF PINYON MICE 125 with afternoon thunderstorms in late summer fully boltedmolted weighing 14114.1 18018.0 g adults were unpublished data the only tree species in the born the previous year and were fully boltedmolted woodland are pinyon and juniper dominant weighing lsiisi18118.1 g shrubs are big sagebrush artemisia tfidentatatridentatdtridentated we anaesthetizeranaesthetized each mouse with Metometofanefane purshia bitterbitterbrushbrush glandulosaglandulose and P triden and removed a 202.0 X 10 cm patch of dorsal tafatata rubber rabbitbrushrabbitbrush chrysothamnus nau fur from between the shoulders down to just selsusseosusseosus green rabbitbrushrabbitbrusbrabbitbrush C viscidiflorusviscidiflorus above the skin we then adhered the transmit- mormon tea ephedra virviridisidis cactus opuntia ter to the back with cyanoacrylate each mouse and echinocereus sppapp and squaw apple pera was kept in a protected box for 10 15 min and phyllusphyllum sp dedecker 1984 more detailed I1 released at its initial point of capture after its descriptions of the study area are provided in recovery from anaesthesia hall 1992 we obtained the ist relocation fix on each METHODS radio tagged animal during the daytime about 6 h after the radio was attached we used a radio telemetry portable receiver telonics inc mesa AZ and to catch mice for attachment of radio trans- communications specialists inc orange CA mittersmitters we set sherman live traps 76767.6 X 898.9sg and 2 and 3 element yagi antennas after loca- X 22922.9 cm on two 4 ha grids and four 18isha ha ting a signal we walked to the fix or den site grids at about 2500 in elevation in the pinyon and recorded the azimuth and distance in to a juniper vegetation the large grids pinyon and trap location on the grid and then determined cedar 81 trapsgridtraps grid with 25 m spacing were coordinates for the fix location we obtained established in 1988 as part of a longtermlong term proj- 1 3 fixes every night and I1 den site was re- ect on mice the small grids 100 trapsgridtraps grid with corded every day for each individual as long as 15ism m spacing were established in 1991 as con- the transmitter remained attached forty three trol grids used in an experimental study mor- percent of the telemetry relocations were col- rison 1988 hall 1992 these grids were used lected when there was no or very little moon- for a total of 870 trap nights from 1991 to 1993 light ie around the time of a new moon and specifically to provide animals for telemetry only 16 were collected when the moon was additional trapping was conducted on the sites 60 100 full and overhead at night we homed to determine population abundances and habi- in white and garrott 199042 on each mouse tat use morrison et al 1991 hall 1992 by following the transmitted signal until we were we checked traps between midnight and within 10 in of the animal this was accom- first light rather than at dawn to minimize the plished by keeping our lights off or at low power stress of affixing transmitters on nocturnal mice and minimizing all noise as we approached with to identify species surety we examined radio tagged individuals specimens at the museum of vertebrate zool- we radio tagged a total of 30 pinypinyonon mice ogy university of california berkeley and from july 1991 through july 1993 of these 21 compared them with specimens we collected were tagged between july and december 1991 from 1988 to 1991 morton et al 1995 in addi- 2 in january 1992 and 7 between may and tion there were only 2 species of peromyscus july 1993 of the males and inhabiting our study area P truel and P man sixteen mice were iculatus which were not difficult to distin- 14 females 20 were adults and 10 subadultssubadults guish from each other thus we are confident characteristics of nighttime relocations and in our identifications of pinyon mice daytime den relocations were described for 26 we radio tagged subadult and adult mice and 25 animals respectively to assess habitat for which transmitters were 10 of their characteristics we recorded 11 substrate types body masses transmitters with acrylic potting in on or under which each fix occurred table averaged 202.0 g smiSM 1 mouse style AVM instru- 1 we also recorded the plant species nearest ment co livermore CA ages of animals the fix site ie the primary plant the plant were distinguished based on our previous work species near the primary plant but fartherearther with the populations morrison et al 1991 mor- from the fix site ie the secondary plant and rison and hall unpublished data subadultssubadults the size small 2 in medium gi91212.1 494.9 in or were born the current year and were nearly large 5 in of the nearest plant 126 GREAT BASIN naturalist volume 57

TABLE 1 substrates used for den sites by pinyon mice P 0570.57ost df 3 4 and so we pooled all grids andor associated with relocation sites of micemlee in the together to make sample sizes larger for the white mountains of california 1991 1993 other analyses we then conducted t tests zar substrate 1984126 of home range sizes between sexes dead tree base andor exposed roots and ANOVAs of home ranges among seasons live treeti ee base summer july august fall september octo- live treeti ee lipuiptipup in ber winter december january rock outcrop slab shelfsheifshelo all downed log for statistical analyses we used SPSS snag PC norusis 1992 and considered p5Ppa 0050.05oos tree trunk to be significant dead branch or mistletoe ball in live tree shrub base hole RESULTS stump nighttime relocation characteristics most nighttime relocation sites were associated with singleleafpinyonsingleleafsinglesingieleaf pinyon trees 42 utah analyses jumperjuniper trees 31 bitterbitterbrushbrush 11 and rock outcrops 8 across all grids male mice used DEN AND SITES we used log relocation shrub bases and downed logs less than expected linear analyses sokal and rohlf 1981747 to whereas female test for patterns of association among combi- mice used these substrates nations of plant and substrate variables recorded more than expected loglinearloglinear analysis CG at nighttime and daytime relocation sites for 18218.2 FP 00020.002 df 8 adult mice used these analyses we evaluated each combination shrub bases less than expected subadult mice of dependent and independent categorical vari- more than expected G 19619.6 P ooi0010.01 df ables eg sex of mouse by substrate age of 8 subadult mice were associated with jumperjuniper mouse by nearest plant season by plant size trees more often than expected whereas adult A loglinearloglinear analysis tests for independence of mice again showed the opposite pattern G igi191 categories by calculating odds ratios from resid- 191p0004df619119.1 P 00040.004 df 6 uals of observed and expected values and then den site characteristics determining the goodness of fit of the resulting ratio model sokal and rohlf 1981748 the average distance moved between con- the analysis is useful for uncovering relation- secutive den sites was 47047.0 in s 1060logo10.60 in ships among variables in a multmultiwayiway crosstab n 17 mice with 2 den sites males moved alationulaulationtion similar to multiple comparison proce- an average of 82082.0 in s 166516.65 in n 6 and dures for multmultiwayiway analyses of variance females moved 31031.0 inm s 137613.76 in n 8 HOME RANGES we used program mcpaalmcphal the difference between the sexes was significant vers 1221.22 M stuwe and C E blohowiak t 2382.38 P 0040.04 df 12 males moved a conservation and research center national maximum of 160 in and females a maximum of zoological park smithsonian institution wash- 126 in between consecutive dens the mini- ington DC to determine the home range sizes mum movement was 0 in ie when mice of mice for which we collected 8 total fixes stayed in the same den 2 nights in a row although our relocation data were few per we recorded data for 72 den sites 60 in mouse we used harmonic mean HM analyses 1991 2 in 1992 and 10 in 1993 animals with dixon and chapman 1980 to calculate home 2 den sites averaged 303.0 18181.8 different den range sizes for each mouse individually this locations during the time we followed them method includes less unused space in the home most dens were located in or under rock ledges range area than the minimum convex polygon outcrops slabs or shelves 28 and in live method hayne 1949 and it has greater utility trees 25 they were secondarily found in because it approximates the size of the activity downed logs dead branches of otherwise live centers of animals ie areas with highest trees and at shrub bases 8 each the near- activity intensities est plants to these dens were most commonly HM values of the mice did not differ signif- pinyonpinyon pines 50 and junipersjunijuniperuspers 32 icantly among the 6 grids iway1 way analysis of male mice used live tree bases more than variance ANOVA zar 1984163 F oti0710.71 expected and dead branches in live trees less 199711997 DEN SITES AND MOVEMENTS OF PINYON MICE 127

TABLE 2 vegetative characteristics associated signifi- TABLE 3 vegetative characteristics associated more or cantly more or less often than expected with den sites less often than expected with den sites used by pinyon used by pinyon mice in the white mountains california mice in summer winter and fall 1991 1993 1991 1993 summer fall winter sex variable uselusea variable useusea use use MALE live trees asbnsbns live trees shrub bases ns dead branches in live trees holes ns pinyon trees snags ns bitterbrushbitterbrusbBitterbrush shrubs pinyon trees ns small trees bitterbrushBitterbrush shrubs ns large trees small trees ns medium trees ns FEMALE use live trees use of variable by mice was significantly greater than expected use was significantly less than expected loglinearloglinear analysis P 005oos0 05 dead branches in live trees busnonsignificantbusns nonsignificant result inin loglinearloglinear analysis P 0050oos05 pinyon trees bitterbrushBitterbrush shrubs small trees large trees summer the opposite pattern occurred in fall useailseaulse use of variable by mice was significantly greater than expected G 17417.4 df 4 P 0.002 3 use was significantly less than expected loglinearloglinear analysis P 005 0002 table home ranges we analyzed a total of 134 fixes among 30 than expected whereas females showed the radio tagged mice ranging from 4 to 31 fixes opposite pattern loglinearloglinear analysis G 25.7257 257 per animal xT 10 585.8 per animal fixes P 00040.004 df 10 table 2 male dens were were collected about 10 h apart s 212.1 h associated commonly more with pinyon trees over an average of 5 d s 3 1 d and less commonly with bitterbrushbitterbrush shrubs of5ds31d than for animals with 8 total fixes n 15 7 expected whereas females again showed the females 8 males home range sizes did not opposite pattern G 10.6log P 0.03003 4 106 003 df differ among seasons for males F 1821.82 P finally female dens associated were with small 026 df 2 5 or females F 4774.77 FP 0090.09 trees more often and large trees less often than df 2 4 harmonic mean areas averaged 1.717 expected 17 males showed the opposite pattern ha s 2972.97 ha for all mice combined home of habitat use G lis11811.8 P 00030.003 df 2 range sizes of male mice did not differ from 2 table those of female mice in any season t tests FP also there were differences among age values 0080.08 combined across all seasons classes adult used mice dead branches in live male HM areas averaged 292.9 4274.27 ha and trees less than expected but subadult mice female HM areas averaged 080.8 0760.76 ha sea- used them more than expected G 235 C 23523.5 P sonal home range sizes ranged from 0010.1oi 1 ha fall 0.009 df 10 also 0009 adult mice used medium males and winter females to 555.5ss ha summer sized trees less often whereas subadult mice males table 4 used them more often G 71ti7.1 P 0030.03 df 2 discussion among seasons summer fall winter live tree and shrub bases and holes were used more characteristics of the habitat used by pinyon often and snags were used less often during mice at night and during daytime denningderming con- summer in fall live tree and shrub bases and cur with previously published data regarding holes were used in the opposite pattern in habitat use for this species hoffmeister 1951 winter snags were used in the opposite pattern 34 1981 douglas 1969461 ribble and samson G 34434.4 df 20 P 0020.02 table 3 also in 1987 we also substantiated the claim that pin- summer den sites were associated with pinionspinyons yon mice use multiple daytime sites douglas more than expected and with bitterbitterbrushbrush less 1969464 a characteristic shared by several than expected in fall the opposite pattern was other species of ofperomyscusperomyscus stickel 1968388 exhibited G 17717.7 df 6 P 00070.007 finally however male and female pinyon mice exhib- small trees were used less than expected and ited differential habitat use at daytime sites medium sized trees more than expected in males used tall pinyon trees and the bases of 128 GREAT BASIN naturalist volume 57

TABLE 4 seasonal differences in home range sizes of reduced need to move widely in search of male and female pinyonpryonpmyon mice in the white mountains mates changes in food availability for both california fromblom 1991 to 1993 sexes andor changes in environmental tem- harmonic mean values peraperaturestures among other factors eg scheibe sex season ahaxhaT ha s n 1984a our average home range area estimate for mallMALIMALE summer 55 4954 95 4 all P truel combined was 171.7 ha which was fall 01 004 2 much larger than telemetry estimates for P winter 05 059 2 leucopusleucopus by madison 1977 010.1oiol ha mineau overall 29 427 8 and madison 1977 11liii1.1 ha and wolff 1985 FEMALEFEMALL 0060.06 ha and for P maniculatus by wolff 1985 summer 16 067 2 0050.05oos ha previous estimates for pinyon mice fall 07 053 3 based on live trapping were also smaller rang- winter 01 006 2 ing from 0430.43 douglas 1969439 to 0480.48 ha overall 08 076 7 scheibe 1984b for adult male pinyon mice vs our 292.9 ha and from 0280.28 scheibe 1984b to 0370.37 ha douglas 1969 for adult females vs 0.808 ha live trees whereas females used dead branches our 08 we think the large home and mistletoe balls in small but otherwise range estimates live trees and also bitterbitterbrushbrush shrubs scheibe for pinyon mice in the white mountains in this study could have due to 3 and ofarrellOFarrellrelireIl 1995 likewise found differences been interacting factors have in habitat use between male and female pinyon first drought may influenced how mice in california and suggested that this was pinyon mice utilized space A recent california related to the constraints of reproduction on drought lasted from 1986 to the winter of females versus the relative lack of constraints 1992 93 and pinyon mice were at low abun- on males on the other hand scheibe 1984b dances and experiencing low and abbreviated found no significant differences in the home productivity during this period morrison et al range habitat characteristics of male and female 1991 morton et al 1995 morrison and hall pinyon mice in california however because submitted manuscript second the drought he considered only the species of plants pres- was combined with poor pinyon seed crops in ent it was not surprising that both sexes were the white mountains which probably further associated with the same vegetation contributed to reducing the densities of mice we also observed seasonal variation in den morrison et al 1991 morrison and hall sub- site use with greater variability in substrate mitted manuscript with decreased food and use in the summer than in the fall and winter water availability pinyon mice may have had scheibe and ofarrellOFaroearrellrelireIl 1995 similarly found to forage widely leading to the large home that female pinyon mice were consistently ranges we observed associated with particular habitat characteris- finally although some authors have sug- tics during reproductive months males how- gested that a large number of live trapping re- ever were associated with a wide breadth of captures in a trap session adequately represent habitat characteristics especially in the spring micromicrohabitathabitat and space use by small mammals and summer but did show increased habitat wolff 1985 desy et al 1989 others have sug- specificity in the fall and winter additionally gested that radio telemetry provides a more the den sites of male pinyon mice were located complete and accurate assessment of these farther apart than the dens of females males parameters cranford 1977 bergstrom 1988 and females also tended toward having differ- douglass 1989 in our study mice used an ent home range sizes similar to the findings of average of 171.7 ha over about 5 d and many mice scheibe 1984b As with habitat use these used areas off the 181.8isls and 4 ha grids hall sexual differences may result from the relative 1992 hall and morrison submitted manuscript selectivity of male and female mice based on this suggests that if we had relied only on their needs for finding mates and for tending trapping data to estimate the home range sizes nests respectively scheibe and ofarrellOFaroearrellrelireIl 1995 of mice we would have underestimated the increased habitat specificity in the fall and areas used the large home ranges were prob- winter as we observed could be tied to males ably not an artifact of the radio transmitters 199719971 DEN SITES AND MOVEMENTS OF PINYON MICE 129

studies on birds have demonstrated that trans- hoffmeister D F 1951 A taxonomic and evolutionary mitters tend to cause reduced home range study of the pinon mouse peromyscus truet illinois biological sizes rather than larger home range sizes eg monograph volume 21 1981 peromyscus trueithuei mammalian species 161 all hooge 1991 if there are any effects at the 1 5 choice of whether or not to use telemetry there- HOOGEHOOCE P N 1991 the effects of radio weight and harnesses fore has important implications for the deter- on time budgets and movements of acorn wood- minationmi of habitat use and may also influence peckers journal of field ornithology 62 230 238 calculations of survival immigration trappa KLEIN H G AND J N LAYNE 1978 nesting behavior in four species of mleemice journal of mammalogy 59 ability micemlee bility eg krebs and boonstra 1984 and dis- 103 108 persal of small mammals in grid based studies KREBS C J AND R BOONSTRA 1984 trappabihtytrappabilityTrappability estimates for mark recapturee data canadian journal of zool- acknowledgments ogy 62 2440 2444 MADISON D M 1977 movements and habitat use among we thank the staff of the white mountain interacting peromyscus leuleucopuscopus as revealed by radio telemetry canadian field naturalist 91 273 281 station bishop research california espe- MCCABE T T AND B D BLANCHARD 1950 three species cially D trydahl and E phillips and several of peromyscus rood associates santa barbara CA field assistants especially PE aigner L baker 136 appp and L nordstrom for their logistical and tech- MERRITT J FE AND J M MERRITT 1978 seasonal home nical support we also thank 2 anonymous re- ranges and activity of small mammals of a colorado subalpine forest acta theriologicathenologica 23 195 202 viewers for their helpful comments on earlier MESERVE P L 1977 three dimensional home ranges of drafts of this manuscript this work was sup- erlcriericncetidcricetidcetid rodents journal of mammalogy 58 549 558 ported by funding from J verner at the pacific MINEAU P AND D MADISON 1977 radio tracking of per- southwest forest and range experiment sta- omyscus leucopusleucopus canadian journal of zoology 55 tion fresno california and white mountain 465 468 MORRISON M L 1988 the design and importance of long- research station fellowships to LSH term ecological studies analysis of vertebrates in the inyo white mountains california USDA forest ser- literature CITED vleevice general technical report RM 166 MORRISON M L M L MORTON L S HALL J L HARNER BERGSTROM B J 1988 home ranges of three species of AND J J KEANE 1991 population biology of small chipmunks tamias as assessed by radiotelemetryradiotelemetry and mammals in the inyo white mountains california grid trapping journal of mammalogy 69 190 193 pages 246 255 in C A hall jr V doyle jones and CRANFORD J A 1977 home range and habitat utilization B widawski editors natural history of eastern cali- by neotomafuscipesneotoma fufuscipesscipes as determined by radioteleme fornia and high altitude research white mountain try journal of mammalogy 58 165 172 research station symposium volume 3 university DEDECKER M 1984 flora of the northern mojave desert of california press los angeles california california native plant society berkeley MORTON M L M L MORRISON L S HALL AND M E 164 appp PEREYRA 1995 life history paiapalaparametersmeters in mice ex- DESY E A G 0 BATZLI AND L JIKE 1989 comparison of posed to a prolonged drought southwestern natu- vole movements assessed by live trapping and radio ralist 40 18 28 tracking journal of mammalogy 70 652 656 NORUSIS M J 1992 SPSSPC base system user s guide DIXON K R AND J A CHAPMAN 1980 harmonic mean version 505.05 0 SPSS inc chicago IL 910 appp measure of animal activity areas ecology 61 OTARRELLOFARRELL M J 1978 home langeiangerange dynamics of rodents 1040 1044 in a sagebrush community journal of mammalogy DOUGLAS C L 1969 comparative ecology of pinyon 5965759 657 668 micemlee and deer micemlee in mesa verde national park RIBBLE D 0 AND F B SAMSON 1987 microhabitatmicrobabitatMicrohabitat asso- colorado volume 18 university of kansas publica- ciationsciati ons of small mammals in southeastern colorado tions lawrence with special emphasis on peromyscus rodentia DOUGLASS R J 1989 the use of radio telemetry to eval- southwestern naturalist 32 291 303 uate micromicrohabitathabitat selection by deer mice journal of SCHEIBE J S 1984a the effects of weatherofweather sex and season mammalogy 70 648 652 on the nocturnal activity of peromyscus truet roden- FRANK P A AND J N LAYNE 1992 nests and daytime tia southwestern naturalist 29 1 5 refugia of cotton micemlee peromyscus gossypinusgossypinus and 1984b sexual differences in the home ranges of golden mice ochrotomysOchrotomys nutnuttnuttalltnuttallinuttalliaalltailtalittalli in south central peromyscus trueithuei and dipodomys panamintpanamintinusinus florida american midland naturalist 127 21 30 rodentia southwestern naturalist 29 7 13 HALL L S 1992 experimental manipulation of food re- SCHEIBE J S AND M J OFARRELLOTARRELL 1995 habitat dynam- sources of micemlee in a pinyonpryonpmyon juniperjumper woodland un- icslesies in peromyscus truet eclectic females density published master s thesis university of california dependence or leproductivereproductive constraints journal of berkeley ilg119119ppappp mammalogy 76 368 375 HAYNE D W 1949 calculation of size of home range SOKAL R R AND F J ROHLEROHLF 1981 biometry and2nd edi- journal of mammalogy 30 1 18 tion W H freeman and co new york 859 appp 130 GREAT BASIN naturalist volume 57

stallSTAIISTAH C D 1980 vertical nesting distribution of two 1989 social behavior pages 271 291 in CG L kirk- species of peromyscus under experimental conditions land jr and J N layne editorsediedltoistols advances in the journal of mammalogy 61 141 143 study of peromyscus rodentia texas tech university STICKELSTICKELLL Ff19681968 home langeiangerange and travels pages 37311373 411 press lubbock in J A king editor biology of peromyscus roden- WOLFF J 0 AND D S durbDURRDUKR 1986 winter nesting behav- tia special publications of the american society of lorior of peromyscus leuleucopuscopus and peromyscus manicumanibumamou mammalogists 2 1 593 latus journal of mammalogy 67 409 412 taitltantTAITI M J 1981 the effect of extra food on small rodent WOLFF J 00.OANDBAND B hurlbut71982HuRLBUTHURLBUTT719821982 day refuges of pero- populations I1 deermiceDeermice peromyscus maniculatusmamculatus myscus leuleucopuscopus and peromyscus maniculatusmamculatus jour- journal of animal ecology 50 111 124 nal of mammalogy 63 666 668 WHITE G C AND R A GARROTT 1990 analysis ofwildlifeofwildlife ZAR J H 1984 statisticalbiostatisticalBio analysis and2nd edition pren radioradlo tracking data academic press san diego CA tice hall inc englewood cliffs NJ 718 appp WOLFF J 0 1985 the effects of density food and interspecific interference on home range size in per- received 19 july 1996 omyscus leuleucopuscopus and peromyscus maniculatusmamculatus accepted 28 january 1997 canadian journal of zoology 63 2657 2662 great basin naturalist 572 0 1997 appp 131 141

LATE FALL AND EARLY SPRING BIRD observations FOR mulettmulegtMULEGE BAJA california SUR MEXICO

robert C WhitWhitmorewhitmorelwhitmore1morelmorei1 and R craig Whitmore 2

ABSTRACT observational data from spring and fall 1996 and spring 1997 for the region near eulegemuleg6mulege baja california sur mexico are summarized in tabular form in addition new or noteworthy data for 17 species are annotated to provide clarification of previously published records A uniquely plumaged bird too far south for a female american robin turdus migratormigratoriusmigratoriesius in basic plumage and too far north for the endemic san lucas robin Turdturdusits migratormigratoriusmigratoriesius conconfilsconfconfimsconfinisimsfinisainis and intermediate in coloration between the two was recorded range expansions are documented for several species including european starling sturnus vulgarisduldutvulgans anna s hummingbird calyptecalypter anna western meadowlark sturnellaStumeliamellameila negneglectsneglectalecta white faced ibis plegadis chichi and white fronted goose anser albifronsalbifrons least brebesgrebes tachybaptus dominicus a species of concern which is apparently declining in numbers and belding s yellowthroat geothlypis beldingibeldingi a species endemic to baja california sur were observed in the freshwater marsh during all 3 study periods

key words baja california sur mexico eulegemulegemulegg bird records range expansion desert oasis

the avifauna of baja california hereafter hotel serinidadSerinidad approximately 2 km east of the BC the second longest and most geographi- town supports small aircraft while the nearest cally isolated peninsula in the world grismer commercial air service is to loreto approxi- and mcguire 1993 is one of the most inter- mately 132 linkmkin south its distance from both the esting yet poorly studied in north america northern and southern population centers makes wilbur 1987 howell and webb 1995 late eulegemuleg6mulege accessible only to hardy travelers 19th and early 20th century natural history the area features both a brackish water tidal work in BC has been summarized by E D estuary rio eulegemulegemuleg6 and a small year round nelson 1921 and J grinnell 1928 much of spring fed freshwater lagoon the former is the recent ornithological effort has been con- small 3 km long runs primarily west to east centratedcentrated in either the coast and mountains of and includes a poorly developed saltwater marsh northern BC kiff et al manuscript in prep- ofcordgrassofcordgrasscordgrass spartinaspartinafoliosafolfoifoliosafoliosepoilosaiosa gramineae glass- aration patten et al 1993 unitt et al 1995 wort sallsailsalicorniaSalicomia bigeloviibigelovii chenopodiaceae and lagoons of the pacific coast massey and palacios saltwort batis maritinamaritimamaritima plant names follow 1994 offshore islands anderson 1983 cody roberts 1989 A narrow mangrove commu- 1983 or southernmost regions from la paz to nity principally black mangrove avicennia the cape rodriquez estrella and riverariverarodrirodri germinantgerminansgerminans avicenniaceae and red mangrove quez 1992 carmona et al 1994 guzman et al rhizophora mangle rhizophoraceae occurs 1994 this report summarizes early spring irregularly along the estuary and on several 1996 and 1997 and late fall 1996 observations small islands within its boundaries above the for the area near eulegemuleg6mulege baja california sur high tide zone are scattered stands of mangle hereafter BCS dulce maytenus phyllanthoides celastraceae eulegemulegemuleg6 is located on the east coast of BCS lining the southern edge of the estuary are north of bahia concepci6nconcepcion and south of santa several retirementretiremenC communities consisting of rosalia straddling 2653n26 53n 11158w fig 1 permanent houses and semipermanent domi vehicular access is by means of mexico high- ciles formed by modifying various types of way 1 a reasonably maintained 2glanelane asphalt motor homes and trailers many residents pro- road passable during all but the wettest sea- vide supplemental food for birds in the form sons approximately 800 km south of tijuana of typical backyard grain feeders and hum- baja california norte hereafter BCN and mingmingbirdbird feeders filled with sugar water asso- 500 km north of the cape A landing strip at the ciated with these communities are numerous

division of forestry PO box 6125 west virginia university Morganmorgantowntown WVVVV 26506612526506 6125 portadopdrtado 10 oasis rio eulegemuleg6mulege eulegemulegemuleg6 baja california sur mexico

131 132 GREAT BASIN naturalist volume 57

west of the lagoon an arroyo meanders through the center of a broad valley the arroyo con- 2 tains water only during seasonally heavy rains 3.3 4 portions of the valley are heavily irrigated with

5.5 ancestral water and support citrus orchards 31 corn garlic onions alfalfa some truck crops and pasture annual rainfall averages 12 cm el 8.8 the average computed from numerous years

29 without rain interspersed with an occasional heavy downpour loio10.10 ornithologically the muleg6mulege lleile110 eulege region has re- 2 ceived only anecdotal attention wilbur per- 27 14 sonal communication howell and webb 1992 1 50 though the is home 3 16 even area to species 17 endemic to BC xantus hummingbird nomen- 18 clature follows the 1983 AOU checklistcheck list with 9 25 20 appropriate supplements see appendix for scientific names belding s yellowthroat and 21 gray thrasher in addition recent observations 2 23 indicate that a small colony of least brebesgrebes breed within the freshwater lagoon at eulegemulegemuleg6 howell and webb 1992 personal observation 11411 4 111122 11tio1100 and large concentrations of hooded oriole 1 1 1 breed in the palmar fig I1 outline map of 2 states in los estadesestados anidosunidos field data were collected during february de mexico baja california norte and baja california sur march april and november 1996 and febru- showing appiapplapproximateoximate position of prominent locations 1 ary and march 1997 the data consist of direct tijuana 2 Enensenadasenada 3 maneaderoManeadero 4 san felipe 5 observation enhanced with playbacks of santo domingo 6 bahia dcde san quintin 7 el rosariorosano 8 re- cataviacatavifiacatavma 9 bahia de los angeles 10 guerrero negro corded songscallssongscalls here we summarize new 11 san ignacio 12 santa rosaliarosalfa 13 eulegemulegemuleg6 14 bahia noteworthy or locale specific information for concepcion 15 la purisima 16 comondncomondfiComondn 17 loreto 17 species some of which may be considered 18 ciudad Insurgentinsurgentsinsmgentesinsurgenteses 19 bahia magdalenaMagdalenadaleDa 20 paz la common but for which no eulegemulegemuleg6 records are 21 la laguna 22 san jose del cabo provided in published sources eg nelson 1921 grinnell 1928 wilbur 1987 howell and ornamental plantings and scattered palms in- webb 1992 1995 in addition we provide tab- cluding native mexican fan palm washingtonia ular summarization of our data for all species rorobbustaustal arecaceae and introduced date palm observed during the 3 time periods precise phoenix dactyliferadactylifera arecaceae dense stands latitudelongitudelatitude longitude locations were determined of these species are locally referred to as the using GPS carmincarmin model 100 while approx- palmar imate hereafter apapproxprox locations were deter- the freshwater lagoon approximately 151.5lsis mined from an atlas of surface maps topo- kmkin long and 10 30 in wide depending on sea- graphic international inc 1986 son is formed by damming the main spring primarily to prevent tidal saltwater contamina- ANNOTATED SPECIES ACCOUNTS tion it supports a small community a plantation of date palms a dense stand of rush juncus least grebe observed at the freshwater sp Juncajuncacaejuncaceaecae and limited citrus and mango oasis each day it was visited in spring adults orchards the lagoon is highly impacted by were attracted to tape recordings of their ad- domestic cattle and pigs which run freely and vertisementverti sement and distress calls As many as 6 have trampled significant portions of the edge adults could be seen at I1 time within 50 in of habitat away from the water habitat abruptly the tape player all birds observed in march transforms into sarcosarcocaulescentcaulescent desert wig- were in full adult breeding plumage red eyes gins 1980 dominated by cercidium bursera and lack of a white throat two juveniles were bachyPachypachycereuspachycerenscereus and opuntia wilbur 1987 to the observed on 26 april 1996 this species was 199719971 BAJA california SUR MEXICO BIRD RECORDS 133

not recorded in brackish water of tidal rio from close range 10 in swimming in the eulegemulegemuleg6 based on playbacks we estimate that freshwater lagoon at eulegemuleg6mulege the furthest there were at least 8 pairs of least brebesgrebes in south for which wilbur 1987531987 53 provides the oasis we know of at least I1 successful nest- documentation is bahia san quintin approx ing attempt howell and webb 1992 also ob- 302rn3027n 1155rw11557w1155rW on the pacific coast the served them in the oasis and cited 2 other range map in howell and webb 19951551995 155 observations from 1987 and 1988 this species shows the transientwintertransient winter range of this species was formerly common in the cape region but only on the pacific coast side of the peninsula has declined in that area because of habitat although the west coast of mainland mexico is degradation wilbur 198732 33 since this included oasis is highly impacted by humans and appears anna s hummingbird we found this species to be degenerating and since this situation is to be common at feeders with documented sight common at most of the other BC oases wilbur records continuously from 29 february through 1987 grismer and mcguire 1993 there is early april 1996 when all hummingbird use of little hope for longtermlong term survival of the local feeders diminished howell and webb 1992 population report them in october north of eulegemulegemuleg6 at western grebe one observed at estero guerrero negro approx 2758n27058n IMWW san marcus a saltwater lagoon located approx on the west coast and south at the el tnpuitripuitrippitinpui 20 km northeast of eulegemuleg6mulege approxapprox 2707n27007n resort south ofloretoof loreto approx 26n26 N 11127u11127w lww11204w wilbur 198734 lists this species on the east coast wilbur 19871091987 109log reports that aslawluwoccasional in the gulf of california while the species is an uncommon resident south howell and webb 199597 include it as a non- to 30 on the mainland and to 28 on islands breeding wintering species less common in howell and webb 19954251995 425 state that amissanna s the gulf than on the pacific coast howell and hummingbirds winter south to about guer- webb 1992 provide 5 winter records 3 of rero negro the maximum number we recorded which are from the pacific coast 1 from the at any I1 time was 4 individuals 2 males and 2 interior and I1 from the cape region the loca- females on 3 march 1996 tion closest to eulegemulegemuleg6 is san ignacio approxapprox rufous hummingbird for this report we 2717n2717n112054w11254w assume that individuals with rufous tails and white faced ibis on 12 november 1996 green backs were alienallenailen s hummingbirds while we observed 4 individuals feeding in an irri- those with rufous tails and rufous backs were gated alfalfa field about 8 kmkin west of eulegemuleg6mulege rufous hummingbirds although this may be on 15 november 1996 at rio eulegemulegemuleg6 we ob- unreliable phillips 1975 kaufman 1990 from served 4 feeding individuals on exposed mud I1 march through 25 march 1996 they were flats at low tide it cannot be determined if abundant at feeders with peak numbers 7 these 2 sightingssightings were the same individuals males and 5 females seen at I1 time on 3 wilbur 198751 lists this species as a rare march the last individual 1 I1 female was seen transient providing 3 location records manea on 25 march when present we noted that dero approxapprox 3144n la purisima they dominated all other species at the feed- approxapprox 2612n 11204w112004fw112004lwfW and 3 records at ers including the much larger xantus hum- la paz approxapprox 2408n24 WN iktnw11017w an im- mingmingbirdbird howell and webb 19954281995 428 do not mature was recorded on 11 june 1991 at san include this species on their range map for josejosg del cabo approxapprox 2303n 10943w BCS while wilbur 19871091987 109log 110 states that howell and webb 1992 the range map in it is an uncommon spring and fall tran- howell and webb 1995147 includes all of sient the length of the peninsula none of baja as transientwinteringtransient wintering range but the wilbur s documented sites however include account does not provide documentation for eulegemulegemuleg6 phillips 1975 postulates a spring BCS migration route out of the state of sinaloa into greater white fronted goose on 11 sonora mexico and across the sea of cortez november 1996 nine individuals were in an at approximately 30n30 N our data indicate that irrigated alfalfa field approximately 10 km the crossover is at least 26 and possibly fur- west of eulegemulegemuleg6 they took off flying due east ther south down the valley toward the town on 14 alienallenailen s hummingbird observed almost november 1996 we observed 9 individuals daily from I1 march through 13 march 1996 134 GREAT BASIN naturalist volume 57 departing nearly 2 wk before the last rufous american robin one individual was ob- hummingbird howell and webb 1995 served by 3 people over a 2 h period on 15 428 429 do not include this species in BCS november 1996 at the rancho la kentanaventanaVentana while wilbur 19871101987 iloiio110 lists them as common approximately 10 km west of eulegemuleg6mulege in an to abundant transients in northwestern agricultural area the bird in question was ex- baja california presumably not BCS tremely gray in coloration and was initially phillips 1975 summarizes specimen locations identified by us as a female american robin in for this species noting only 6 spring speci- basic plumage however after examining our mens I1 each on 22 february 1925 at santo field notes field guide illustrations several domingo approx SOMGN 11557wliy57w on the american robin females collected in baja or pacific coast and 25 february 1925 at san nearby desert locales and 8 specimens of san quintin approx 302rn3027n 11557w on the lucas robins collected in baja we are left pacific coast and 4 on 2 march 1945 at isla undecided about the positive identification of cedros a continental island approx 2815n this bird the back wings nape crown and 11520w115 20w on the pacific side there were no forehead were too gray and the breast and specimen records for BCS and none for main-mainmaln abdomen were not rufous enough for a desert land mexico for april and may our data sup- area american robin female in unworn basic plumage abdomen port the contention that alienallenailen s hummingbirds however the breast and san complete their northward migration before were not as washed out as a typical rufous hummingbirds but not the supposition lucas robin in either event the record is of lists that they cross the sea of cortez at about 30 note since wilbur 1987132 american north robins as an uncommon migrant and winter visitor chiefly in the north with costa s hummingbird although it is widely supportive sightings including december at known that this species is the most abundant bahia de angeles approxapprox hummingbird in desert areas of BC we report los 29n in and may at laguna approxapprox 3 unusual locations was la lw2306n nest the ist site an in the cape region in addition23nhe active nest with 2 eggs that was constructed 10936w lists the san lucas robin as native only to on a of fishletfishnet material slung under a piece the mountains of the cape region howell and blue glass fishing float used as a decoration webb 1995592 include similar information hanging on a outside a home patio interest- from the above locations in their range maps ingly nest was adorned with pale blueblue the neither reference includes the area surround- fibers perhaps from a which plastic amatapifiatapipmatafiata ing eulegemuleg6mulege in addition american robins are nearly exactly matched glass of the float the listed as breeding in BCN howell and webb we discovered 2 the nest when it had eggs 1992 in summary the individual we observed both of which hatched and later fledged the was more than 200 km south of the published and2nd and ard3rd nests were located under the range of the american robin and more than edge of television satellite dishes adjacent to 300 km north of the range of the san lucas patios the contents of the ist costagosta s nest were robin the plumage did not fit perfectly unknown but the female was sitting tight from either of the two but was closer to the latter I1 march through 15 march 1996 its ultimate european starling during november we outcome is likewise unknown the and2nd nest observed this species daily in the agricultural contained 2 nestnestlingslings that were being fed as of valley west of eulegemulegemuleg6 this species was less 9 march 1997 common in spring of 1996 but during spring red breasted sapsucker one an adult 1997 many individuals were observed defend- male was observed on 3 consecutive days in ing potential hole nest locations amongst the early march 1997 in the mixed mesquite pal- eardoncardon pachycereusPachy cereus this is south of previ- mar habitat edging rio eulegemulegemuleg6 we place ously published winter sightings at guerrero these observations on record since wilbur negro approxapprox 2758n27058n IMWW wilbur 19871111987 iiiili111 112 states that they are a sparse 1987137 howell and webb 1992 1995613 winter visitor october to february throughout this species appears to be continuing its south- BC and provides only grinnell s 1928 ward colonization route location summaries none of which is within pyrrhuloxia observed commonly during 250 km of eulegemuleg6mulege spring 1996 and 1997 and fall 1996 in a variety 199719971 BAJA california SUR MEXICO BIRD RECORDS 135 of habitats including backyard feeders irri- 1987 this matches the range map given in gated cropland open desert and mixed scrub grinnell 19281761928 176 figure 13 for latitude contrary to grinnell 1928184 who confines 26 27n the range map in howell and webb the species to mesquite prosopis sp aimosmimos identifies the area east of the central moun- oideae associations eulegemulegemuleg6 is near the north- tains given the species songcallsong call especially in ern edge of the year round range of this and spring marchaprilMarch April it should have been easy the following species wilbur 1987148 how- to observe however we have no records from ell and webb 1995682 supporting spring early springspringlatespringwatelate fall 1996 although a single documentation for pyrrhuloxia includes very male did respond to a song playback on 2 dif- rare at santa rosalia approx 2720n ferent days at the freshwater lagoon in early IIGW approx 30 km north of eulegeMumulegemuleg6lege march 1997 song sparrows are known from 3 s fairly common at santa aglendaaguenda apapproxprox additional areas in BC northwest coastal moun- 2703n 11225w between santa rosalia tains south to el rosariorosarlorosano approx 3003n and eulegemuleg6mulegeMulege bahia agua verde approxapprox 11544w11c5044w rio colorado exact location un- 2528n liriywililii111 15w south of loreto and el known but approx 3150n315315ynyN IMW and a triunfo apapproxprox 2345n23 N ikriow11010w south of small population discovered recently in the la paz oasis at cataviacatavifiacatavma approx 2944n 11443w northern cardinal observed on several howell and pyle 1990 dates and at several locations including 3 western meadowlark this species was march 1996 in mixed shrub habitat 7 march heardseenheardheardyseenseen regularly march 1996 and 1997 1996 at a feeder in the oasis rio eulegemulegemuleg6 and and november 1996 in irrigated alfalfa fields captured on 8 march 1997 in a mist net located approx 10 km west of eulegemuleg6mulege they are known in desert scrub habitat 10 km west of town bleedersbreedersbreeders in northwestern BC with winter many of the contour feathers covering the back records south to san ignacio approx 2720n of this individual were either entirely gray or isow11250wesox wilbur 19871601987 160igo 161 howell and tipped with gray typical of desert dwelling webblsox1992 19957381995 738 include all of BCS as cardinals in basic plumage this gave a pale winter range and include I1 summer record cast to the overall ventral appearance northern from ciudad Insurgentinsurgentsinsurgenteses approx 2510n cardinals are common and widespread south lll4yv11145w taking advantage of recent agricul- of santa rosalia approxapprox 2720n27020n IIGW11216w tural development wilbur 1987148 howell and webb 1995 brown headed cowbird although none 681 682 although no records for eulegemulegemuleg6 are were observed during spring 1996 or 1997 a included small flock 20 individuals was observed on clay colored sparrow in november this 13 november and again on 15 november 1996 species was observed on multiple days often foraging in a heavily grazed alfalfa field in the within flocks of lark sparrows and brewer s agricultural valley approx 10 km west of sparrows foraging in the edges of irrigated eulegemulegemuleg6 although the range map in howell fields approx 10 km west of eulegemulegemuleg6 they are and webb 19957421995 742 places the year round 11 common winter visitors to the cape district range of this species throughout BC no docu- october to april only a few reports north mentation is provided wilbur 19871621987 162 lists ofoflat25lat 25 wilbur 1987153 documented this species as a regular winter visitor south to records include catavinacatavifiacanavinaCatavina apapproxprox 2944n the cape district with 1 supportive record for IMW and san ignacio approxapprox 2717n27017n la paz approx 2410n24010n IIOW inm janu- howell and webb s 1995716 ary but as a breeder onlyliowin BCN our data rangelwmap does not include the eulegemuleg6mulege area support the contention that cowbirds are win-win song sparrow the distribution of this spe- ter visivisitantstants to the eulegemuleg6mulege not year round cies in central baiabaja is problematic although residents listed in both wilbur 1987157 158 and howell and webb 1995725 as present at the acknowledgments same latitude as eulegemulegemuleg6 the text description in wilbur lists locations only west of the cen- we thank john and jane boyd russell and tral mountains including records from san joanne evans ted and ann manyk and gene ignacio apapproxprox 2730n 11250w south to and lydia tobias for access to their property comonducomond6compondu approxapprox 2610n26010n lllOW wilbur feeders and for calling our attention to unusual 136 GREAT BASIN naturalist volume 57 observations and anecdotal information we in the sea ofofcoitezcortez university of california press thank S R wilbur D W anderson S N G berkeley los angeles and london GRINNELL J 1928 A distributional howell J guzman E palacios B massey R summation of the orniorni- thology of lower california university of california rodriguez estrella and L FE kiff for advice publications in zoology 32 1 300 encouragement and updated observational GRISMER H L AND J A mcguire 1993 the oases of data we thank K parkes at the carnegie central baja california mexico part I1 A preliminary museum of natural pittsburgh penn- account of the relict mesophilic herpetofauna and history the sylvania status of the oases bulletin of the southern califor access to and advice concerning fornia academy of sciences 92 2 24 specimens of american robin san lucas GUZMAN J R CARMONA E PALACIOS AND M BOJORQUEZ robin and northern cardinal in addition we 1994 seasonal distribution of aquatic birds in estero thank M M campbell and 2 anonymous re- de san josejosg del cabo BCSB C S mexico cianciaciencia mar 20l932009120193201 93 103 viewers for constructive comments for the 1im- in HOWELL S N G AND S WEBB 1992 noteworthy bird provementprovement of earlier drafts we thank L gribko observations from baja california mexico western and J bell for assistance with figure 1 travel birds 23 153 163 funding for robert C whitmore was obtained 1995 A guide to the birds of mexico and northern from the college of agriculture and forestry central america oxford university press new york 851 appp at west virginia thank university finally we HOWELL S N G AND P PYLE 1990 additional notes on ruth M whitmore for continued patience and birds in baja california aves mexicanasMexicanas 290 1 6 7 cheerful support even in the early morning KAUFMAN K 1990 advanced birding petersen field guide no hours this manuscript is published with the series 39 houghton mifflin company boston approval of the director west virginia agri- MA KIFF L FE S R wllburWILBUR E MCMILLAN J C BORNEMAN cultural and forestry experiment station as K AXELSON AND J SCHMIDT in preparation budsbirds of scientific publication 2606 the san pedro martir baja california norte mexico MASSEY B W AND E PALACIOS 1994 avifauna of the wet- CITED lands of baja california mexico studies in avian literature biology 15 45 57 NELSON E D 1921 lower california and its natural AMERICANAMLRICAN ornithologists UNION 1983 checklistcheck list of resources national academy of sciences 16 north american birds ath6th edition alienallenailen press first memoirs 194 appp lawrence kansas 877 appp PATTEN M A K RADEMAKER AND T E WURSTER 1993 1985 thirty fifth supplement to the american noteworthy observations from northeastern baja cali- ornithologists union checklistcheck list of north american fornia western birds 24 89 93 birds auk 102 680 686 PHILLIPS A B 1975 the migrations of alienallenailenallersailers s and other 1987 thirty sixth supplement to the american hummingbirds condor 7719677 196 205 mthologistsornithologists union checklistcheck list of north american oi ROBERTS N C 1989 baja california plant field guide birds auk 104 591 596 natural history publishing company jolla CA 1989 seventh supplement to the la thirty american 309 appp oi nithologistsornithologists union checklistcheck list of north american RODRIGUEZ ESTRELLA R AND L RIVERA RODRIGUEZ 1992 birds auk 106 532 538 kleptoparasitism and other interactionsinter actions of crested 1991 thirty eighth supplement to the american caracara in the cape region baja california mexico nithologistsornithologists union checklistcheck list of north american oi journal of field ornithology 63 177 180 birds auk 108 750 54 topographic international INC 1986 baja topographic 1993 thirty ninth supplement to the american atlas directory box 5794 san clemente CA 92676- nithologistsornithologists list oi union checklistcheck of north american 8794 birds auk 110 675 682 UNFITr P A M REA E PALACIOS E MELLINK L ALFARO 1995 fortiethfolfoi tiethbieth supplement to the american orni unit AND S GONZALEZ 1995 noteworthy records of birds thologists union checklistcheck list of north american birds in northwestern baja california mexico western auk 112 819 830 in birds 26 144 154 ANDERSONANULRSON D W 1983 the seaseabirdsbirds pages 246 264 in WIGGINS 1 L 1980 flora of baja california stanford uni- T J case and M L cody editors island biogeogra- versity press stanford CA 1025 appp phy in the sea of cortez university of california WILBUR S R 1987 budsbirds of baja california university of press berkeley angeles and los london california press berkeley 253 appp CARMONA R J GUZMAN S RAMIREZ AND G FERNAN- DEZ 1994 bleedingbreeding waterwateibirdswaterbirdsbirds of paz bay baja la received 3 january 1997 california sur mexico western birds 25 151 157 accepted 2277 march 19919977 CODY M L 1983 the land birds pages 246 264 in T J case and M L cody editors island biogeography 199719971 BAJA california SUR MEXICO BIRD RECORDS 137

APPENDIX

summary of observational bird data arranged by habitat type abundant A seen daily with little or no effort com- C mon usually seen in appropriate habitat by experienced observer fairly common F seen more than once in appropriate habitat by experienced observer uncommon U seen in appropriate habitat by experienced observer looking specifically for that species raierare R seen once in appropriate habitat by experienced observer

habitat typee pelagic freshwater saltwater irrigated sarcocaulescentSarcocaulescent and shore lagoon3lagoons estuary 3 cropland 3 desert SpecieSpeciessl1 status2 sp fa sp fa sp fa sp fa sp fa least grebe re C C tachybaptus dominicus pied billed grebe re F IT podilymbus podiceps eared grebe mi A A podiceps nigrinigricollismgncolliscollis blue footed booby re C C sula nebouxii brown booby re A A sula leucogasterleuco gaster brown pelican re A A pelecanus occidentalisoccidentoccidentalistalis double crested cormorant re A A phalacrocorax auritusauntusauretus brandt s cormorant re C C phalacrocorax penicillatuspemcillatus magnificent Frigatefngatebirdfrigatebirdbird re A A fregata magnificmagnificentmagnificensens great blue heron re C C C C ardea herodias great egret re C C C C C C Casmercasmerodiusodius alba snowy egret re C C C C C C egretta thula little blue heron bere F F egretta caeruleacaecaeruledoaerulearuieafulearuled coloredtricoloredTn heron re U U egretta tricolor reddish egret re F F egretta rufescentrufescensrufescens cattle egret re F F F F bubulcusBubulcus ibis green backed heron re U U butoridesButorides stristnatusatus black crowned night heron un C C nycticorax yellow crowned night heron un F F nycticorax violaviolacenscens white faced ibis wi U U plegadis chihichahi greater white fronted goose wi R R anser albifronsalbifrons green winged teal wi U U anas crecca mallard wi F F F F anas platyrhynchosplatyrhynchous 138 GREAT BASIN naturalist volume 57

habitat type pelagic freshwater saltwater irrigated sarcocaulescentSarcocaulescent and shore lagoonslagoon3 estuary 3 croplandcropland33 desert SpecieSpeciessi1 status2 sp fa sp fa sp fa sp fa sp fa

northernnoi them pintail wi C C anas abutaacuta blue winged teal wi F F anas discorsdiscorddiscors cinnamon teal wi u u anas septentrionaliumseptentnonalium gadwall wi u u U U anas strepera american digeonwigeon wi C C anas americanaamencanaamencana redhead wi F aythya collanscolcoicollariscollaraslansianslaris ring necked duck wi F F F F aythya amenamericanacana lesserlessel scaup wi C C C C aythya affinis bufflehead wi F F bucephala albeola ruddy duck re C C oxyurajamaicensisoxyura jamaicensis osprey re A A F F A A pandion haliaetushahaetushaliahaliaeetusetus cooper s hawk re U U U U accipiter coopericoocooperiaperi northern harnelharrier re U circus cyaneuscycyaneousaneus harris hawk re U Paraparabuteobuteo uncinctuncinctusus red tailedtalled hawk re C C C C buteobuteojamaicensisjamaicensis amerleanamerican kestrel re C C C C falco sparsparveriussparvenusspderiusverlusveriusarvenus merlin wi u falco columbariuscolumbcolumbariumarius peregrineperegi me falcon re u falco peregrinus california quail re A A A A callipeplaCallipepla californicacahfornicacahcabcalicallcailfornica sordsora re F F U U porzana rohnacarolinaca common moorhenmoorten re F F galbgaibcalhnulaCalhgallinulanula chlochloropusropus american coot re A A C C fulica americanaamencana black bellied plover wi C C Pluviapluvialishs squatarola semipalmated plover wi F F charadriusCharadnus semipalmatus killdeer re C C F F charadriusCharadnus vocivociferusvociferousferus black necked stilt wi F F himantopus mexicanusmexicanus 199719971 BAJA california SUR MEXICO BIRD RECORDS 139

habitat type pelagic freshwater saltwater irrigated sarcocaulescentSarco caulescent and shore lagoonslagoon3 estuary 3 croplandcropland33 desert SpecieSpeciessl1 status2 sp fa sp fa sp fa sp fa sp fa american avocet wi U U recurvirostra americanaamencana greater yellowlegs wi U F U F tringa melanoleucamelanoleuca solitary sandpiper wi F F tringa solitariosolitariasolitaria willet wi C C C C cataptrophorus semipalmatus spotted sandpiper wi F F IT F actipisactitisActitis maculansmaculariamaculandmamacuculandlarialarlaiaria whimbrel wi F F IT F numeniusNumemus phaephaeopusopus long billed curlew wi F F F F IT F numeniusNumemus americamericanosamericanusamencanusanus marbled godwit wi F IT limosafedoalimosa fedoafedda sanderling wi U caicabCalcandriscahdriscalidrisCahdrisidris alba bonaparte s gull wi A A latuslarusLOTUS philadelphia heermanHeermaysmarss gull re C C C C larus beenheenheennanniheermanniheermanninanni yellow footed gull re C C C C larus livens elegant tern re C F C IT stemasterna elelegansdelegansegans white winged dove re F F C A C A zenaida asiaasiaticaasiaticaltica mourning dove re U U U U U U zenaida macrouramacroura common ground dove re F F F F C C F F columbina paspasserinasenna greater roadrunner re F F F F geogeococcyxCeococcyx californicalifornianscalifornianuscahformanusanus great horned owl re F IT F F F F bubo virginianusvirgimanusvirgivirginianuslanusmanusmanos xantus hummingbird re C C C C C C F F hylocharisHylocharis tanxanxantustustiii anna s hummingbird re F F F F calyptecalypter anna costa s hummingbird re A A A A A A A A calyptecalypter cistaecostae rufous hummingbird mi C C selasphorus rufus alienallenailen s hummingbird mi F F selasphorus sasin belted kingfisher wi F F ceryle alcyon gila woodpecker re A A A A C C C C melanerpes uropygiuropygialisalis red napedcaped sapsucker wi U sphyrapicus dehosdebosdariusvariusvanus 140 GREAT BASIN naturalist volume 57

habitat type

pelagic freshwaterfreshpreshwater saltwatersaltvsalav ater irngiengirrigatedatedabed sarcocaulescentsarcocaiSarco eaicaiealclescentnlescentcaulescent and shore lagoonslagoon3lagoiagoionaion3 estuaryestu ary3arya3 croalcroplandcropl andaand33 desertdeslert SpeciesSpeciespecies1sl1 status2 sp fa sp faea sp fa SP fa sp fa

ladder backed Woodpeckwoodpeckerei re F FIT F F ficoidesPicpicoidesoides scalansalaris gilded flicker re F F F F colantescolaptes chryssideschrysoideschrys oides clayglaygray flycatcher wi U F U F empidonax wrightii black phoebe wi C C sansayomisSaysayormsomisorms nigricansnigmgncansricans say s phoebe wi U U U U sayornissdyomisSayornis saya vermilion flycatcher re A A porocephaluspyrocephalus rulubinusrubinusbinus ash throated flycatcher re C C myiarchus cinerascens cassin s kingbird wi C C tyrannus vociferantvociferans violet green swallow re F F C C tachycineta thalassinathalasstnathalassinothalassina tree swallow wi C F tachycineta bicolor western scrub jay re F F F F aphelocomaAphelocoma califcalifomicacalifornicacalifornicapornicaomica common raven re A A A A corvus coray verdin re F F F F C C A A autauriparusflavicepsauripanisAuripanis flamflamcepsceps cactus wren re C C C C campylorhynchus brunneicapillus canyon wren re U U Catcatherpesherpes mexicanusmexicanus marshmaimal sh wren re C C cistothomscistothorusCistocistophorusthorusthoms papalustnspalustrislustris house wren re F F troglodytes aedonabdon blue gray gnatcatcher re F F C C polioptilapohoptila caecaenileacaerulearuleanilearuieafuleanoleanoiea california gnatcatcher re F F A A polioptilapohoptila califcalifomicacahformcaomica ameiamericanamerleanamer leanican robin wi R turdus migratormigratoriusmigratoriesius noinorthernthem mockingbird re F F F F C C mimus polyglotpolyglottospolyglottoustos gray thrasher re U U F F toxoxstoma cinereumcinereum phainopepla re U U F F F F phainopepla miens loggerhead shrike re C C F IT laniuslamus ludovicianusludomcwnusludovic ianuslanus european starling re F F stamusstumus vulgaris 199711997 BAJA california SUR MEXICO BIRD RECORDS 141

habifhabitat at type

pelagic freshfreshwaterpresh water saltwatersaltwsalawratertater irrigirrigatedatedabed sarcocaulescentSarcosarcocalSargarcocal descentcaulescent and shore lagoonslagoon3lagoiagoionaion3 estuestuaryary3arya3 croplandcropichopiandaand33 desertdesert SpecieSpeciessi1 status2 sp fa sp fa sp fa sp fa sp fa yellow rumpedbumped warbler wi C C C C C C dendroica coronata belding s yellowthroat re C C geothlypis beldingibeldingi orange crowned warbler wi F F F F F F hennwermvennivoravermivoraVennVermluoraivora celata northern cardinal re u U U U F F cardinalis pyrrhuloxia re F F F F F F cardinalis sinuatus green tailed towhee wi C C F F pipilo chloruruschlorurus clay colored sparrow un U spizella hallidapalpallidalida song sparrow un R melospiza melodia brewer s sparrow wi F F spizella brebreweribrewereweri black chinned sparrow re U spizella atrogularis lark sparrow wi A A chondestes grammacusgrammacus black throated sparrow re C C amphispiza bilineatebilineatabilineata white crowned sparrow wi A A A A C C zonotrichia leucleucophrysophrys western meadowlark un C C sturnella negneglectsneglectalecta brown headed cowbird un U molothrus ater hooded oriole re A A A A F F icterus cucullacucullatustus house finch re C C C C C C F F carpodacus mexicmexicanusanus lesser goldfinch re F F carduelis pspsaltriaaltria house sparrow re A A A A passer domesticusdomesticus icommonlcormnon and scientific names follow american ornithologists union checklistcheck list of north american birds 1983 and supplements 1985 1987 1989 1991 19931995 2rearee year round resident mi migrant wi winter resident un status uncertain including associated edge habitat within 50 m ofwatelof water creatgreat basin naturalist 572 0 1997 appp 142 148

interannualINTERANNUAL ABUNDANCE OF NONNATIVE FATHEAD MINNOWS pimephales PROMELAS IN UPPER KLAMATH LAKE OREGON

david C simoni and douglas FE markiel

ABSIRACIABS TRAur since its introduction about 20 yr ago fathead minnow pimephales kromelaspropromelasmelas has become very abundant in upper klamath lake oregon in 1991 mean beach seine catch per unit effort CPUE was 214 compared to 25 for native blue chub gila coecoerulearulea the next most abundant species in 45 trap net samples collected in 1992 fathead minmin- now constituted 59 of the fishes caught in agency lake subbasssubbasmsubbasinsubbasubbasinsm 27 in upper klamath lake and 17 in tributary inflow habitats fromplomprom 1991 to 1995 fathead minnow declined and the abundance of some native fishes increased introduction to klamath basin was coincident with USU S environmental protection agency promotion of fathead minnow as biobioassayassay subjects upper klamath lake fathead minnow have incomplete lateral lines and males have mandibular tuberclestubercledtub ercles diagnostic of the northeastern subspecies although the origin as bait bucket transfer forage fish or laboratory release cannot be determined with certainty the possibility of laboratory release suggests modification of bioassaybioassay protocols to requirelequire destruction of test 01or excess subjects

key words fathead minnow oregon nonnativenormative species species interactions baitfishbai0tshbattbaitfish introductions laboratory release

fathead minnow pimephales kromelaspropromelasmelas is fathead minnow has been introduced in broadly distributed east of the rocky mountains many western states as a bait and forage fish with a native range from great slave lake state sanctioned propagation and distribution canada to chihuahua mexico and eastward began in adjacent states of idaho about 1945 to the appalachians vandermeer 1966 lee simpson and wallace 1978 and california in and shute 1980 hubbs and black 1947 rec- 1953 shapovalov et al 1959 published infor- ognized 2 broadly distributed subspecies P p mation suggests early introductions of fathead kromelaspropromelasmelas a midwestern to northeastern form minnow as bait and forage fish were P p con with an incomplete lateral line and nuptial pertus the southwestern form evans and tuberclestubtubercledercles on the lower jaw of breeding males douglas 1950 miller 1952 minckley 1973 and P p confertusconfertus a southwestern form with a the northeastern form of fathead minnow P complete lateral line and no nuptial tubtuberclestubercledercles p kromelaspropromelasmelas was promoted as a biobioassayassay subject on the lower jaw by the US environmental protection agency fathead minnow was first collected from EPA beginning in the late 1960s EPA 1972 oregon in spencer creek a small tributary to brauhn and schoettger 1975 klamath river 27 km downstream of upper in this paper we document relative abun- klamath lake on 13 may 1974 andreasen dance of fathead minnow in upper klamath 1975 first reported in upper klamath lake in lake from 1991 to 1995 discuss possible im- 1979 ziller 1991 it was occasionally captured pacts on native fish and compare the fish fauna there in 1982 by oregon department of fish before and after its introduction we also com- and wildlife ODFW by 1983 fathead min- ment on diagnostic morphological characters now was captured in each ODFW trap net set of upper klamath lake fathead minnow and J ziller fisheries biologist ODFW personal suggest possible origins communication and by 1993 we had captured the species from klamath marsh national wild- description OF AREA life refuge 50 km northeast of upper klamath lake and gerber reservoir 50 km southeast upper klamath lake the headwaters of the of upper klamath lake unpublished data klamath river is in south central oregon east

dcpjitmentdepartnient of fisheries and wildlife 104 nash hall oregon state university corvallis OR 97331380397331 3803

142 199719971 KLAMATH FATHEAD MINNOWS 143 of the cascades fig 1 at 1262 m it drains an average of 333.3 1 5 s 1411.41 tuberclestubtubercledercles and 9415 km2 of mixed forest grass shrub flatland rio grande specimens had no chin tubtuberclestubercledercles and marshes johnson et al 1985 the william- vandermeer 1966 showed a sharp break in son river is the major tributary providing lateral line completeness with populations north about half the inflow while its northern sub and east of kansas being 222.2 49949.9 complete basin agency lake is fed by the wood river and those to the southwest 63063.0 95095.0 com- johnson et al 1985 it is the largest lake in plete he considered this northeast vs south- oregon with a mean summer surface area of west pattern to present a classical pattern of 27811 ha 141 km of shoreline and a mean subspecific variation he also showed that man- depth of 242.4 m US army corps of engineers dibular nuptial tuberclestubercledtubercles on breeding males 1978 johnson et al 1985 the lake is hyper were virtually absent south of kansas and high eutrophic highly turbid and caustic ph typi- to the east with 2 populations around lake cally 90go909.0 with wide diel fluctuations in dis- michigan having mean chin tubercle counts of solved oxygen from near depletion to super- 434.3 and 11411.4 vandermeer 1966 upper klamath saturation bortleson and fretwell 1993 high lake fathead minnows appear to be the north- ph and anoxia are thought to be partially eastern subspecies P p kromelaspropromelasmelas rather than responsible for the declining populations of 2 the southwestern form P p confertusconfertus but this endangered catostomids lost river sucker identification should be considered tentative deltistesDeltistes luxatusluxatus and shortnoseshorthoseshortnose sucker chas until the systematics of fathead minnow is bet- bistesmistes brevirostris scoppettone and vinyard ter resolved 1991 A ard3rd catostomid in the klamath basin klamath larglargescaleescale sucker catostomus sny METHODS deri was formerly a US fish and wildlife beach seine service category 2 candidate species beach seine sampling was conducted in 1991 subspecies identification 1993 and 1995 as part of a longtermlong term sampling program of sucker year class strength ten sites we identified the subspecies of fathead min- in upper klamath lake and 5 in agency lake now in upper klamath lake using criteria of fig 1 were sampled with a 61 m long seine hubbs and black 1947 a complete lateral line with 48 mm bar mesh and a 2 x 2 X 2 m bag A and absence of mandibular nuptial tuberclestubercledtubercles sampling unit was a 14 circle arc that sampled on breeding males to diagnose P p confertusconfertus 30 m2ma sites were sampled weekly from 25 july and an incomplete lateral line and presence of to 26 september 1991 though the full com- mandibular nuptial tuberclestubtubercledercles on breeding males plement of sites was typically not sampled as to diagnose P p kromelaspropromelasmelas we implemented other aspects of our catosto- the following museum specimens were ex- mid research sites were sampled monthly in amined oregon klamath drainage oregon july and august 1993 and biweekly in july and state university OS 4944 OS 7953 OS 7955 august 1995 OS 12506 OS 14116 OS 14326 OS 14327 analysis of variance ANOVA was used to minnesota great lakes drainage OS 14274 test for differences in relative abundance among from stocks maintained by EPA environmen- years for fathead minnow blue chub gila tal research laboratory duluth and new coeruleacoerulea tui chub cilacilacliagila bicolor and ageageo 0 mexico rio crandegrande drainage museum of sucker both species catch per unit effort southwestern biology university of new mex- CPUE because catch data frequency distri- ico MSB MSB 735 butions were highly positively skewed data average lateral line completeness pored were 109loglogg transformed CPUEI to better total lateral line scales was 41 range 6 83 meet assumptions of normality required for standard deviation s 0180.18 in klamath 41 ANOVA least significant difference LSD test 7 98 s 0290.29 in great lakes and 95 was used on mean logg109log chuelcpuelcpue1CPUEl values to 81 100 s 0070.07 in rio grande fathead identify interinterannualannual differences in abundance minnows breeding males from upper klamath for significant ANOVA models all analyses lake had an average of 434.3 2 7 s 1451.45 tu were performed using statgraphicsStatgraphics version 7 bereles on the chin great lakes specimens had manuguistics 1993 144 GREAT BASIN naturalist volume 57

WA wood riveriv OR williamson river

CA agency B sprague river T ason T C lake B T T recreation B creek T crystal creek B

T T

B B T T T B T B B B T

upper B klamath

lake B

klamathkjamath fallsfalisfails

fig 1 map showing upper klamath lake and agency lake oregon shoreline beach seine sites are indicated by a B and trap net sites are indicated by a T

trap net usually set overnight and fished for approximately 24 h but set times ranged from 155iss15.5 to trap net sampling was conducted from 21 99.0990 h we found no significant P 0050.05oos r agency 990 april to 22 october 1992 sites were in 00370.037 relationship between number of hours lake the northern half of upper klamath lake fished and logelogg transformed number of fish and tributary inflows lower williamson river caught so we calculated trap net CPUE as thomason creek crystal creek and recrea- number per net tion creek fig 1 trap nets were 65 mm bar the only available data for evaluating his- mesh had a single 242.4 X 23 m lead two 242.4 X torical change are 1964 and 1965 surveys in 107 m wings and a 121.2 X 12 m square frame upper klamath lake by vincent 1968 vin- with two 10 cm throats lake trap nets were cent s data and analyses are difficult to inter- set offshore in open water river mouth trap pret because he reported pooled results from nets were set in mid channel trap nets were a gang of 3 gill nets and a single hoop net his 199719971 KLAMATH FATHEAD MINNOWS 145 gill nets had stretched mesh sizes 32 45 57 4waw 2waw and 152 mm but he gave no mesh size for the fathead IN blue chub hoop net with I1 exception he reported results minnow 120 as percent composition rather than in termsofterms of 2waw effort we make comparisons of our trap net data to vincent s data but caution that our data 40 are not directly comparable to his and only LU ol010 C 19911981 im1893 1985 18811991 1993 1995 crude comparisons are useful &IL im im im

RESULTS beach seine ge 60 76 50 was tuitintultun ageageo 0 fathead minnow the most common chub 40 species in upper klamath beach seines suckers lake 30 during 1991 CPUE 2144214.4 coefficient of 20 variation CV 270 followed by blue chub 10 CPUE 24924.9 CV 190 and tui chub 4 0 96gg CPUE 969.6 CV 266 fathead minnow iggi19911981 100319931893 im1895 iggi1991 im im oos density declined significantly P 0050.05 each year year from 1991 to 1995 while blue chub and ageageo 0 sucker increased significantly P 0050.05oos fig 2 mean and 95 confidence intervals of beach from 1991 to 1995 fig 2 tui chub CPUE seine CPUE for fathead minnow blue chub tuitintultun chub and treadedtrended upward each year but was not signifi- ageageo 0 sucker from upper klamath lake 1991 1995 cant P 005oos0.05 except for yellow perch perca flavescentflavescensflavescens in 1993 other species never ex- catch rates was much higher in tributariestributaries than ceeded 2 of the total catch in any year yellow in lake samples table 1 perch was 7 of the 1993 catch all of which compared with vincent s 1968 data our were ageageo 0 included other native species trap net data table 2 are noteworthy in the marbled sculpin cottus klamathensisk1amathensis slender coftuscottuscoitus presence of fathead minnow however with- sculpin cottus tenuis and klamath scul- lake out knowledge of the mesh size of vincent s pin Cottcottustis princeps nonnative species included trap nets we cannot determine whether fathead yellow bluegill perch lepomis macrochirusmacrochires minnow were vulnerable to his gear in our trap and pumpkinseed lepomis gibbosusgibbo sus nets fathead minnow constituted 17 of the agency in lake beach seine samples fat- fish in tributatributariesries 27 in upper klamath lake head minnow CPUE also decreased signifi- and 59 in agency lake because fathead min-min cantly P 0050.05oos from 1991 CPUE 2196219.6 now now represents a substantial portion of the CV 179 to 1995 CPUE 1085108.5 CV fish fauna percentages of other species must 346 with 1993 intermediate CPUE 1858185.8 decline if densities remain constant vincent CV 161 changes in abundance of native 1968 mentioned tui chub was caught at a species were unclear with no significant P maximum rate of about 5 per hour h in july 0050.05oos differences among years for blue chub tui and december based on his relative species chub or ageageo 0 sucker CPUE in agency lake composition table 2 vincent s maximum rate trap net for blue chub must have been about 83h8 3hah our 1992 upper klamath lake trap net catch fish were captured during 45 trap net sam- rates were 11iili1.11 1 tui chubachubh and 18918.918 9 blue chubb ples 11 from upper klamath lake 13 from indicating that tui chub may have declined after agency lake and 21 from tributary inflow the introduction of fathead minnow however areas fathead minnow was the most abundant we found up to a 5 fold increase in intermterannualinterannualannual species captured in agency lake and2nd in beach seine abundance for tui chub from 1991 abundance in upper klamath lake and ath4th in CPUE 969.69gg 6 to 1995 CPUE 48648.648 6 and a abundance in tributariestributaries table 1 agency 4 fold increase in blue chub from 1991 CPUE lake trap nets caught fewer species 10 than 2492424.99 to 1995 CPUE 105 in our surveys upper klamath lake 13 or tributariestributaries 14 thus apparent changes in abundance from the but total catch rates were higher variation in 1960s in both chub species could be attributed 146 GREAT BASIN naturalist volume 57

IABLITABLE 1 species mean catch number per net median TABLE 2 comparison of percent species composition catch number per net and coefficient of variation CV for 1992 trap net samples from this study with data of the mean foifolforhorbor fish caught in trap nets from upper kla- reported from upper klamath lake by vincent 1968 math lake agency lake and ti ibutarytributary inflow habitats in UKL upper klamath lake AL agency lake and TI 1992 tributary inflow habitats species mean median CV species TI AL UKL vincent or species group 1968 UPPERUPPLH klamarilklamvriiKLAMAril LAKELAKL blue chub 4841 4170 88 lampreys 2 I1 I1 I1 fathead minnow 1752 600 125195 trouts I1 tuitintulten chub 410 240 133 brown bullbuilbulibullheadheadbead 13 I1 1 5 klamath lake sculpin 65 10 175 tuitultuichubchub 10 9 7 35 marbled sculpin 57 40 144 blue chub 36 20 62 58 slender sculpin 14 0 244 fathead minnow 17 59 27 pacific lamprey 05 0 228 suckers 1 I1 I1 I1 suckelsuckerss1sa 04 0 185 sculpstulpinssculpinssculpmsins 13 2 4 1 yellow pelpeiperchch 01 0 331 yellow perch 8 9 1 2 bibrownown bullhead 01 0 332 sunfish 1 1 I1 pumpkinseedpumpkmseed 01 0 332 bluegill 01 0 332 7150 6250 82

agencyAGI NC Y LAKELAKL attributed the decline of lost river and short fathead minnow 5598 3560 log109 nose suckers to competition andor predation blue chub 1355 880 104 by nonnative species most nonnative species tun chub 592 700 77 tuitintul in upper klamath lake bluegill pumpkinseed yellow perchpelper clioiioil 218 0 249 marbled sculpin 23 10 146 yellow perch and brown bullhead ameiurusameirusAmeirus klamath lake sculpin 13 0 193 nebulosus are relatively rare and density pacific lampilampreyey 111 1 0 180 dependent impacts on suckers might be small 205 brownbi own bullhead 03 0 however fathead minnow is still abundant even sucsuesuckershsuckelkershs 01 0 361 syr the slender sculpin 01 0 361 after a 5 yr decline although fathead min-min pumpkinseed 0 0 now reportedly consumes algae higher plants bluegill 0 0 zooplankton and insects becker 1983 duns- 7815 7930 81 moor 1993 has experimental evidence that it IIUBUIARY INFLOWinain1 LOW iiabi1a1iiabrrxrs s feeds on sucker larvae in aquaria additionally blue chub 1058 130 237 franzin and harbicht 1992 found walleye stiz brownbi own bullhead 390 0 309 ostostedionedion vitreum larvae in fathead minnow tuichubtuitul chub 174 70 203 stomachs from a manitoba irrigation drainage fathead minnow 96 30 149 has a density dependent yellow pelpeiperchch 46 20 128 if fathead minnow marbled sculpin 38 10 154 impact on suckers or other native fishes we slickersuckerssc 28 10 206 might expect a compensatory response to the Klarnathklamathklarnath lake sculpin 25 0 201 fathead decline we observed from 1991 199 minnow pumpkinseed 10 0 mechanisms influencing pacific lampreylampi ey 07 0 183 to 1995 extraneous slender sculpin 01 0 458 the direction of change offishof fish abundances from bluegill 01 0 458 1991 to 1995 are unknown because upper kla- 1873 970 168 math lake is managed by the USU S bureau of reclamation as an irrigation reservoir lake levels lost riverRIVLI indand shortshortnoseslioitnoscshorthosenose luekersticker blostI1 ost riverRIVLI sticker are regulated however there is no active man- largescale guckel alostulostoslrivciriver shorshortrioseslioitnosctriose indand kliimtlikharriath largluesluesciloescaleeileciloetie suckelsuckersueker agement to reduce fathead minnow abundance only 2 active management strategies have been to fathead minnow introduction natural popu- implemented as a consequence of endangered lation variation or susceptibility to poorer water species listing of lost river and shortnoseshorthoseshortnose quality castleberry and cech 1993 sucker 1 closure of a snag fishery on adult sucker and 2 prolonged elevation of spring- discussion time water levels the ist strategy would seem to have little impact on fathead minnow abun- with the possible exception of native cato dance the and2nd inundates sucker spawning stostomidsmids the fish community of upper klamath gravels and shoreline vegetation and was de- lake appears resilient williams 1988 partly signed to provide cover to reduce larval sucker 199719971 KLAMATH FATHEAD MINNOWS 147 vulnerability to predation however prolonged be argued that states govern access distribu- springtime water levels could also promote fat- tion and handling of fishes within their borders head minnow abundance by providing com- but a laboratory s record for following EPA pro- plex structural spawning habitat such as under tocols may be a greater incentive than poorly surfaces of rocks logs and vegetation scott enforced state regulations and the costs of and crossman 1973 becker 1983 although disastrous introductions far outweigh the min- the trends evident in figure 2 suggest that ele- imal costs associated with destruction of test vated springtime water levels had the desired organisms all protocols for bioassaybioassay of exotic effect by increasing young sucker abundance organisms should specify destruction of all test they did not increase fathead abundance and all excess individuals at the end of the native fishes have responded positively to bioassaybioassay declines of nonnativesnonnatives in other systems such as the great lakes jude and tesar 1985 eck acknowledgments and wells 1987 north carolina streams lemly 1985 and great smoky mountains national this work was supported in part by con- park streams moore et al 1984 however we tracts from the klamath falls office of USBRUSER found no documentation of responses of west- the denver office ofusbrof USBR the portland office ern US native fishes where spread of non ofusof US fish and wildlife service and the ore- natives has seriously compromised many gon state university agriculture experiment native fish populations minckley and douglas station we are grateful to mark buettner and 1991 rinne and minckley 1991 we have sharon campbell USBR for logistics support demonstrated in upper klamath lake that and information and larry dunsmoor kla- decreasing abundance of nonnative fathead math tribe for discussions of klamath fathead minnow is associated with increasing abun- minnows we thank dan logan erik lesko dance of some native fishes and marcus beck for field assistance fathead finally diagnostic characteristics of klamath minnows from EPA ERL duluth were pro- fathead minnows indicate they are of north- vided by vince matson this is oregon state east origin possibly including the great lakes university agriculture experiment station con- region where EPA began use and promotion tributiontribution no 11128 of fathead minnow as a standard bioassaybioassay subject Govcovgovernmentalemmental and other laboratories using literature CITED fathead minnow in biobioassayassay work as well as baitshopsbaitshops obtain supplies from multiple sources ANDREASEN J K 1975 occurrence of the fathead minnow often the lowest known pimephales kromelaspropromelasmelas in oregon california fish and bidder thus it is not game whether there have been laboratory releases 613 155 156 BECKER G C 1983 fishes of wisconsin university of of fathead minnow and if so whether these wisconsin press madison could be distinguished from baitfishbaitfish release BORTLESON G C AND M 0 FRETWELL 1993 A review however their detection in oregon was coin- of possible causes of nutrient enrichment and decline cident with the promotion of fathead minnow of endangered sucker populations in upper klamath lake oregon USU S geological survey water by EPA and 21 29 yr after their introduction resources investigations report 93408793 4087 portland as baitbaitfishfish in neighboring idaho and california OR shapovalov et al 1959 simpson and wallace BRAUHN J L AND R A schoettger 1975 acquisition 1978 and culture of research fish rainbow trout fathead minnows channel catfish and bluegillsblue gills ecological potential for biobioassay subjects minnows the nonnative assay research reports EPAE PA 6603 75 011 to be released and the irony that it might castleberry D T AND J J CECH JR 1993 critical happen in the furtherance of environmental pro- thermal maxima and oxygen minima of five fishes tection suggests a prudent countermeasure from the upper klamath basin california fish and game 78 145 152 although the EPAEPXs s published protocols spec- DEGRAEVE G M ET AL 1991 variability the perfor- ify incoming in initial quarantine of incomingincomin 9 fish they do mance of the seven day fathead minnow pimephales not specify disposition of excess or test fathead kromelaspropromelasmelas larval survival and growth test an intra minnow except in some larval assaysessays where and interlaboratory study environmental toxicology specimen fixation in formalin is part of the and chemistry 10 1189 1203 dinsmoorDUNSMOOR L 1993 laboratory studies of fathead min- assay brauhn and schoettger 1975 norberg now predation on catostomid larvae klamath tribes and mount 1985 degraeve et al 1991 it can research report KT 93 01 chiloquin OR 148 GREAT BASIN naturalist volume 57

ECK G W AND L WELLS 1987 recent changes in lake in response to removal of sympatricsympatnc rainbow trout in Michimichigangarisgarlss fish community and their probable causes great smoky mountain national park journal of the with emphasis on the role of the alewife alosa tennessee academy of science 594 76 77 pseudoharengus canadian journal of fisheries and NORBERG T J AND D I1 MOUNT 1985 A new fathead aquatic sciences 44 supplsuppiauppl 2 53 60 minnow pimephales kromelaspropromelasmelas subchronicsubchronic toxicity EPA USU S environmental protection AGENCY 1972 test environmentalenanenvn onmental toxicology and chemistry 4 recommended biobioassayassay procedure for fathead minmin- 711 718 nows pimephales kromelaspropromelasmelas rafmesquerafinesqueRafinesque chronic tests RINNE J N AND W L MINCKLEY 1991 native fishes of national water quality laboratory duluth MN and land a dwindling resource of the desert south- EVANS W A AND P A DOUGLAS 1950 notes on fishes west general technical report RM 206 USU S depart- recently introduced into southern california cali- ment of agricultureofagriculture forest service rocky mountain fornia fish and game 36 435 436 forest and range experiment station fort collins FRANZIN W G AND S M HARBICHT 1992 tests of drift CO 45 appp samplers for estimating abundance of recently hatched SCOPscoppettonescoppetronePETrONE G G AND G VINYARD 1991 life history walleye larvae in small rivers north american jour- and management of four endangered lacustrine nal of fisheries management 12 396 405 suckers pages 359 377 inireoneonn W L minckley and J E HUBBS C L AND J D BLACK 1947 revision of cera deacon editors battle against extinction native fish tichthysttchthystichthys a genus of american cyprinid fishes mis- management in the american west university of cellaneouscel publications of the museum of zoology arizona press tucson and london universityuniveiuniver sity of michigan no 66 SCOTT W B AND E J CROSSMAN 1973 freshwater fishes JOIINSONJOHNSON D M R R PETERSENPELERSEN D R LYCAN J W SWEET of canada fisheries research bulletin of canada M E NEUHAUS AND A L SCHAEDEL 1985 atlas of ottawa 184 1 966 oregon lakes oregon state university press corvallis shapovalov L W A DILL AND A J CORDONE 1959 A JUDE D J AND F J TESAR 1985 recent changes in the revised check list of the freshwater and anadromous inshore forage fish of lake michigan canadian jour- fishes of california california fish and game 45 nal of fisheries and aquatic sciences 42 1154 1157 159 180 lelLEELLL D S AND J R SHUTE 1980 pimephales kromelaspropromelasmelas SIMPSONSIMPSONJJ CANDRC AND R L WALLACE 1978 fishes of idaho rafmesquerafinesquerabinRafinesque fathead minnow page 341 in D S lee university press of idaho moscow C R gilbert C H hocutt R E jenkins D E US ARMY CORPS OF ENGINEERS 1978 klamath river mcallister and J R stauffer jr editors atlas of basin oregon reconnaissance report san francisco north american freshwater fishes north carolina district biological survey publication 1980121980 12 north car- vandermeer J H 1966 statistical analysis of geographicofgeographic olina state museum of natural history variation of the fathead minnow pimephales bromeprome lemlyLLMLY A D 1985 suppression of native fish populations las copela 1966 457 466 by green sunfish in first order streams on piedmont VINCENT D T 1968 the influence of some environmental northn01 th caicalcarolinaolina transactions of the american fish- factors on the distribution offishesof fishes in upper klamath eries society 114 705 712 lake unpublished master s thesis department of manuguistics INC 1993 statgraphicsStatgraphics version 7 rock- fisheries and wildlife oregon state university ville MD corvallis MILLER R R 1952 bait fishes of the lower colorado WILLIAMS J E 1988 endangered and threatened wild- river from lake mead nevada to yuma arizona life and plants shortnosesbortnoseshorthoseshortnose and lost river suckers with a key for their identification california fish houghton s goldenrod and pitcher s thistle final and game 38 7 42 rules federal register 53137 27129 27133 MINCKLEYMINCKLLY W L 1973 fishes ofarizonaof arizona sims printing co ZILLER J S 1991 factors that limit survival and produc- inc phoenix AZ tion of largemouth bass in upper klamath and MINCKLEY W L AND M E DOUGLAS 1991 discovery agency lakes oregon information report 91691 6 fish and extinction of western fishes a blink of the eye in division oregon department of fish and wildlife geologic time pages 7 17 min W L minckley and J E portland deacon editors battle against extinction native fish management in the american west university of ari- received 12 february 1996 zona press tucson and london accepted 23 january 19919977 MOOREMOORL S E G L LARSENLARSLN AND B RIDLEY 1984 A summary of changing standing crops of native brook trout great basin naturalist 572 V 1997 appp 149 154

WINTER HABITAT SELECTION BY reintroduced PRONGHORN ON ANTELOPE ISLAND GREAT SALT LAKE UTAH

melissa J klikiiKilKilgorekilgorelkilgore1kilgoreegorel1 and W sue Fairbanks 2

ABSTRACT the recent and future introduction of several ungulate species on antelope island necessitates knowledge of habitat use by each species in this study habitat preferences of reintroduced pronghorn antilocapra americanaemenamenoanacanaoane on antelope island were evaluated during february marchmaich 1993 and january march 1994 elevation slope physiography aspect and habitat type of sites used by prongpronghornhornborn were compared to similar data collected from random points during the severe winter of 1993 pronghorn preferred terrain that was 1281 1380 m in elevation and was relatively flat or at the base of a hill slopes greater than 30 were avoided south facing slopes were preferredprefened west facing slopes were avoided pronghorn preferred sagebrush habitats and avoided grasslands during the mild winter of 1994 pronghorn showed preferences for slightly higher elevations avoided slopes greater than 30 but used other habitat features in pioproportionproportion to their availability future winter studies of pronghorn should include considerations of snowfall patterns and the availability versus the abundance of sagebrush

key words pronghorn antelope antilocapra americanaamencana habitat selection winter habitat habitat availability

with the introduction ofpronghornofpronghorn into new utah provided an excellent opportunity to areas many questions must be answered con- study winter habitat selection of pronghorn cerning their winter habitat preferences factors the recent and future introduction of other such as vegetation availability and diversity ungulaungulatesungulatedtes on the island necessitates knowledge snow depth and physiography may play an im- of critical pronghorn winter habitat to mini- portant role in their survival past habitat stud- mize potential interspecific competition iden- ies have considered these factors but many tification of these critical areas also can be neglected to examine a particular feature s used in planning construction of hiking and availability when results were interpreted but bike trails to avoid human use of prime winter see amstrup 1978 clary and beale 1983 ryder habitat the objective of this study was to and irwin 1987 availability is important be- determine which factors affect winter habitat cause observed habitat use by pronghorn may preferences of pronghorn and thus to provide simply reflect the occurrence of habitat types data for better management of pronghorn on in the area only when animals use certain fea- antelope island tures out of proportion to their availability can they be said to prefer or avoid those features STUDY AREA marcum and loftsgaarden 1980 optimal winter habitat with ample vegeta- antelope island state park in the south- tion is critical for pronghorn survival A severe eastern region of the great salt lake is about winter with heavy snowfall and low tempera- 16 km from salt lake city utah fig 1 the tures can result in higher mortality from mal- island was closed to the public from 1983 to nutrition martinka 1967 van wormer 1969 1993 because of rising lake levels that flooded and may lead to a low production of young the the 11611ilg 6 km causeway leading to the island following spring martinka 1967 additionally the 10409 ha island is topographically di- deep snow can hinder winter movements by verse ranging in elevation from 1280 to 2011 m pronghorn van wormer 1969 mitchell 1980 jones 1985 utah department of natural kindschy et al 1982 affect vegetation avail- resources 1988 average annual precipitation ability hovey and harestad 1992 and increase is 43 cm with 10810.810 8 cm falling january march predation risk van wormer 1969 national weather service forecast office the reintroduction of pronghorn onto ante- NWSFONWSFOI salt lake city international air- lope island state park in the great salt lake port salt lake city utah most of the island

ldepartmentepaitment of zoology weber state university ogden UT 84408250584408 2505 present address 2675 NE lancaster st 52 corvalliscoicol vallis OR 97330 2depaltment2department of zoology weber state university ogden UT 84408250584408 2505 author to whom correspondence should be addressed

149 150 GREAT BASIN naturalist volume 57

N duceddeuced to the island in may 1993 and 10 fawns born in may and june 1993 were included fence with gates with the original 23 pronghorn in the 1994 field season prior to release onto the island each animal was fitted with a solar powered buffaloB radio transmitter advanced telemetry systems point P inc box 398 isantibisanti MN 55040 mounted on a standard livestock ear tag pronghorn were tracked february march r M 6 1993 and january march 1994 in all weather 10 conditions using radiotelemetryradiotelemetry to locate 2 3 d week TU groups data were collected per between 0700 and 1800 h no animal was included in more than I1 group on the same day and groups were noted as separate only if they were separated by 05050.5 km pronghorn observations stopped when winter groups began to 0 1 km disperse and bucks began establishing territories 31 march 1993 and 20 march 1994 the visual location of each group was marked on a topographic map vege- fig 1 antelope island state park study area in the great dominant salt lake utah Piongpronghornpionghornpronghorn moved from north to south tation physiography and elevation were noted through several open gates along the fence on the north later the site was revisited to measure slope side oftheodtheof the island using a clinometer and aspect north 315 44 east 45 134 south 135 224 or west 225 314 slope was categorized into 2 groups is grassland with patches of sagebrush arte- is 30 and 30 because slopes 30 were sppapp and a few small and marsh misia riparian scarce this also facilitated comparison with areas streams run on the island intermittent previous studies kindschy et al 1978 in and summer seasons average tem- in spring categorization of physiography and vegeta- peraperaturestures for january february and march of tion type occupied was subjective but consis- 1993 were 3 ac 1 14cac and 7 ac respec- 39c9c 4c 75c5c tent we categorized the physiography of each tively snowfall was 1278127.8127 8 cm in january 33533.533 5 point as relatively flat little or no slope slope cm in february and a trace in march for win- in of hill top of hill base of hill or terrace ter 1994 temperatures averaged 2 ac in jan- dom- 27c7c inant vegetation was categorized as grassland uary 1 ac in february and 7 ac in march 18c8c 76c6c few or no sagebrush bushes or trees sage- snowfall was 15715.715 7 cm 38138.138 1 cm and 797.97 9 cm respectively NWSFO brush some grass and forbs sagebrush dunes antelope island was named by john C fre- sand dunes where sagebrush predominates or shrub woody other than sage- mont in the 1840s for the herds of pronghorn vegetation only inhabiting it however the last pronghorn on brush we used grassland and sagebrush statistical analysis due to lim- the island was sighted in 1870 M A larsson habitats in the ited availability of other types personal communication the island also is vegetation even inhabited by about 500 bison bison bison and when all other types were combined into a 70 mule deer odocoileus hemionus with pop- single group ulatulationsions of coyotes caniscams latranslalatranotrans bobcats lynx air temperature at the time of the sighting snow rufus and golden eagles aquila chrysaetos of each pronghorn group was recorded as potential predators of pronghornofpronghorn depth at the site was recorded by relating the depth of snow to the length of the prongpronghornhordss MATERIALSMAFERIALS AND METHODS hind legs wind direction and velocity data were obtained from the closest weather station twenty three pronghorn were captured with to the island the salt lake international air- a net gun in summit county utah and report NWSFO about 26 kinkm southeast afanteofanteof ante- leased on antelope island 30 january 1993 lope island wind conditions at the weather three bucks from morgan county utah intro station and on the island were comparable as 199711997 WINTER HABITAT SELECTION BY PRONGHORN 151 both occur on the valley floor equidistance pronghorn observations at different aspects from the mountain range readings are taken using kruskal wallis aspect groups were com- hourly at the station and we used values clos- bined into north and east facing and south- est to the times of pronghorn group sightings and west facing due to small numbers of sites used by pronghorn were compared to pronghorn observations at each aspect north 76 randomly selected sites to determine habi- and east facing aspects were grouped because tat preferences we determined the number of these slopes are usually colder than south- and random sites by estimating that 20 sightings of west facing aspects on which snow melts more pronghorn groups would be made per month quickly snow depth in areas used by prong- over the 2 winter study period when deter- horn was summarized but not analyzed statis- mining random point distribution we consid- tically because availability data were collected ered the whole island because it was impossi- in the summer ble to define a priori the boundaries of available range RESULTS random points were chosen using a random number table and a grid 60 x 60 in super- in winter 1993 47 observations of prong- imposed onto a 75 minute series topographic horn groups were made each group consisted map once the location in the field was of 2 23 pronghorn mean s 14 8 ani- approached as closely as possible with a topo- mals forty pronghorn groups were observed graphic map we selected the exact spot for in winter 1994 groups consisted of 1 36 data collection by releasing a rock on a string pronghorn mean s 11 llI1 I1 animals that had been swung overhead at each of these snowfall was heavysiisilsllllsllduring 1993 totaling random points the elevation slope aspect 1613161.3 cm in january and february NWSFO dominant vegetation and physiography were by 6 march snow was patchy and by 15 recorded march it remained only at higher elevations availability of habitat features as determined snow depth at the sites where pronghorn were by random point data was compared to actual observed ranged from 0 to 10 cm pronghorn pronghorn sightings with chi square analysis were found in areas free of snow 48 of the to determine if pronghorn were using a site in time and in areas with snow 52 of the time proportion to its availability use availability n 21 observations of groups snowfall on aX of 0050.05oos or not in proportion to availability the island was minimal during winter 1994 a use or availability if pronghorn used a total of 61761.7 cm in january february and habitat less than expected they were avoiding march NWSFO when snow did fall it melted the feature if the habitat was used more than within 1 2 d consequently pronghorn were expected the pronghorn were choosing the observed in snow only 4 times feature following a significant difference as during 1993 pronghorn groups occupied indicated by chi square we analyzed elevation sagebrush habitats in greater proportion than physiography slope dominant vegetation and their availability and grasslands less than their aspect data with bonferroniBonferroni Z tests to deter- availability fig 2aaa x2xa2 434.3 df 1 P 0050.05oos mine which categories of each feature were in contrast no preferences were observed in used significantly more or less than expected pronghorn use of vegetation types during 1994 based on availability marcum and loftsgaar- xaoc221dfx2 212.1 df 11p005P 0050.05oos den 1980 in all bonferroniBonferroni Z tests 97 con- during winter 1993 pronghorn preferred fidence intervals were used for individual cate- elevations of 1281 1380 in and avoided those gories and simultaneous a varied with the from 1481 to 1880 in fig 213 x2xa2 158iss15.8 df number of categories in the analyzed feature 3 FP 0010.01 during 1994 pronghorn pre- wind velocity and current temperature each ferred elevations of 1381 1480 in and avoided 2 were compared to physiography data and ana- all higher elevations Y 989.8 df 3 P lyzed using the kruskal wallis test to determine 0050.05oos during 1993 pronghorn groups used if pronghorn use of physiographic features was south facing aspects in greater proportion than related to weather conditions to test whether their availability and west facing aspects less pronghorn were likely to use certain aspects than their availability fig 2cac x2xay2 969.6gg df 3 more than others when wind velocities were P 0050.05oos pronghorn groups showed no pref- high we calculated median wind velocities for erences or avoidancesavoidances in use of aspect during 152 GREAT BASIN naturalist volume 57

A loo100

60 1993 observations RANDOM POINTS 40 1994 observations W 0

0 GRASSLAND SAGEBRUSH DOMINANT vegetation C loo100 loo100 80 80 60 60 40 40 20 W 20 C D 0 0 NORTH EAST UTH WEST 1281 m 1381 m 1481 m 1581 m 31544 45 134 1w224 225 314 1380m 1480m 1580m 1880m ASPECT D ELEVATION E 100 100 80 i 80 caCD 60 60 40 oa 40 01LU 20 in ilnailga 20 0 FLAT BASE SLOPE TOP inTERRACE 0 30 30 physiography SLOPE lridividualindividual p003

fig 2 use versus availability of habitat characteristics by prongpronghornhornborn on antelope island great salt lake utah during the winters of 1993 and 1994 use referslefersrehers to prongpronghornhornborn observations availability refers to collected random point data 1993 n 47 observations 1994 n 40 observations the winter of 1994 xax22 11liii1.1 df 3 P temperature and pronghorn use of physio- 0050.05oos graphic features were not related 1993 H relatively flat areas and bases of hills were 5935.93 df 4 FP 0050.05oos 1994 H 3843.84 df 4 used during 1993 in greater proportion than FP 0050.05 nor was the use of physiographic their availability slopes of hills were avoided features related to wind velocity 1993 H fig 21ad2d x2xa5c2 22422.4 df 4 P ooi0010.01 in con- 4334.33 df 4 FP 0050.05 1994 H 4574.57 df 4 trast pronghorn groups exhibited no prefer- FP 0050.05 wind velocity during 1993 was not ences or avoidancesavoidances of physiographic features correlated with aspects of terrain used by during the winter of 1994 xax2 404.0 df 4 P pronghorn H 0590.59 df 1 FP 0050.05 how- 005oos0.05 pronghorn groups avoided slopes 30 ever during the winter of 1994 pronghorn use during the winters of 1993 fig 2eae x2xa2 666.6gg of north and east aspects was associated with df 1 P 0020.02 and 1994 xax22 51sisl5.1 df 1 higher wind velocities H 4174.17 df 1 FP P 0050.05oos during both 1993 and 1994 current 0050.05 the median and modal wind direction 199719971 WINTER HABITAT SELECTION BY PRONGHORN 153 was 150 ie from the southeast therefore survival during a severe winter moss and use of north and east aspects was probably not vaughn 1996 future plans for the island due to thermoregulatorythermo regulatory constraints should include habitat restoration with the planting of sagebrush on south facing aspects discussion additionally studies that incorporate the geographic information system cisgisCISGIS with habitat pronghorn exhibited more habitat prefer- preferences of all park ungulaungulatesungulatedtes would be ences and avoidancesavoidances during winter 1993 than beneficial in planning the development of trail 1994 due to heavy snowfall that limited vege- systems and other human uses on the island tation availability use of the lowest elevations eg koeln et al 1994 lachowski et al 1994 and relatively flat areas during 1993 may have bosakowski et al 1995 moss and vaughn in aided pronghorn in predator avoidance and press detection van wormer 1969 bruns 1977 As many as 15 mule deer and 100 bison were vaughan 1986 pronghorn use of slopes 30 seen in the vicinity of prongpronghornhom during winter was consistent both years and with previous in addition bighorn sheep ovis canadensis studies amstrup 1978 kindschy et al 1978 are to be introduced to the state park in janu- and also may reflect predator avoidance behav- ary march 1997 thus competition for food iors however we have no direct evidence to and space may be an important consideration support or refute this explanation as there was especially during severe winters pronghorn no winter mortality despite frequent sightings appear to graze compatibly with bison deer of coyotes in the area and bighorn sheep yoakum 1980 and in some particularly during a severe winter snow areas mule deer and prongpronghornhom have little depth can affect the amount of forage available overlap in habitat use during winter wood for ungulate consumption hovey and harestad 1989 however with few sagebrush areas 1992 pronghorn apparently preferred sage- limited space and increasing ungulate popula- brush habitats in 1993 because grassland areas tions on the island it is critical to continue to were buried in deep snow pronghorn in alberta monitor habitat use to examine future compe- were seen pawing through snow to reach food tition for food and space even when other vegetation was available above the snow ogara 1978 but we did not ob- acknowledgments serve that behavior in this study additionally funding and logistical support were pro- higher concentrations of fats and proteins in in vided by the zoology at weber sagebrush than in cured grasses may make department state university and antelope island state park sagebrush a preferred winter forage martinka S greathouse M tsukamoto C fowerspowers and 1967 sundstrom et al 1973 but the absence S wead provided valuable field assistance of habitat preferences in 1994 when there was special thanks to M larsson J fillpot M little snow suggests accessibility was the main rorick K zuby and the staff on antelope preference factor island for the help and support of this project pronghorn preference for south facing S amstrup C marti N smith G wurst S aspects may to release be related their near an zeveloff and 2 anonymous reviewers made known point 1 east west ridge as buffalo fig helpful comments on earlier drafts of this the severe winter deep snow and uncertainty manuscript of being in a new habitat could have caused the animals to remain in this location where sage- literature CITED brush was readily available on a south facing slope however their preference for south fac AMSTRUP S C 1978 activities and habitat use of prong ing aspects during winter 1993 was consistent homsbornshorns on montana wyoming coal lands pages 270306270 306 with previous studies clary and beale 1983 in proceedings of the eighth biennial pronghornpronghom antelope workshop jasper alberta canada the warmer aspects provide faster snowbeltsnowsnowmeltmelt bosakowski T A D RAMSEY AND D G SMITH 1995 thus increasing food availability GIS analysis of land use patterns and nesting den- on 4 august 1994 most of buffalo point sity of Swainswainsonsorss and red tailed hawks in northern burned including almost all of the sagebrush utah journal of raptor research 29 50 abstract BRUNS E H 1977 winter behavior of prongpronghornshorns in few other areas on antelope island have south relation to habitat journal of wildlife management facing aspects and sagebrush for pronghorn 4156041 560 571 154 GREAT BASIN naturalist volume 57

CLARY W P AND D M BEALEBLALE 1983 pronghorn reactions MITCHELL G J 1980 the pronghorn antelope in alberta to winteiwinterwintee sheep grazing plant communities and university of regina saskatchewan 165 appp topography in the great basin journal of range moss S AND D M VAUGHN in press using a GISCIS to management 36 749 752 assess winter habitat sites of pronghorn antelope on HOVEYHOVLY FE W AND A S HARESTAD 1992 estimating effects antelope island utah bulletin of the association of of snow on shrub availability for black tailed deer in north dakota geographers southwestern british columbia wildlife society OGARA B W 1978 antilocapra americanaamenoanacana mammalian bulletin 20 308 313 species 90 1 7 lonesionesJONES C D 1985 A manual of the vascular flora of ante- RYDER T J AND L L IRWIN 1987 winter habitat rela- lope island state paikpalkpark davis co utah unpublished tiontionshipsships of prongpronghornshorns in southeentralsouthcentralsouth central wyoming thesis brigham young university provo UT 110 appp journal of wildlife management 51 79 85 klndsiiyrkindschyKINDschyCuy R RRCC sunds1romandjSUNDSTROM AND J D YOAKUM 1978 SUNDSTROM C W G HEPWORTH AND K L DIEM 1973 Rangerangewildlifercingewildlifewildlife interrelationsintenanten elatelationsrelationsionslons prongpronghoiprongbornpronghornhornbornhoihol n antelope abundance distribution and food habits of the pages 216 269 in proceedings of the eighth biennial pronghorn wyoming game and fish commission pronghornpronghoihor n antelope workshop jasper alberta cheyenne 61 appp canada UTAH department orOF NATURAL RESOURCES 1988 geol- 1982 wildlife habitats in managed rangelands ogy and antelope island state park utah utah the great basin of southeastern oregon prongpronghornshorns department of natural resources utah geological USDA forest selvleeservice general technical report and mineral survey 18 appp PNW 145 pacific northwest forest and range VAN WORMER J 1969 the world of pronghorn J B lip- experimental station portland OR 18 appp pincott company philadelphia PA 191 appp koelnKOLLN G T L M COWARDIN AND L L STRONG 1994 VAUGHAN T A 1986 mammalogy ard3rd edition saunders geographic information systems pages 540 566 in college publishing fort worth TX 576 appp T A bookhout editor research and management WOOD A K 1989 comparative distribution and habitat techniques foifor wildlife and habitats ath5th edition the use by antelope and mule deer journal of mammal- wildlife society bethesda MD 740 appp ogy 70 335 340 LACHOWSKI H M T WIRTH P MAUS AND P AVERS YOAKUM J 1980 habitat management guides for the 1994 remote sensing and GIS thentheir role in ecosys- american pronghorn antelope USU S department of tem management journal of forestry 928 39 40 interior bureau of land management technical MARCUM C L AND D 0 loftsgaarden 1980 A non note 347 denver CO 77 appp mapping technique for studying habitat preferences journal of wildlife management 44 963 968 received 18 september 1996 MARIINKAMARTINKA C J 1967 mortality of northern montana accepted 4 february 1997 prongpronghornshorns in a severe winter journal of wildlife management 31 159 164 great basin naturalist 572 0 1997 appp 155 162 classification AND ordination OF COLEOGYNE communities IN SOUTHERN NEVADA

simon A leillellleli and lawrence R walkerlwalkerswalker1

ABSTRACT woody plant community composition was analyzed throughout the range coleogyneofcoleogyneof ramosissima in the spring and sheep mountain ranges of southern nevada the lower coleogyne elevational boundary was analyzed in detail in lucky strike canyon on the eastern edge of the spring mountains TWINSPAN 2 way indicator species analysis identified 4 piprimaryunary species and stand groups from the 2 mountain ranges extensive survey and 4 primary species and stand groups in the detailed study at the lower coleogyne ecotone intensive survey analysis of DECORANA detrendeddetrended correspondence analysis results indicated that elevation and soil depth were the environmental factors most significantly associated with distribution of species and stand groups in the extensive survey elevation was the only sig- nificantnificant physical factor associated with distribution of species and stand groups in the intensive survey five vegetation zones from the 2 mountain ranges were identified based on their dominant species in 15 extensive transects coleogyne were subdivided into pure stands and upper and lower ecotonesecotones for further investigation of species distribution and envi- ronronmentalmental factors selected environmental factors appear to play an important role in structuring the mojave desert vegetation zones in southern nevada

key words coleogyne ramosissima torrtomtowtorn classification ordination vegetation zones lucky strike canyon spring mountains sheep range

three dominant vegetation types character- distribution of coleogyne in utah may thus ize southern nevada larrea tridentata cov depend largely upon abiotic factors however ambrosia dubosadumosa payne valley floors and lower little is known about the factors that deter- mountain slopes coleogyne ramosissima torr mine the pattern of coleogyne distribution in mid elevation slopes and pinus monomonophyllaphylla southern nevada torrtorn & fremrem juniperus osteosperma little in this study we determined the spatial upper elevation slopes the coleogyne vege- arrangement of larrea ambrosia coleogyne tation zone ranging from 1200 to 1550 m in and pinus juniperus vegetation along an ele- elevation in southern nevada is characterized vatvationalional gradient and the percent cover for all by a closely spaced matrix of coleogyne with a woody perennial species we also classified scattered distribution of other shrub species vegetation using TWINSPAN and determined previous studies have shown that air tempera- the relationship between the distribution of tures in the coleogyne community in utah range desert plant species groups and environmental from 24 to 47c47 C korthuis 1988 korthuis factors based on correlations with axis values 1988 proposed that the upper limit of coleoboleo from DECORANA gyne distribution may be set by low air tem- peraperaturestures and that cold air draining from adja- STUDY AREA cent mountain slopes may limit coleogyne establishment on basin floors alternatively the this study was located in the spring 360n lower limit of coleogyne may be determined 11530w and sheep 3550n 11535w by low soil moisture in utah bowns 1973 mountain ranges approximately 65lsswkm north- annual precipitation ranging from 180 to over west of las vegas nevada precipitation patterns 270 mm appears to be required for develop- of southern nevada include summer storms and ment of coleogyne stands in the mojave desert winter rains summer storms generally occur hunter and mcauliffe 1994 shallow soils are in july and august and can sometimes be local typical of coleogyne communities and may and intense winter rain is widespread and may partially determine its abundance and distri- last up to several days snow is frequent at high bution in utah callison and brotherson 1985 elevations particularly in the pinus juniperus

idepartment of biological sciences university of nevada las vegas las vegas NV 89154400489154 4004

155 156 GREAT BASIN naturalist volume 57 woodlands and upper coleogyne ecotonesecotones pre- diameters were calculated by computing the cipitation is positively correlated with increase average of the longest and shortest dimension in elevation rowlands et al 1977 of the plant cover elevation and aspect of southern nevada is an area of temperature each plot were measured we visually estimated extremes with a mean minimum winter tem- percentage of rock soil and vegetation cover peraperatureture of 10clog and summer temperatures and assigned each plot to I1 of 6 cover categories of above 47c47 C temperature means and ex- absent 1 5 6 25 26 50 51 75 and tremes are negatively correlated with increas- 76 100 we classified topography of each plot ing elevation rowlands et al 1977 relative as slope terrace or dry wash repositioning humidity is low 20 is common in the sum- when cliffs rocky areas and streamstreambedsbeds were mer months resulting in exceedingly high encountered new plots were positioned hori- evaporation zonzontallytally approximately 20 m from the transect at an identical elevation to avoid these habitats METHODS ground surface was characterized as desert pavement loose rocks sand and sand with broad patterns of vegetation zonation were boulders examined with 12 elevational transects located using TWINSPAN hill 1979a we classi- on the spring mountains and 3 on the sheep fied sampled plots into vegetation types and range in summer 1993 the transects con- generated groups of species based on vegetation tained 9 16 circular 100 m2ma 5655 65 m radius similarities among the sampled stands each plots spaced at a fixed elevational interval of stand group represented a vegetation zone 65 m average number of plots per transect was from the lowest to the highest elevational 12 transects were located east west and south groups in the part of the spring and sheep of the spring mountains and east and west of mountain ranges sampled stand group 1 was the sheep range each transect included 2 characterized by larrea ambrosia shrublandsshrublands plots in the larrea ambrosia shrubland just while stand group 4 was characterized by pinus below the coleogyne shrubland mean elevation juniperus woodlands using TWINSPAN analy- 1155 40 m lei and walker 1997 extended sis we divided the sampled plots into 2 groups throughout the entire coleogyne zone mean each of which was then divided again and so elevation 1560 45 m lei and walker 1997 on the dichotomy was terminated if there and included 2 plots in the pinus juniperus were 4 or fewer plots in a group kent and woodland just above the coleogyne zone mean coker 1992 kent and coker 1992 further elevation 1870 50 m lei and walker 1997 proposed that the size of the eigenvalue gen- A portion of the lower ecotonal area of cole erated at each dichotomy by TWINSPAN anal- obyneogyne was examined in detail at the location of ysis reflects the importance of each compo- one of the original transects on the eastern side nent in explaining the total variation within the of the spring mountains six replicate 100 m2ma data set the eigenvalue ranging from 0 to 1 circular plots were located at each of 6 eleva- was largest at the initial dichotomy and became tions at 30 m elevational intervals in lucky smaller with each successive dichotomy one strike canyon between 1160 and 1310 m ele- sampled plot from red rock canyon of the vation the lowest elevation included 6 plots spring mountains was eliminated due to the just below the lower elevational limit of cole existence of numerous rare riparian species obyneogyne the remaining 5 elevations represented we used DECORANA hill 1979b to gen- increasing levels of coleogyne density but the erate species and stand ordination scores coleogyne shrubland extends to 1600 m in these scores were based on the percent cover lucky strike canyon lei and walker 1997 value of each woody species each point on data collected min these intensive plots were the diagram corresponds to a species and dis- also collected in the extensive plots that cov- tances between points on the graph are approx- ered the entire elevational range of coleogyne imations of their degree of similarity kent within each plot we recorded the presence and coker 1992 distances between points on of all woody perennial species 10 cm tall the diagram increased as species distributions including subshrubssubshrubs subshrubsSubshrubs aiealeare plants that diverged and as species occupied different have suffrutescent stems at the base with herba- vegetation zones similarly stand ordination ceous stems making up the canopy canopy scores were also generated by DECORANA 199719971 COLEOGYNE GRADIENT ANALYSIS IN SOUTHERN NEVADA 157 and the interpretation resembled species ordi- RESULTS nation scores distances between points increased as stand distributions diverged stand four major species groups from the 56 woody scores could be matched with several environ- taxa were identified by TWINSPAN analysis mental variables to detect vegetation and plot for the extensive transects in the spring and variation in relation to the environment envi- sheep mountains fig 1 species in group A are ronronmentalmental variables used in our study included typical of larrea ambrosia stands found at the elevation topography soil depth plot aspect lower limit of coleogyne they include lar- type of ground surface and percent soil and rea tridentata ambrosia dubosadumosa ephedra neva rock cover stand aspect could not be correlated densisbensis yucca schidigeraschidigera and acamptopappus because all stands occurred on northwest fac shockleyishockleyi these species tend to occupy slopes ing slopes in lucky strike canyon similarilysimilaritySimilarily and terraces at low elevations species in group percent rock cover could not be correlated B are typical of nearly monomonospecificspecific stands of because all stands had a rock cover of over coleogyne and include yucca brevifoliabrevifolia prunus 75 the ist and and2nd ordination axes were fasiculatafasiculata and thamnosma montana species orthogonal and indicative of different sources in group C typical of the upper coleogyne of environmental variation ecotone are represented by atriplex canescentcanescenscanes cens classification and ordination techniques Gutiergutierreziagutierrezbarezia sarothraesarothrae and chrysothamnus nau were applied to a total of 180 sampled plots on selsusseosus although Gutiergutierreziagutierrezbarezia sarothsarothraerae exists in the 15 elevational transects from the spring coleogyne stands it is particularly abundant at and sheep mountain ranges extensive plots the upper coleogyne ecotone species in group and 36 plots at the lower elevational limit of D are found in the pure pinus juniperus wood- coleogyne in lucky strike canyon intensive lands with artemisia tridentata as the major plots understory species pinus juniperus are the 2

SPRING AND SHEEP MOUNTAIN RANGES

101.0lo 0 9

018 070.7 060.6og 0 5 eigenvalue 04040.4

0 3 020.2 F L010.1oi 0.0 I1 E7 sadoi AMD U GRSP ACSH crrcpr A PC P attATC ACGR ATCA ARPU ARTR AGUT AMUT FONE DYCO LATR KRPA EPNE OPRA MESP LYAN CHLI SAME PRFA OPBA CHNA ERAN JUOS PIMO CEGR RHTR MATO GACO PSFR OPPO COME THMO YUBA GUSA EPVI CELE STPA KRPA ENVI YUBR STPI FAPA OPAC HYSA PSCO MIFR MOUT

A B C D

SPECIES GROUPS

fig 1 dendrogram of species groups identified by TWINSPAN from 15 elevational transects in the spring and sheep mountain ranges main groups were arranged by elevation from the lowest group A to the highest group D species are arranged alphabetically within groups see appendix I1 for species abbreviations 158 GREAT BASIN naturalist volume 57

dominantcodominantco genera in these woodlands with with coleogyne as the dominant species and juniperus osteosperma more abundant at lower crayla spinosa as a common associated species elevations and pinus monomonophyllaphylla at higher ele- hymenocleaHymen oclea salsola exists in or near the edges vationsvations associated species including fallugiaFallugia of washes within the coleogyne dominated parparadoraparadoxaadoxa ephedra virviridisidis Cercocercocarpuscarpus ledi stands in this group folius and forsellesiaFors ellesia nevadensis are also pre- analysis of the 180 extensive and 36 inten- sent but with significantly fewer individuals sive plots using DECORANA resulted in sig- four main species groups from 26 woody nificantnificant segregation of stand groups from taxa were identified by TWINSPANTWIN SPAN analysis TWINTWINSPANSPAN along axis 1 but not along axis 2 for the intensive plots at the lower boundary fig 3aaa pearsons correlation analysis indi- of the coleogyne shrubland in lucky strike cated that axis I1 of the stand ordination is sig- canyon fig 2 species in group A are typical nificnificantlyantly correlated P 00010.001 with eleva- of larrea ambrosia stands dominated by lar- tion soil depth topography and percent soil rea tridentata and yucca schidigeraschidigera species in cover in the spring and sheep mountain ranges group B typical of the lower half of the lower table 1 axis 2 is significantly correlated with coleogyne ecotone and dominated by ambrosia elevation topography and type of ground sur- dubosadumosa are generally short shrubs species face table 1 in lucky strike canyon both in group C are typical of the upper half of axis 1 r 0900.90ogo P 00010.001 and axis 2 r the lower ecotone with yucca brevifoliabrevifolia and 0400.40 P 00010.001 of the stand ordination are acamptopappus shockleshockleyiyi as the most abun- significantly correlated with elevation table dant species species in group D are typical of 2 DECORANADECOBANA also showed a significant stand the nearly monomonospecificspecific coleogyne stands group segregation along axis 1 but not along

LUCKY STRIKE CANYON

riot 101.0loio 09og090.9 08080.8 07070.7 06og060.6 05050.5 eigenvalue 04040.4 F 03030.3 02020.2 oloi0010.11 10 00oo000.00 OPEC ENVI LATR FAPFAPA AMDU KRPA YUBA ACSH CORA ECPO SAME ERFA OPRA OPAC CELA LYAN GRSP HYSA PSFR PSCO EPNE MATO OPBA YUSC MESP YUBR

A B C D

SPECIES GROUPS

fig 2 dendrogram of species groups identified by TWINSPAN from 6 elevations throughout the lower coleogyne ecotone in lucky strike canyon major groups were arranged by elevation from the lowest group A to the highest group D species are arranged alphabetically within groups see appendix I1 for species abbreviations 199719971 COLEOGYNE GRADIENT ANALYSIS IN SOUTHERN NEVADA 159

1 pearsons 500 1 1 TABLE 1 results of correlation analysis by A SPRING AND SHEEP MOUNTAIN RANGES matching the ist and and2nd axis of stand ordination scores STANDS acquired fromhiomhrom DECORANA to various physical factors in 400 the spring and sheep mountain ranges fig 1aaa r is the 0 GROUP 1 coefficient of linear correlation 0 GROUP 2 300joo u GROUP 3 factor ist axis and2nd axis V 0 0 m GROUP 4 X elevation 0790 79 0241024 2001 soil depth 054054lsl 0034 percent soil cover 0.35003535 0 lons10IONS 0 0 035 00 topography 033 0240.240 24 1 100 00 0 ground surface 0OIONSionslonsIQNS 020020ns plot aspect 010ns0 1 0 03ns003ns percent rock cover 003ns0 03 010ns0 10 0 I1 L- I goo 0 200 00 600 800 P ooi0010 01 AXIS 1 pp00010001 NS nonsignificant

600goo B SPRNGSPRING AND SHEEP MOUNTAIN RANGES and soil depth in the spring and sheep moun- SPECIES 500 tain ranges four major species groups were 0 GROUP A also distributed along the lower coleogyne eco- 400 THMOTHMOOO OPBA 0 GROUP B 0 WSEEPNE canyon C tone in lucky strike and were domi- EULA 0a GROUP MESP by dumosa 300 0 GROUP D nated larrea tridentata ambrosia dubosa SAMEOGDSAME- QS AMDU E coRA LYA 10 buosjuos IATR yucca brevifoliabrevifolia and coleogyne ramosissima X poPIMO ARTR 0 latr lucca 200 0 0 PAPAFAPA wiirYUBR YUSC respectively determined by elevation EPVII 1 I1 0 OGRSPGRSP CAN 0 100 CHNAg YUBA in southern nevada coleogyne shrublandsshrub lands 0 ENVI gusan HYSA establish at mid elevations with well drained apeceopeceOPECO0 0 0 SADO colluvial slopes between elevations of 1300 prfaePRFA and 2100 coleogyne stands rarely establish loo1001 in 200 0 200 400 600 boo800 above 2000 in unless they are situated on south AXIS 1 facing slopes coleogyne sometimes establishes below 1300 on north facing slopes and fig 3 ordination of stand A and species B groups in lei determined by TWINSPAN from the spring and sheep walker 1997 callison and brotherson 1985 mountain ranges across the entire elevational range of noted that coleogyne is the most abundant coleogyne some uncommon species and overlapping species in its community and may contribute points were eliminated for ease of interpretation and visumisuvisu- over 75 of the total vegetation cover while alizationaliza inm the ordination diagram see appendix tion species associated contribute less than I1 for species abbreviations species may 15 our data confirmed that the coleogyne plant community is nearly monomonospecificspecific with axis 2 in lucky strike canyon fig 4aaa correl- other species comprising less than 28 of the ations of stand groups vary with each environ- total number of species among the 15 transects mental factor figures 313 and ab4b413 illustrate the on 2 mountain ranges our data show that distribution of TWINSPANoftwinspan species groups and coleogyne is a dominant shrub in its vegeta- individual species along DECORANA axes I1 tion type in terms of height and cover many and 2 in the spring and sheep mountain ranges shrub species occur around the periphery of and lucky strike canyon respectively reflect- coleogyne canopiescanopies because coleogyne roots ing differences in the occurrence of varioussarioussarlous do not release chemical toxins to repress the plant species over different elevations and vege- growth and development of other nearby species tation zones sampled in southern nevada bowns and west 1976 which is in contrast to larrea roots mahall and callaway 1992 cacti discussion are sparsely distributed in the coleogyne shrub lands many cacti are vulnerable to prolonged four major species groups that were domi- winter freezing temperatures larson 1977 nated by ambrosia dubosadumosa coleogyne ramo the greatest woody plant diversity is found in sissimamissimasissima CutierrezcutlercutierreziaGutiergutierreziagutierrezbareziaia sarothraesarothrae and pinus mono or near edges of washes within the larrea phylla respectively were found to be distrib- ambrosia community and the lower coleogyne uted along gradients determined by elevation community 160 GREAT BASIN naturalist volume 57

tableIABLE 2 results of pearsons coicol relationcorrelation analysis by 300joo 1 1 matching the ist and and2nd axis of stand ordination scolesscores A LUCKY STRIKE CANYON STANDS 0 GROUP I1 acquired from DECORANA to various physical factors in 250 the lower coleogyne elevational boundary at lucky strike 9 GROUP 23 0a canyon fig 2aaa r is the coefficient of linear correlation A 0 GROUP 3 200 m GROUP 4 A GROUP 5 factorpactol ist axis and2nd axis 411 A c A A A GROUP 6 LI 150 6& elevation 0900.900ogo90 0400.400 40s1 A 0 ground 0 28 0 26ns026ns 0 OF6 A smsurfaceface 028ns & D peipel cent soil cover 024ns0 24 0 28ns028ns 100 0 A soil depth 017ns017 0 03ns003ns CL 0 topography 0003ns0 003 033033ns 50 A plot aspect 0 percentpeipel cent rock cover 0n 1 n 1 i 0 50 100 115050 200 250 300 P 005oos0 05 iai9 p001p 0001 AXIS 1 NS nonsignificantnonsinitil nt coefficicocflincntt c iniiotcannot helielye computed mdand coicoleolconfatedcondatedcon iddateditceitcd

500 LUCKS the lower coleogyne boundary generally B luckiLUCKY STRIKE CANYON SPECIES 400 SAME 0 contains relatively high species richness this 0 FAPApa ERFA GROUP A vegetation zone consists of floras from both 300 0 0 PSPRFSFR MESP EPNEepe 0 0 9 GROUP 8 larrea ambrosia and coleogyne shrublandsshrub lands 0 YUBR YSAHYSA a GROUP C ECPO 0 such floras include yucca schidigeraschidigera and kra- 200 YUSC iiilii111 ENVI ACSH 0 B a GROUP D 0 0a COCORARA meria parvifoliaparvifohaparvifolia from larrea ambrosia shrub X 0 1 KRPA QCI LYAN lands and Y bremfoliabrevifoliabrembrevifolia and grabiagrayia spinosa from 100 OPRA AMDUDU MATO GRSP coleogyne shrublandsshrublands species commonly asso- 0 OPEC 0 0 0a M YBAYUBA 0 latroLATRQ OPBAgp OPAC 0a ciated with coleogyne begin to establish as PO0 0 0 100 PSCO elevation increases along the ecotone A sig- EULA nificantnificant overlap of exists presumably 0 species 1 200 due to the relatively uniform habitat and topog- 200 loo100 0 loo100 2 00 3 00 400 500 raphy throughout the lower ecotone in lucky AXIS 1 strike canyon and on the 2 mountain ranges increasing diversity may partially relate fig 4 ordination of stand A and species 13 groups species determined by TWINSPAN along the lower coleogyne to increases in soil moisture and soil organic boundary in lucky strike canyon across the lower eco- matter and to decreases in soil compaction tone of coleogyne see appendix 1 for species abbrevia- and soil temperatures in lucky strike canyon tions lei and walker 1997 coleogyne gradually becomes less abundant as elevation increases and it disappears rapidly forbs and cryptogamic crusts in the coleogyne when artemisiaarthArtemista tntritrldentata appears harsh and zones moreover west 1983 stated that com- prolonged cold winter airali temperatures appear position and productivity of annuals in cole to play a major role in preventing the desert obyneogyne communities vary from year to year shrubs on lower mountain slopes from migrat- because they rely heavily on the amount of ing upslope bowns 1973 proposed that cold precipitation bromus rubens and B tectoriumtectorumtectorum air temperatures in winter months may be a are the primary winter ephemerals they tend factor limiting the distribution of coleogyne at to form carpetcarpetlikelike vegetation among shrubs in its upper elevational boundary the spring seasons during wet years although A negative correlation exists between the there is a zone of overlap between the 2 bro- abundance of grasses and shrubs in coleogyne mus species B rubens is more abundant at the vegetation zones grasses forbs and crypto- lower coleogyne ecotone while B tectoriumtectorumtectorum is gamic crusts are more common on deeper soils more common at the upper ecotone beatley where clay silt and mineral nutrients are more 1976 abundant and shrubs are more common on DECORANA analysis of axis I1 of stand or- shallow and sandy soils callison and brother- dinationdination reveals significant correlations between son 1985 for these reasons edaphic factors the distribution of sample stands and eleva- largely control the distribution of shrubs grasses tion and soil depth in the spring and sheep 199711997 COLEOGYNECOLEOCYNE GRADIENT ANALYSIS IN SOUTHERN NEVADA 161 mountain ranges extensive survey decreased and in structuring vegetation zones inm south- air and soil temperatures increased precipita- ern nevada relationships between the distri- tion and increased soil moisture are associated bution of sample stands and various physical with increases in elevation for the extensive factors are purely correlative experimental transects factors such as topography and per- functional and ecosystem approaches are cent soil cover show significant but weak cor- required to further understand relationships relations with stand ordination scores indicat- between distributions of plant communities and ing some influence of these parameters on the environmental factors in southern nevada final groupings of stand and species identified moreover extensive studies of these physical by TWINSPAN analysis soil depth is a signif- factors in coleogyne shrublandsshrublands across its entire icant physical factor limiting coleogyne distri- geographical range in the southwestern deserts bution coleogyne communities exhibit rela- are required to determine ecological require- tively shallow soil depth due to calichebaliche layers ments of coleogyne and the specific environ- 30 50 cm beneath the soil surface west 1983 ment it occupies the area 0 10 cm below the soil surface contains few coleogyne roots only roots of annuals acknowledgments were identified in this zone bowns and west 1976 shrubs can trap aeolian materials and we express our gratitude to yin chin lei greater root activity and weathering cause a de- and steven lei for collecting vegetation and pression of the petropetrocalciccalcic layer directly under environmental data anna sala assisted with shrubs west 1983 root biomass is primarily the multivariate analyses TWINSPAN and located between 10 and 30 cm and is nega- DECORANA we also thank the motor pool tively correlated with soil depth in coleogyne and department of biological sciences of the communities bowns and west 1976 soil depth university of nevada las vegas for providing is shallowistshallowestshallowest in coleogyne communities but logistical support deepest in pinus juniperus woodlands shallow soils result in low rootshootroot shoot ratio and limited literature CITED root development in the coleogyne communities west 1983 BEATLEY J C 1976 vascular plants of the nevada test site and central southern nevada ecological and DECORANA results of axis I1 of stand ordi- geographical distributions national technical infor- showed nation scores a significant correlation mation service USU S department of commerce between the distribution of sample plots and springfield VA elevation at the lower coleogyne ecotone of BOWNS J E 1973 an autecological study of blackbrushblackbrush lucky strike canyon DECORANA axis 2 did coleogyne ramosissima torr in southwestern utah not correlate significantly with any physical fac- unpublished dissertation utah state university logan tors except elevation habitat and topographtopographyy BOWNS J E AND N E WEST 1976 blackbrushBlackbrush coleoncoleog did not change significantly to enhance hetero- yne ramosissima torr on southern utah rangelands geneous environmental conditions throughout utah agricultural experiment station research the lower coleogyne elevational boundary report 27 department of range science utah state hence elevation is the most essential inde- university logan pendent variable to detect stand and species CALLISON J AND J D brotherson 1985 habitat rela- tiontionshipship of the blackbrushblackbrusbblaekblackbrush community coleogyne 1 groupings on axis I in lucky strike canyon ramosissima of southern utah great basin natural- and the spring and sheep mountain ranges in ist 45 321 326 southern nevada tables 1 2 HILL M 0 1979a TWINSPAN a fortran program for certain physical factors appear to play an arranging multivariate data in an ordered two way essential role in limiting coleogyne distribution table by classification of individuals and attributes ecology and systematics cornell university ithaca to a well defined elevational band between NY approximately 1050 and 2150 in in the spring 1979b DECORANA a fortran program for de and sheep mountain ranges near las vegas trendedtreaded correspondence analysis and reciprocal aver- nevada studying an elevational gradient anal- aging ecology and systematics cornell university ysis of the mojave desert plant communities ithaca NY HUNTER K L AND J R mcauliffe 1994 elevational would contribute to an understanding of whichofwhich shifts of coleogyne ramosissima in the mojave desert physical factors are most important in deter- during the little leeiceiee age quaternary research 42 mining the current distribution of coleogyne 216 221 162 GREAT BASIN naturalist volume 57

klnlkenrklinl M AND P COKER 1992 vegetation description and amelanchier utahensis koehne T AMUT analysis a piacticalpractical approach belhaven press lon- arctostaphylos pungentpungens HBK S ARPU don UKU K artemisia tntritrldentata gray S ARTR korthuisKoHn iuisluisluls S L 1988 coleogyne ramosissima in william atriplex canescentcanescenscanes cens pursh S ATCA C fischereischerelscherpischerFischeischerseher compiler the fire effects information atriplex conferticonfertifoliaconfertifohafolia wats S ATCO system database USU S department of agriculture ceanothus greggiigreggngregen gray S CEGR forestfoiest service intermountaininteiantei mountain research station fire Ceratceratoidesoides lanata pursh S CELA science laboratory missoula MT magnetic tape Cercocercocarpuscarpus ledifoliusledifohuslediledlfolius nutt T CELE leeisreelsreeis 9 track 1600 bpi ASCII with common LISP chilopsis linearislinearis caacav T CHLI present chrysothamnus nauseosus pall S CHNA LARSON P J 1977 the desert of the southwest sierra coleogyne ramosissima torr S CORA club books san francisco dowaniacowania mexicansmexicanamexicana torr S COME lelLEILLI S A AND L R WALKER 1995 composition and dis- dyssodia coopericooperia gray S DYCO tributiontrib ution of blackbrushblackofblackbrushbrush coleogyne ramosissima com- echinocactus polycephaluspolycephalous eng munimunitiesties in southern nevada pages 192 195 in bruce &abig&bigbig su ECPO A roundy E durant mcarthur jennifer S haley encelia virginensisvtrginensis blake S ENVIEN VI and david K mann compilers proceedings wild ephedra nevadensis wats S EPNE land shrub and and land restoration symposium ephedra vindis cov S EPVI 19 21 october 1993 las vegas NV general techni- eriodictyon angustifohumangustifolium nutt S ERAN cal report INT GTR 315 US department of agri- eriogonumfasciculatumenogonumfasciculatum benth S ERFA culture forest seiviceserviceSeivice intermountain research sta- fallugiaFallugia parparadoraparadoxaadoxa endl S FAPA tion ogden UT forsellesiaForsellesia nevadensis gray S FONE lelLEILLI S A AND L R WALKERWALKLR 1997 biotic and abiotic fac gaura coccineacoccinea nutt S GACO toitoltorss influencing the distribution of coleogyne com- crayiagrabiagrayia spinosa hook S GRSP munimunitiesties in southernn nevada great basin naturalist CutierrezcutlercutletcutierreziaGutiergutierreziagutierrezbareziaia sarothraesarothrae pursh Ss GUSA 5716357 163 171 hymenocleaHymenoclea salsola T & G S HYSA MAHALL B E AND R M CALLAWAY 1992 root commu- juniperusJumperus osteosperma torr T JUOS ninication mechanisms and intraintracommunityintianti community distribu- krameria parviparvifoliafolia benth S KRPA tions of two mojave deseitdesertdeceit shrubs ecology 73 larrea tntritrldentata cov S LATRLATE 2145 2151 lepidiumfremontiilepidium fremonfremontefremontntn wats S LEFR ROWLANDS P CG II11 JOIINSONJOHNSON E RITTER AND A ENDO lycium andersoandersoniiandersonnniinil gray S LYAN 1977 the mojave desert in M barbour and J machaeranthera tortifoliatortifohatortifolia gray S MATO major editors tenestnalterrestrial vegetation of california Menodora spinescentspinescensspinescens gray S MESP john wiley and sons new yoliyoriyork mirabilisfroebeliimirabihs froebfroebehiebiehi behr S MIFR wesrWESTWLST N E 1983 coloradocoloichloi ado plateau moravianmohavianMohavian blackbrushblackbrush Mormortoniamortomamontomatonia utahensis cov S MOUT semisemidesertdesert in temperate desert and semi deserts opuntia acanthocarpa engelm elsevier scientific publishing company amsterdam & bigel S OPAC netheinesheinetherlandslands opuntia basilarisbasilaris engelm & bigel S OPBA received 27 october 1995 opuntia echinocarpaechino carpa engelm accepted 13 january 1997 & bigel su OPEC opuntia polyacanthapoly acantha engelm & bigel su OPPO opuntia ramosissima engelm & bigel su OPRA APPENDIX I1 pinus monomonophyllaphylla torr & frem T PIMO prunusfasciculataprunus fasciculatafasciculatefasciculata wats S PRFAPRITA list of woody perennial species with abbreviations psilostrophe coopericooperia gray S PSCO found on 15 elevational transects from the larrea ambrosia psorothamnusfremontiipsorothamnus fremonfremontefremontntn S PSFR shrubshishl ublandsuplandsshrublandslands to the pinuspinns juniperus woodlands in the spring rhus tntritrllobata nutt S RHTR and sheep mountain langesranges and on the 6 elevations across Salazanalazaria mexicansmexicanamexicana torr S SAME the lower coleogtnecoleogyne boundary in lucky strike canyon salvia domi kell S SADO symbols of lifeformslifeilfeoflifeformsformsborms T tree S shrub Ss subshrub Stanstanleyastanleybleya pinnatepinnata pursh Ss STPI su succulent stephanomenastephanomeriaStephanomeriaomena pauciflora torr S STPA thamnosma montana torr species lifeformLifeform abbiabblabbreviationeviaevlaeviationaviationtion & frem S THMO yucca baccatabaccala torr S YUBA acacia greggngregen gray T ACGR greggii yucca brevifoliabrevifokabrevifoliafokaaoka engelm T YUBR acamptopappus shockleshockleyi gray S ACSH yi yucca schidigeraschidigera roezlroedl S YUSC agave utahensis gray S AGUT ambrosia dubosadumosa gray S AMDU ambrosia erioerloeriocentraenoenocentracentra glaycraygray S AMER great basin naturalist 572 0 1997 appp 163 171 BIOTIC AND ABIOTIC FACTORS influencing THE distribution OF COLEOGYNECOLEOGVNE communities IN SOUTHERN NEVADA

simon A leillellleli and lawrence R walkerlwalkerswalker1

ABSTRACT coleogyne ramosissimarainosissima is a desert shrub that forms a nearly monomonospecificspecific shrubland and occupies a well defined elevational band between approximately 1050 and 2150 m on mountain ranges in southern nevada the coleogyne shrubland shares relatively broad upper and lower ecotonesecotones with pinus jumpersjuniperusjumpems and larrea ambrosia plant communities respectively we characterized the extent that environmental factors correlate with coleogyne density and examined variation in biotic and abiotic factors along the lower elevational boundary of coleogyne in lucky strike canyon near las vegas nevada coleogyne density was positively correlated with gravimetric soil moisture soil organic matter coleogyne water potential coleogyne stem and leaf phosphorus and coleogyne leafleafbiomassbiomass however coleogyne density was negatively correlated with soil temperatures soil compaction and coleogyne stem and leaf nitrogen coleogyne stem biomass and elongation were generally negatively correlated with coleogyne density coleogyne density was weakly correlated with soil ph soil depth soil nitrogen soil and leafphosphorus these variables did not exhibit a consistent pattern with increasing elevation edaphic factors particularly soil moisture and soil organic matter appear to play a major role in determining the distribution of coleogyne shrubshrublandslands in southern nevada

key words coleogyne ramosissima soils lower coleogyne ecotone lucky strike canyon mojave desert

coleogyne ramosissima torr blackbrushblackbrush is silt and mineral nutrients are more abundant a member of the rosaceae family and is the callison and brotherson 1985 shrubs on the only species in the genus coleogyne coleogyne other hand are more common on shallow and primarily establishes along the upper and cen- sandy soils bowns 1973 suggested that low tral portions of the colorado river drainage soil moisture limits the lower elevational and is an endemic yet widespread species in boundary of coleogyne communities in utah the southwestern united states bowns 1973 hunter and mcauliffe 1994 proposed that an previous studies have shown that larrea tri increase in precipitation is the paramount fac- dentata ambrosia dubosadumosa plant communities tor permitting the downslope establishment of appear to dominate between 900 and 1200 in coleogyne in the vicinity of searchlight in elevation and are replaced by coleogyne com- southern nevada coleogyne establishes as low munitiesmunities from 1200 to 1500 inm elevation in the as 1080 in whereas around death valley of mojave desert beatley 1969 1976 coleogyne southern california coleogyne is absent below communities are replaced by finnsfinusrinusfenuspinus mononwnophyllamonophyllaphyliaphylla elevations of 1350 in hunter and mcauliffe 1994 juniperus osteosperma woodlands at elevations current geographic variation in precipitation appears to control the coleogyne distri- of 1500 1800 in coleogyne forms nearly mono bution at its lower elevational we specific stands in much of its range with rela- boundary examined abiotic factors that might limit cole tively broad upper and lower ecotonesecotones that ogyne its lower overlap adjacent obyne at elevational limit in southern communities by as much as nevada 100 in in elevation in southern nevada lei and walker 1995 coleogyne generally grows historical biobiogeographyBlO GEOGRAPHY well on sand sandy loam and loam less well on gravel and clay loam and poorly on dense pankratpackrat neotoma sppapp midden records from clay korthuis 1988 A negative correlation the state of nevada reveal that woodland vege- exists between the abundance of grasses and tation persisted to the end of the early holo- shrubs within the coleogyne shrublandsshrublands for in- cene at elevations as low as 1250 in in the stance grasses forbs and cryptogamic crusts mojave desert betancourt et al 1990 As are more common on deeper soils where clay annual temperatures rose pinus juniperus

idepartment of biological sciences university of nevada las vegas las vegas NV 89154400489154 4004

163 164 GREAT BASIN naturalist volume 57 trees migrated to higher elevations on desert the last pluvial period 20000 10000 yr BPBE mountain slopes and were replaced by desert bradley 1964 shrubs betancourt et al 1990 coleogyne first coleogyne shrubshrublandslands generally share rela- appears in the fossil record in 3 late wisconsin tively broad lower ecotonesecotones with larrea neotoma deposits in the frenchman flat area ambrosia in southern nevada lei and walker of southern nevada approximately 10000 yr 1995 lucky strike canyon represents the ago and should have extended then to about typical vegetation and landscape conditions 700 in wells and berger 1967 the entire prevailing in the region the aim of this study coleogyne zone and at least the upper part of was to determine the elevational range and the larrea ambrosia zone around frenchman coleogyne density at its lower ecotone this flat were occupied by evergreen juniperusJumperus study also involved an investigation of biotic woodlands during the late pleistocene wells and abiotic factors along the lower coleogyne and jorgensen 1964 the arrival of xerophytic elevational boundary in lucky strike canyon shrubs and the extinction of woodlands in low elevations in the mojave desert occurred about STUDY AREA 8000 yr ago and vegetation zones migrated upslope significantly around 1000 yr ago anand lucky strike canyon is located approximately 65 km northwest of vegas continue through the present time betancourt las 3610n between and kyle canyons on etetalal 1990 11510w lee the east facing slopes of the mountains coleogyne can be regarded as a paleoen spring in southern nevada coleogyne community demic species exhibiting little variability and the in lucky strike canyon is characterized by perhaps on its way to extinction bowns 1973 hot summers above 40 C and cold winters shrubs are considered relict because they 40c the below loc temperature means and extremes were probably once more widespread than at 10c are negatively correlated with elevation sum- present and their current distribution repre- mer rainfall usually occurs during thunder- sents a inm their range with time restriction storms in july and august and comes from the bowns 1973 paleopaleoecologicalecological evievl however evi- gulf of california drawn into the desert by dence reveals the coleogyne ecotone larrea strong convectional currents rowlands et al has undergone frequent elevational migrations 1977 summer storms are often local and in- during the quaternary period pendleton et al tense winter rains on the contrary are mild and from also 1995 evidence packratpankrat biddensmiddens widespread and come from the pacific ocean coleogyne has repeatedly moved up shows they may last up to several days snow is fre- and down elevational gradients in response to quent at high elevations in southern nevada van climatic shifts phillips and devender 1974 particularly at the upper coleogyne ecotone pendleton cole and webb 1985 et al 1995 with pinus juniperus woodlands A relative pendleton et al 1995 hypothesize that cole humidity of5ofofa 20 is common in summer sea- obyneogyne is currently migrating into areas where sons high evaporation and low precipitation it is adapted to edaphic and climatic conditions create a typical andaridanidannd environment with an coleogyne shrublandsshrublands appear to be migrating average annual rainfall of 200 mm toward higher elevations on desert mountain slopes in southern nevada and northward into METHODS southern parts of the great basin desert B pendleton personal communication 1994 the measurements were made in 1993 and ecology of coleogyne seedlings at higher ele- 1994 across the lower ecotone of coleogyne in vations of the mojave desert and min the great lucky strike canyon we established six 100 basin desert may have different dynamics than m2ma circular plots 565 m radius along a tran- lower elevation populations in the mojave sect in the center of lucky strike canyon at desert A global climatic shift toward lowered each of 6 elevations 1160 1310 in 292.9 km in annual precipitation and more extreme fluctua- distance at 30 m intervals for a total range of tions of temperature and rainfall began in the 150 in in elevation and a total of 36 plots tertiary and continued through the quaternary across the lower coleogyne ecotone the lowest to the present bowns 1973 the current com- of the 6 elevations represents larrea ambrosia position and distribution of plant communi- stands below the range of coleogyne shrubs ties in southern nevada has developed since the remaining elevations represent increasing 199719971 COLEOGWECOLEOGYNE distribution IN SOUTHERN NEVADA 165 levels of coleogyne density but the coleogyne chamber and nitrogen gas twenty terminal community extends to over 1600 m elevation twigs from 1 coleogyne bush approximately in lucky strike canyon lei and walker 45 60 cm canopy diameter were harvested in 1995 june 1994 for analysis of growth the growth soil samples at 2 depths 0 7 and 7 15 cm of coleogyne normally begins in march and were collected from random locations within ceases in june west 1983 twig elongation each of the 36 plots with core widths of 12 cm was measured from march to june 1993 stems all soils were passed through a 2 mm sieve and leaves of coleogyne were collected and before analysis soil and air temperatures and then oven dried for 72 h at 60c60goc C before weigh- soil moisture were measured every 6 wk from ing and analyzing for total nitrogen and phos- may to august 1993 soil moisture was also phorus following acid digestion page 1982 measured every 6 wk from may to august 1994 biomass of current year stems and leaves was we took soil temperature readings using a soil determined after oven drying for 72 h at 60c60goc C thermometer with the probe at the soil surface statistical analyses and at 5 10 and 15 cm below the surface in open areas and beneath shrub canocanopiespies air one way analysis of variance ANOVA fol- temperatures at 161.6iglg m were recorded concur- lowed by a tukey s multiple comparison test rently with soil temperatures we weighed soil analytical software 1994 was used to detect samples dried them for 72 h at 110c and then differences among elevations in the lower ele- reweighed them to determine gravimetric soil vatvationalional limits of coleogyne and to compare moisture oven dried soils were then used for site means when a significant elevation effect soil ph organic matter and nutrient analyses was detected pearsons correlation analysis soil organic matter and water potentials of was performed to correlate biotic and abiotic coleogyne shrubs were measured in may july factors with coleogyne density soil moisture and august 1994 soil organic matter was soil organic matter and soil and plant nutrient determined from ash remaining after soils percentages were arcsine transformed prior to were heated to 550c550 C for 4 h we measured soil statistical analysis multiple analysis of variance compaction in open areas in august 1994 using MANOVAMAN OVA was used to detect significant a penetrometer that was inserted into the soil effects of elevation depth month time and after we had removed stony surface pavements year on soil moisture soil organic matter soil total kjeldahl nitrogen TKN total kieiKielkjelkjeldahldahldahi temperature and water potential of coleogyne phosphorus TKP soil depth and soil ph mean values are expressed with standard errors were measured in august 1993 total nitrogen and significance was determined at P 0050.05oos and phosphorus were determined on oven dried soils following acid digestion using a RESULTS modified kjeldahl method page 1982 for rapid flow through analysis of soil extracts the mean density of coleogyne increased soil depth was estimated by determining the significantly P 000010.0001 as elevation increased depth to which a steel rod could be pounded across the lower ecotone fig 1 soil temper- into undisturbed soils soil ph was determined atures figs 2aaa ab2b declined significantly from a slurry consisting of equal parts of soil with increasing elevation across the lower eco- and distilled water no edaphic measurements tone by month and depth P 000010.0001 and were made during winter months due to the were cooler under shrubs than in the open P presence of snow on the ground at the upper 0050.05oos all possible interactions open space portions of the lower coleogyne ecotone shrub canopy soil depth month and eleva- in june 1993 within each of the 27 plots tion were also significant P 0010.01ooi temper- that contained coleogyne we measured the atures were usually cooler at higher elevations following characteristics of coleogyne water and under shrub canocanopiespies potential stem growth and nutrient status coleogyne density was negatively correlated TKN TKP and biomass production of stems with air temperatures r 0950.95ogs P 00010.001 and leaves we measured water potentials on across the lower boundary in lucky strike 10 to 20 cm long terminal branches of cole canyon air temperatures were also signifi- obyneogyne shrubs at predawn and midday in may cantly different P 000010.0001 among the 3 and august 1994 using a portable pressure summer months data not shown 166 GREAT BASIN naturalist volume 57

70 OPEN SPACE d

60 1 1 1 1 60 77 A p7pa77 0 CcmM 15 cm a a b c c 50 50 b 0 cd 7 40 40 bc uj 7 of A d zU P 30 f Lj 30 z a 20 0 ob 20 10 V a a 10 001i i 1100 1150 1200 1250 1300 13501550

ELEVATION m 0 1111000 1150 1200 1250 1300 1350 fig 1 density of coleogyne along its lower ecotone n ELEVATION m 6 mean s narrow vertical bars denote standard eierrorsroirol s of the means columns labeled with different letters aiealeare significantly different at P 0050.050oos05 UNDER SHRUB CANOPY

60 1 1 1 soil moisture table 2 0 7 cm soil depth B 0 cm 15 cm increased with elevation P 0 0001000010.0001 but also 50 varied by month may july and august P 000010.00010 0001 and year 1993 1994 P 0 0001000010.0001 a 40 significant for soil LU interactions moisture were of a ab bc c C c detected soil depth 0 7 and 7 15 2 between a a a a a cm and year P 00500.05oos05 soil depth and month 30 P 0.00010 0001 and between soil depth month HE 00001 L and elevation P 0.05000505 W 005 j 20 A similar pattern of soil moisture was seen V at depths of 7 15 cm as well as among the summer months and between the 1993 and 10 1994 years soil moisture percentage was the highest in the month of august 1993 data not 0 shown because of summer storms that occurred 1100 1150 1200 1250 1300 1350 from late july through mid august ELEVATION m coleogyne shrubs were significantly more water stressed more negative water potential fig 2 soil temperatures at the soil surface and at depths of 15 cm in open spaces A and under shrub values fig 3 in august of 1994 at lower ele- in canopiescano pies B in august 1993 n 36 in each treatment 0.0010 001 mid- vations at both predawn P 0001oolooi and mean sys5sa narrow vertical bars denote standard errors of day P 000010.00010 0001 water potentials were posi- the means columns labeled with different letters are sig- tively correlated with coleogyne density at nificnificantlyantly different at P 00500.05oos05 predawn and midday r 0950.950 95 P 00010.0010 ooi001 r 0970 97 P 00010.0010 ooi001 respectively significant interactions were detected between water potential the factor limiting growth and development of and month may and august 1994 P 0 0001000010.0001 coleogyne all possible interactions month ele- coleogyne shrubs experienced more water stress vation and time were also significant P 0050.05oos in may than in august although a similar pat- coleogyne density was positively correlated tern was detected between these months bowns with soil organic matter r 0930.93 P 00010.001 1973 proposed that shedding of leaves is an soil organic matter at 0 7 cm depth was sig- adaptive feature for conserving water which is nificantlynificantly greater as elevation increased table 199719971 COLEOCWECOLEOGYNE distribution IN SOUTHERN NEVADA 167

0 TABLE 1 pearsons correlation coefficients of coleogyneofcoleogyne 0 ramosissima density to various biotic and abiotic factors of 1 coleogyne all factors were measured during 1993 except soil compaction august 1994 nitrogen and phosphorus 2 values were determineddetel mined by kjeldahl digestion

zWLU 3 factor r F 0 soil organic matter 0-L 4 J 0W a a abhbchab bc c 0 7 cm 093 lilliitil111LLJ 7 15 cm 5 089 soil compaction 089 I31 1 c soil nitrogen UJ 6 I d Z 0 7 cm 037 a a b 7 15 cm 7 039 0 coleogyne nitrogen 0LU

4 1 stems J 8fi I I1 predawn midday 086 0 leaves 087 soil phosphorus 1100 1150 1200 1250 1300 1350 0 7 cmem 026ns 7 15 cm 0680681 ELEVATION m coleogyne phosphorus stems 099 fig 3 predawn and midday water potentials of coleoncoleog leaves 080 yne shrubs across the lower ecotone in august 1994 n coleogyne biomass 15 in each treatment mean s narrow vertical bars stem 092 denote standard errors of the means columns labeled leaves 050 with different letters are significantly different at P coleogyne stem elongation 052 005 soil depth 048 soil ph 0 7 cm 042ns0 42ns 7 15 cm 0 62ns062ns

2 P 000010.0001 A similar pattern was seen at P 0050oos05 depths of 7 15 cm data not shown there p001P ooi001 p 0001 was also a significant interaction P 005oos0.05 NS nonsignificant between elevation and soil depth coleogyne density was negatively correlated with soil compaction table 1 greatest soil stem nitrogen table 1 coleogyne leaf nitro- compaction occurred at larrea ambrosia stands gen content consistently higher than stem at lower elevations and least compaction oc- nitrogen did not differ significantly along the curred in nearly monomonospecificspecific coleogyne stands lower elevational limit of coleogyne table 3 at higher elevations table 2 P 000010.0001 P 0050.05 total soil kjeldahl nitrogen TKN at 2 depths total phosphorus content in coleogyne stems did not exhibit a consistent pattern with eleva- did not increase significantly table 3 P tion P 0050.05 and coleogyne density was 0050.05oos with elevation along the lower ecotone weakly positively correlated with total soil but coleogyne leaf phosphorus content did nitrogen at 0 7 cm or 7 15 cm table 1 the increase significantly table 3 P 0050.05 with upper 7 cm generally had higher TKN than elevation the soils 7 15 cm deep total soil phosphorus mean soil depth did not differ significantly TKP in the upper 7 cm of the soil did not with increasing elevation P 005oos0.05 but was vary with elevation P 0050.05oos and coleogyne weakly negatively correlated with the density density was not significantly correlated with of coleogyne table 1 although it declined total soil phosphorus table 1 the highest slightly with elevation the deepest soil was TKP values occurred at an elevation of 1280 found at an elevation of 1190 m the lowest m which was just below the nearly pure elevation at which coleogyne occurred coleogyne stands the lowest TKP values were coleogyne density showed a weak negative in larrea ambrosia stands 1160 m correlation with soil ph at depths of 0 7 cm total nitrogen content in coleogyne stems table 1 and did not vary with increasing ele- declined significantly table 3 P 005oos0.05 as vation P 0050.05 soils near the center of the elevation increased and coleogyne density ecotone 1210 m tended to have higher ph was negatively correlated with total coleogyne values than soils at other elevations 168 GREAT BASIN naturalist volume 57

TABLETABLL 2 soil moisture and soil organic matter at depths of 0 7 cm soil compaction kgcm2 and soil nutrients TKN and TKP at 0 7 and 7 15 cm depths along the lower coleogyne ecotone mean values of soil moisture organic matter and nutrients are expressed in percentages and weiewelewere arcsine transformed columns labeled with different letters are significantly differentdiffeidiffee ent at P 005oos0 05

soil TKN elevation soil moisture soil OM soil compaction soil TKP in kgcm2 0 7 7 15 0 7 7 15 1160 ogga 76a 007a 004a 082a0 82a 063a0 63a 1190 103ab 29a 65ab6 sab 008a0 08a 003a0 03a 071a0 71a 0 65ab065ab 1220 143abc143dbc 43ab 69ab6 gab 006a0 oga 003a0 03a 067a0 67a 0 73ab073ab 1250 170bc 45ab4 sab 58bc5 nbc8bc 006a0 oga 002a0 02a 074a0 74a 0 72ab072ab 1280 1 79bc179be 57ab5 aab7ab 64ab6 aab4ab 010a 006a0 oga 0ogga69a 100bloobI1 oobbob 1310 200c 68b socsoe5 oc 009a 005a0 osa 081a0 sia 0 82ab082ab

mean stem biomass of coleogyne tended maximum in early summer in southwestern significantlynonnonsignificantly FP 005520.0552 to decrease with utah bowns 1973 this pattern was more elevation table 3 and to be negatively corre- prominent in larrea ambrosia stands 1160 m lated table 1 with coleogyne density mean in elevation where coleogyne shrubs were leaf biomass of coleogyne table 3 showed no absent and apparently could not tolerate a statistical significance across the elevational combination of relatively high soil tempera- boundary but coleogyne density showed a tures and low soil moisture soil temperatures weak positive correlation with coleogyne leaf in open areas displayed a greater fluctuation biomass table 1 unlike stem biomass leaf and were consistently and significantly warmer biomass tended to increase as elevation in- than soil temperatures under shrub canopiescanopies creased up to 1300 m shreve 1924 suggested that insolation is of mean stem elongation of coleogyne 3873873.87 more importance than air temperature in deter- 0430.43 cm table 3 from march to june did not mining soil temperature little variation was display a definite pattern across the lower eco- observed between temperatures at 15 cm depth tone of coleogyne table 1 P 0050.05 the across the ecotone beneath canopiescanopies and in lowest mean stem elongation took place near open spaces much of the fluctuation that the center of the ecotone whereas the highest occurs during late spring and summer can be rate occurred in larrea ambrosia stands cole attributed to changes in soil moisture content obyneogyne density was not significantly correlated bowns 1973 bowns and west 1976 reported with stem elongation table 1 that soil temperatures in coleogyne communities start to increase in february and march discussion peak in june or july but decline in august due to wet soils soil temperatures in our study did abiotic factors appeared to limit the distri- not decline in august despite moderately wet bution of coleogyne at its lower elevational soils during the thunderstorm season soil mois- boundary in lucky strike canyon in southern ture and temperatures at the nevada test site nevada previous studies have shown that low in mercury in southern nevada do not always soil water content influences the distribution correspond to the lower temperature and in- of coleogyne at its lower ecotone in utah creased precipitation associated with increases bowns 1973 the appearance and persistence in elevation but they are influenced by phys- of coleogyne at its lower elevational limit in iographyio and soil properties rickard and searchlight in southern nevada is probably re- murdock 1963 lated to an increase in precipitation hunter and air temperatures in winter may influence mcauliffe 1994 precipitation is the major con- the distribution of larrea ambrosia at their trol of soil water content however no one has upper elevational boundary beatley 1974 tested which edaphic factor may be most impor- stated that the average extreme minimum air tant in determining the distribution and density temperatures on all larrea ambrosia sites are of coleogyne in southern nevada above 17c variation in air temperatures soil temperatures were negatively correlated along the lower boundary of coleogyne in this with coleogyne density soil moisture reaches study was 1c1ac C because the plots covered only a minimum while soil temperature reaches a 150 m in elevation nevertheless coleogyne 199719971 COLEOGYNECOLEOCYNE distribution IN SOUTHERN NEVADA 169

table 3 coleogyne stem elongation cm n 20 per bush total coleogyne stem and leaf nitrogen TKN and phosphorus TKP and coleogyne biomass g along the lower coleogyne ecotone n 27 mean values of nutrient status are expressed in percentages and were aresinearcsine transformed columns labeled with different letters are significantly different at FP 005oos0.050 05 elevation stem elongation stem leaves biomass g m cm TKN TKP TKN TKP stems leaves 1190 423a 0 58ab058ab 043a0 43a 086a0 86a 0ogga92a 018a0 18a 003a0 03a 1220 405a 060a 046a 087a 121a olsaoisa 004a 1250 357a 050ab 056a 076a0 76a 093a0 93a 014a0 14a 004a0 04a 1280 385a 040b0 40b ossa0 ssa 081a0 siasla 135ab 0015a15a 005a0 osa 1310 383a 041b0 41b 062a0 62a 075a0 75a 217b 0013a13a 003a0 03a densities were negatively correlated with air between elevations of 1220 and 1310 m beatley temperatures 1966 tends to form carpetcarpetlikelike vegetation among rickard and murdock 1963 stated that soils shrubs in the spring season of wet years it of coleogyne communities of yucca and french- was not abundant in larrea ambrosia stands man flats in south central nevada have more the presence of cryptogamic crusts can also available moisture than larrea ambrosia soils increase organic matter in soils loope and and that available moisture is greater in the gifford 1972 coleogyne zones in lucky strike subsurface 10 30 cm than in the surface canyon exhibited a higher organic content than 0 10 cm the lower limits of coleogyne dis- larrea ambrosia communities presumably be- tributiontribution may be a result of low soil moisture cause of abundant bromus ephemerals bowns and west 1976 soil moisture in our coleogyne density was generally negatively study was positively correlated with coleogyne correlated with soil compaction variations of density coleogyne growth ceased in mid june soil compaction may play a secondary role in presumably due to limited soil water content limiting the distribution of coleogyne at its soil moisture is greatest prior to the beginning lower elevational boundary soils at higher of coleogyne growth in mid march exhaustion elevations tend to have lower compaction pri- of soil moisture coincides with cessation of marily due to better infiltration also they are growth of coleogyne in mid june west 1983 more permeable to air and water required by in fact coleogyne underwent summer dor- plant roots bowns 1973 mancy with no growth around mid june no algae mosses and lichens that form crypto- water potential readings of coleogyne shrubs gamic crusts appear to fix nitrogen in coleoncoleog were recorded in july 1994 bowns and west yne communities loope and gifford 1972 and 1976 concluded that summer dormancy of may increase fertility of the associated soils coleogyne is a result of soil moisture depletion callison and brotherson 1985 total soil nitro- rather than high air temperatures and no chill- gen did not exhibit a definite pattern across ing treatment is required to break dormancy the lower coleogyne ecotone total phosphorus plants exhibiting drought induced summer remained fairly constant coleogyne leaves had dormancy have evolved to take advantage of a higher content of nitrogen and phosphorus water when it is present and to become dor- than stems which corresponds with bowns mant when water is scarce bowns 1973 1973 coleogyne leaves consistently stored soil organic matter increased with elevation more nutrients than stems coleogyne stems across the lower coleogyne boundary deposi- and leaves were relatively low in phosphorus tion of older and outermost coleogyne leaves nevertheless provenza 1978 proposed that at the onset of summer dormancy and decom- phosphorus content is highest during the grow- position of winter ephemerals can generate a ing period but appears to decrease as the cole considerable amount of organic matter for the obyneogyne growth period continues through june soil in coleogyne communities bowns 1973 coleogyne stem biomass g per branch de- west 1983 stated that composition and pro- clined as coleogyne became denser with ele- ductivity of annuals vary from year to year be- vation on the other hand coleogyne leaf bio- cause they rely heavily on precipitation bromus mass g per branch increased slightly as the rubens red brome grass a dominant winter species became denser with elevation this ephemeral of many larrea coleogyne ecotonesecotones phenomenon may indicate a production of more 170 GREAT BASIN naturalist volume 57

current season leaves as stem biomass declines plateau in southern utah pendleton et al 1995 with elevation seeds are generated only in wet years when the total rate of current season stem elon- rains arrive during the winter or during spring gation of coleogyne did not vary significantly growth bowns 1973 survival of seedlings during the growing season throughout the lower is exceedingly poor young seedlings usually elevational boundary nevertheless relatively die as a result of inadequate soil moisture and high air and soil temperatures may determine older seedlings are eaten and uprooted by the rate of stem elongation in coleogyne peri- rodents and rabbits bowns and west 1976 ods of warm weather result in abundant growth on the contrary seedling survival can be rela- while cool periods retard the growth rate tively high at high elevational sites of the col- bowns 1973 orado plateau in southwestern utah coleogyne density was weakly negatively coleogyne density and distribution appear correlated with soil depth across the lower to be influenced by edaphic factors particu- boundary at lucky strike canyon soil depth larly soil moisture and soil organic matter declined slightly from the first appearance of coleogyne also exhibits variations in stem elon- coleogyne to the nearly pure coleogyne stands gation as well as nutrient content N and P however callison and brotherson 1985 sug- and biomass production of stems and leaves gest that shallowness of soils is an important across its lower elevational boundary lucky feature of coleogyne communities and may par- strike canyon is representative of the vegeta- tially determine the abundance and distribution and landscape conditions prevailing in tion of coleogyne rooting patterns of coleoncoleog southern nevada experiments and establish- yne enable the plant to extract water more ment of longtermlong term plots at various sites are efficiently from shallow and sandy soils than necessary to further understand the complex from deep soils korthuis 1988 the majority relationships between the distribution of cole of coleogyne roots are found at depths of 10 30 obyneogyne and associated biotic and abiotic factors cm and balichecaliche layers are considered a major in southern nevada obstacle to many roots in southwestern utah bowns 1973 hence roots of many shrubs run acknowledgments horizontally and contribute to a general decrease in root biomass moving from plants we thank yin chin lei and steven lei for toward open areas between plants bowns collecting soil samples john boilingbolling assisted 1973 soil depth is a significant factor associ- with soil and plant nutrient analyses TKN ated with coleogyne density and distribution TKP helpful comments by richard hunter only when considering the entire mountain and burton pendleton greatly improved the range but not across its lower ecotone in manuscript the UNLV motor pool and depart- southern nevada lei 1995 ment of biological sciences provided logistical soil ph did not differ significantly between support and the UNLV graduate college pro- larrea ambrosia stands and the lower coleoncoleog vided partial financial support yne ecotone soil ph in coleogyne communities in utah ranges from 787.8 to 858.5 with no definite literature CITED trends within the profile and sites korthuis 1988 soil ph of lower coleogyne limits ranged analytical SOFTWARE 1994 statistics 414.14 1 an interactive statistical program for microcomputers analytical 78 80 only from 787.8 to 808.0 high ph values are re- software st paul MN lated to elevated levels of calcium and sodium BARBOUR M G 1969 age and space distribution of the in soils callison and brotherson 1985 hence desert shrub larrea divaridivancatadivaricatadivaricatedivancatacata ecology 50 679 685 soil ph did not play a vital role in limiting BARBOUR M G AND J MAJOR 1977 terrestrial vegeta- coleogyne distribution at lucky strike canyon tion of california john wiley and sons new york BEATLEY C 1966 ecological status of introduced brome of coleogyne seedlings J the scarcity at its grasses bromus sppapp in desert vegetation of south- lower elevational boundary of lucky strike can- ern nevada ecology 47 548 554 yon is indicative of infrequent reproduction 1969 biomass of desert winter annual plant pop- lei personal observation 1994 however we ulationsulations in southern nevada oikosbikos 20 261 273 observed greater seedling densities at higher 1974 effects of rainfall and temperature on the distribution and behavior of larrea divaridivancatadivaricatadivaricatedivancatacata cre- elevations survival seedling can be relatively osote bush in the mojave desert of nevada ecol- high at high elevational sites of the colorado ogy 55 245 261 199719971 COLEOGYNECOLEOCYNE distribution IN SOUTHERN NEVADA 171

1976 vascular plants of the nevada test site and USU S department ofagriculture forest service inter- central southern nevada ecologic and geographic mountain research station ogden UT distributions national technical information service LOOPE W L AND G FE GIFFORD 1972 influence of a soil USU S department of commerce springfield VA micromicrofloralfloral crust on select properties of soils under betancourt J L T R VAN DEVENDER AND P S MARTIN pinyon juniper in southeastern utah journal of soil 1990 packratpankrat biddensmiddens the last 40000 years ofbioticof biotic and water conservation 27 164 167 change university of arizona press tucson PAGE A E 1982 methods of soil analysis part 2 ameri- BOWNS J 1973 an autecological study blackbrushofofblackbrushblaekblackbrush cole can society of agronomy madison WI obyneogyne ramosissima torr in southern utah unpub- PENDLETON B K S E MEYER AND R L PENDLETON lished doctoral dissertation utah state university 1995 blackbrushBlackbrush biology insights after three years logan of a longtermlong term study pages 223 224 in bruce A BOWNS J E AND N E WEST 1976 blackbrushBlackbrush coleoncoleog roundy E durant mcarthur jennifer S haley and yne ramosissima torr on southern utah rangelands david K mann compilers proceedings wildlandwildwindlandland utah agricultural experimental station research shrub and and land restoration symposium 19 21 report 27 utah state university logan october 1993 las vegas NV general technical BRADLEY W G 1964 the vegetation of the desert game report INT GTR 315 USU S department of agricul- range with special reference to the desert bighorn ture forest service intermountain research station transactions of the desert bighorn council 8 43 67 ogden UT PHILLIPS A M AND T R VAN DEVENDER 1974 pleisto- CALLISON J AND J D brotherson 1985 habitat relation- packratpankrat biddensmiddens from the lower grand canyon ship of the blackbrushblaceblackblackbibrushush community coleogyne ramo cene of arizona journal of the academy of science sissimamissimasis sima of southern utah great basin naturalist 45 arizona 321 326 91179 117 119 PROVENZA F D 1978 getting the most out blackbrusbofblackbrushof COLE K L AND R H WEBB 1985 late holocene vege- utah science 39 144 146 tation changes in Greengreenwaterwater valley mojave desert RICKARD W H AND J R MURDOCK 1963 soil moisture california quaternary research 23 227 235 and temperature survey of a desert vegetation mosaic HUNTER K L AND J R 1994 elevational mcauliffe ecology 44 821 824 shifts of coleogyne ramosissima in the mojave desertdeseitdeceit ROWLANDS P G H JOHNSON E RITTER AND A ENDO during the little ice age quaternary research 42 1977 mojave desert in M barbour and 216 221 the J major editors terrestrial vegetation of california KORTHUIS S L 1988 coleogyne ramosissima in william john wiley and sons new york C Fischerscheiseher compiler fire effects information fischer the sys- SHREVE FE 1924 soil temperature as influenced by alti- tem database U S of US department agriculture for- tude and slope exposure ecology 5 128 136 est service intermountain research station inter- WELLS P V AND R BERGERBERGEB 1967 late pleistocene history mountain missoula MT fire science lab magnetic of the coniferous woodland in the mojave desert tape reels 9 track 1600 bpi ASCII with common science 155 1640 1647 LISP present WELLS P V AND C D JORGENSEN 1964 pleistocene LEI S A A analysis 1995 gradient of coleogyne comminicommunicommuni- woodratwoodral biddensmiddens and climatic change in the mojave ties in southern nevada unpublished mastelmaster s thesis desert a record of juniper woodlands science 143 university of nevada las vegas 117 appp 1171 1173 LEI S A AND L R WALKER 1995 composition and distri- WEST N E 1983 colorado plateau moravianMomohavianhavian blackblackbrushblackbrusbbrush bution of coleogyne communities inm southern nevada semisemidesertdesert pages 399 411 in temperate deserts and pages 192 195 in bruce A roundy E durant semi deserts elsevier scientific publishing company mcarthur jennifer S haley david K mann com- amsterdam netherlands pilers proceedings wildlandwildwindlandland shrub and and land restoration symposium 19 21 october 1993 las received 27 october 1995 vegas NV general technical report INT GTR 315 accepted 27 january27january 1997 creatgleatgreat basin naturalist 572 C 1997 appp 172 177

VARIATION IN germination RESPONSE TO temperature AND WATER availability IN blackbrushBLACKBRUSH COLEOGYNE ramosissima AND ITS ecological significance

simon A leillellleli

absausABSTRACT1 ragl blackbrushblaekBlackblackbibrushush coleogyne ramosissima tontorr is a dominant desert shrub in a distinct mid elevational vegetation belt between creosote bush barsagebursage larrea ttltntridentatadentato ambrosia dubosadumosa shrubland below and big sagebrush pinyon pine utah juniper artemisia tntritrldentata pinus monomonophyllaphyliaphylla juniperus osteospemiaosteosperma woodland above in the mojave desert seed germination patterns of blackbrushblackbrush seeds collected from 2 elevations 1200 and 1550 m in 5 mountain langesranges within the blackbrushblackbrush shrublandsshrub lands were investigated morphological features of blackbrushblackbrush seeds including weight length and width were not significantly different P 0050.05oos0 05 among elevations and mountain ranges in the mojave desert germination of blackbrushblackbrush seeds was optimal when preceded by a prechill period of 4 6 wk seeds incubated at loomroom temperature germinated poorly seeds collected at warm low elevation sites appeared to be less dormant required less prechill time germinatedgeiger meatedmmated faster and showed a higher overall germination response at low temperature relative to cold high elevation sites frequencies of watering also determined the germination response watering at 2 wk intervals revealed the greatest germination some ecotypic variation among populations establishing at different elevations was evident with legardregard to dormancy duration and germination response at certain constant temperatures

key words blackblackbrushbrush germination elevation prechill temperature wateringwateringfrequencyfrequency ecotype mojave desert

control and timing of the germination pro- moist chilling at 4cac without light bowns cess are the keys in survival of plant populations 1973 bowns and west 1976 this phenome- whose principal mode of reproduction is from non could be advantageous because seeds can seed harper 1977 meyer et al 1989 the germinate from relatively deep in soils that selection pressure operating at this phase of rodents leave behind in the cache bowns and the life cycle must be strong since only a tiny west 1976 seeds collected from low eleva fraction of seeds survive to maturity meyer et al tion sites are less dormant than seeds from 1989 investigations between population dif- high elevation sites in southern utah and ferencesferences in germination strategy within black nevada pendleton et al 1995 the correla- brush coleogyne ramosissima are infrequent tion between dormancy status and elevation of by holding aspects of life history and genetics collection site may indicate that blackbrushblackbrush background more or less constant germination has evolved ecoecotypestypes at the germination level strategy variation of blackbrushblackbrush in relation to pendleton et al 1995 temperature and moisture availability may be the aim of this study was to discover the more evident existence of possible geographical and eleva- blackbrushBlackbrush was selected as a representative tional ecoecotypestypes within blackbrushblackbrush populations species for such a study because little atten- that represent specific physiological adapta- tion has been focused on its germination biol- tions to micromicroenvironmentsenvironments in the various ogy previous studies have been limited to rel- mountain ranges across the mojave desert atively small geographical areas in southern seeds of 8 isolated populations of blackbrushblackbrush utah and nevada A significant relationship from 5 major mountain ranges with different exists between collection site elevation and seed elevations were collected to examine variation germination response at 5 15c15 C with and within germination response to different tempera- out a short 2 wk chill pendleton and meyer ture conditions and watering frequencies seed 1994 pendleton et al 1995 germination of morphology among the 8 populations was also blackbrushblackbrush seeds requires a cold stratification measured

1 department of biological sciences university of nevada las vegas laslasvegaslosvegasvegas nevada 89154400489154 4004

172 199719971 blackbrushBLACKBRUSH COLEOGWECOLEOGYNE SEED germination 173

METHODS TABLE 1 location ofblackbiushofblackbrusb seed collections on 8 major mountain ranges ai rangedarranged alphabetically in southern seed collection sites utah nevada and northwestern arizona applappioximateapproximateAppi oximate elevation of each collection site is shown four blackblaekblackbrushblackbrnshbrush blackbrushBlackbrush seeds from 8 populations were populations were sampled within the spring mountains collected by dr burton pendleton and his col- collections were made at the full establishment of black in the leagues in the mojave desert table 1 to deter- brush shrublandsshrublands and were utilized in temperature and moisture experiments mine germination strategies and requirements seeds of 4 populations were collected within elevation of collection the mountains table 1 blackbrushBlackbrush location site spring m often forms a well defined mid elevational band between creosote bush bursagebarsage below mccullough range NV 1200 mormon range 1550 and big sagebrush utah juniper UT pinyon pine sheep range NV 1550 woodland above elevations of blackbrushblackbrush spring mountains NV shrublandsshrublands ranged from 1000 to 1850 inm above kyle canyon 1550 sea level the vegetation of this community lee canyon 1550 mt potosi 1200 was dominated by a closely spaced matrix of red rock canyon 1200 low blackblackbrushbrush with a scattered distribution of virgin mountains AZ 1200 other woody taxa bowns 1973 seeds were obtained from dr pendleton and his colleagues of the shrub sciences laboratory in provo utah they collected at 2 eleva- dishes approximately 25 seeds were placed tions 1200 and 1550 in on 5 mountain ranges on each culture with 30 replicates in each of in late june through july 1994 to represent an the 6 major experimental conditions seeds elevational gradient covering a relatively small were incubated at room temperature 24c24 C geographical area major mountain ranges and as well as in a cool chamber ac4c and 14c14 C collection sites were selected on the basis of for 8 wk without light but were briefly exposed full establishment of blackbrushblackbrush shrubshrublandslands to soft white fluorescent light as germinated and seed availability respectively flowering seeds were counted and recorded seeds from of blackbrushblackbrush generally occurred in may and 8 populations were subjected to 3 different relied heavily on winter and spring precipita- water frequencies every 3 and 2 wk and once tion anthesis occurring over a period of 1 2 a week with an amount of 5 ml and to 4 dry wk was not synchronous throughout the ele- prechill periods no chill prechill at 2 4 and vations of blackbrushblackbrush shrublandsshrub lands pendleton 6 wk the germination of each population was 1994 seeds at higher elevations generally had observed within 2 wk of the initial experiments a shorter flowering period and were freshly germination percentages were recorded at collected in late june whereas seeds at lower weekly intervals for 8 consecutive weeks and elevations had a longer flowering period and mean germination values were arcsine trans were freshly collected in july 1994 pendleton formed emergence of radiclesradiradichescles was the criterion personal communication 1994 for germination morphological features such as seed weight and dimensions were also used to germination experiments discover variations among the 8 isolated black approximately 9600 blackbrushblackbrush seeds were brush populations across the mojave desert initially utilized 3 different temperatures in statistical analysis and 3 different watering frequencies with all possible combinations of these 2 treatments one way analysis ofvarianceof variance ANOVAAN OVA fol- under laboratory conditions some seeds were lowed by a tukey s multiple comparison test incubated at 4cac WC14 C and room tempera- analytical software 1994 was used to detect ture 24c24 C without dry prechill while others differences among seed morphological traits were preceded by dry chilling treatments at from 8 isolated populations and to compare 2cac for 2 4 and 6 wk prior to initiating ger- means of morphological traits from the 2 ele- minationmination experiments in a laboratory at the vations and several geographical locations UNLV campus during experimental treat- multiple analysis of variance MANOVA was ments blackbrushblackbrush seeds were covered by 2 employed to detect significant effects of eleva- layers of moist filter papers in closed petri tion temperature and watering frequency on 174 GREAT BASIN naturalist volume 57

tableTABUTAHLL 2 mean weight length and width with standard errors of blackbrushblackbrush seeds collected from 8 mountain ranges in southernsouthein utah nevada and northwestern arizona N 100 per population in each measurement mountain ranges aiealeare arranged alphabetically and 4 blackbrushblackbrush populations were from the spring mountains mean values followed by the same lettelletterietter within columns aiealeare not significantly different at P 0050oos0.0505 mountain ranges followed by asterisks indicate high elevation sites length mm width mm weight population mg mean ST mean sxax mccullough range 22 513a 017 368a oilolioii0110.11 mormon range 24 sisa 012 383a 012 sheep range 23 siga 017 379a 016 spring mountains NV kyle canyon 24 sita 019 382a 015 lee canyon 23 514a 016 375a 013 mt potosi 24 513a 016 369a oilolioii011 red rock canyon 22 502a 015 365a 014 virgin mountains 23 515a 018 378a 015

seed germination rate mean values of seed di- temperature on the contrary began to experi- mensionsmensions were presented with standard errors ence fungal growth approximately 2 wk from and significance was determined at P 0050.05 the initial experiments with an exception of the 6 wk dry prechill period table 5 the fungi RESULTS AND discussion became significantly more evident throughout the cultures after 4 wk and seed viability was seeds were collected from 8 isolated popu- severely damaged no seeds survived beyond lations in southern utah nevada and extreme 6 wk after the initial experimental treatments northwestern arizona table 1 morphological table 5 characteristics of blackbrushblackbrush seeds were not cold stratified seeds experiencing no dry significantly different FP 005oos0.05 table 2 in prechill yielded fairly low germination per- terms of weight length and width among the centagescentages from both elevations although seeds 8 populations across the mojave desert table from lower elevational sites tended to have 2 germination experiments under laboratory higher percentages table 3 collections made conditions showed various responses to 3 dif- in different mountain ranges across the mojave ferent temperature conditions and 3 watering desert from the same elevations behaved sim- frequencies all possible interactions elevation ilarly in germination pattern and response temperature and watering frequency were sig- tables 3 5 however seeds collected from nificantnificant P ooiool0010.01 on germination rate germi- warmer lower elevational sites 1200 m had a nation of blackbrushblackbrush seeds was significantly higher germination response after a chill as greater P 0050.05 incubating at 4cac cold short as 2 wk than those seeds collected from moist stratification than incubating at 14c14 C colder higher elevational sites 1550 m at a and 24c24 C without dry prechill and preceded constant low temperature my results support by 3 chill periods table 3 seeds from all the suggestion by meyer and pendleton 1990 populations responded well to cold stratifica- that germination patterns vary as a function of tion especially when they were preceded by climate and elevation within a species seeds chill treatments of 4 and 6 wk cold stratifica- also responded to a long chill but often yielded tion at 44cac C without light is required for black a lower germination percentage brush seeds to break dormancy and germinate seeds collected from colder higher eleva- bowns 1973 bowns and west 1976 radicle tional sites responded poorly to short chill but protrusion was from the narrow bent portion responded well to both intermediate 4 wk and of a seed bowns 1973 A cold stratification long 6 wk chill periods at 3 constant temper- requirement may function as an effective mech- atures tables 3 5 long chill was required anism to prevent germination at a season when for complete removal of dormancy for seeds prevailing conditions are unfavorable for seed- found at high elevations meyer and pendle- ling survival capon et al 1978 seeds from ton 1990 seeds may break dormancy in early all populations that were incubated at room spring after experiencing winter chill under 199719971 blackbrushBLACKBRUSH COLEOCWECOLEOGYNE SEED germination 175

TABLE 3 mean germination response ofblackbrushofblackbrushblackblaekbrush seeds at 4cac without light and dry prechill as well as preceded by 2 4 and 6 wk of dry chilling from 8 isolated populations seeds were subjected to 3 different watering frequencies at I1 2 or 3 wk intervals all mean germination values are expressed in percentages and were arcsine transformed N 1000 per population per treatment mountain langesranges followed by asterisks indicate high elevation sites row values followed by different letters aieateare significantly different at FP 0050.050oos05

wateringwatering frequencyfrequc ncy per treatmenttreatmeimelit

c population noN 0 prechill11 2S wk chillchiehiehl 11 4 wk chillchi 11 gwi6t wk chillchiehiehl 11

1 2 3 1 2 3 1 2 3 1 2 3 mccullough iselse 29d lseise 29d 37c 34cd gob 94ab 92b 97a 98a 97a mormon loeioe 12e loeioe 20d 40c 39c 82b glab 86b 94a 96a gsa sheep 14f 18e 12f 23d 34c 31c 85b 93ab 87b 96a 97a 94ab kyle canyon af8f ige lof 22d 34c 30cd 85b 92ab 89b 91ab 96a 97a lee canyon 109log 14f 13f 21e 32d 28de sic 92ab 88b gob gsa 93ab mt potosi 19f 25e 19f 39d 48c 34d 86b 89b 88b 95ab 99a 96ab red rock 22f 27e 199 35d sse 34d 93b 97a 94ab 95ab 97a 98a virgin 19f 18f ilg 32e 47c 41d gib 93ab 93ab 94a gsa 96a

TABLE 4 mean germination response blackbrushofblackbrushof blackblaekbrush seeds at 14c14 C without light and dry prechill as well as preceded by 2 4 and 6 wk of dry chilling from 8 isolated populations seeds were subjected to 3 different watering frequencies at 1 2 or 3 wk intervals all mean germination values are expressed in percentages and were arcsine transformed N 1000 per population per treatment mountain ranges followed by asterisks indicate high elevation sites row values followed by different letters are significantly different at P 00500.05oos05

wateringwatering frequencyfrequebrequemey per treatmenttreatme nt

rilrii 11 11 i 11 population noN0 prechprechillill 2i wk chillchiehiehl 41 wk chillchiehiehl 6gwiwk chiehiehlchill

1 2 3 1 2 3 1 2 3 1 2 3 mccullough lgig lgig og 19ef 24e 17f 62d 72c 69c gib 97a 94ab mormon lgig 4gag og 12f 22e 15f god 67c 64cd 86b 94a 91ab sheep range of if of 13e ige 13e 56d 65c 59cd 87b 93a 90ab kyle canyon af2f af2f if 13e 21d 17de 64c 74b 70bc 88a 89a goa lee canyon isig lg isig 14f 21e 18ef 64d toctoe 67cd 89ab 93a 88b mt potosi 3gag 3gag 3gag 21f 26e 22ef 67d 76c 71cd 88ab 92a gia red rock af3f af3f af3f 18e 25d 22de 71c 83b 77c 84b 94a 93a virgin 3gag 2gag 2gag 19f 27e 23ef 73d 79c 77cd 89b gsa glagia

TABLE 5 mean germination response blackbrushofblackbrushof blackblaekbrush seeds at 24c24 C without light and dry prechill as well as preceded by 2 4 and 6 wk of dry chillogchillmgcbilling from 8 isolated populations seeds were subjected to 3 different watering frequencies at 1 2 or 3 wk intervals all mean germination values are expressed in percentages and were arcsmearcsinearceme transformed N 1000 per population per treatment mountain ranges followed by asterisks indicate high elevation sites row values followed by different letters are significantly different at P 005oos0 05

wateringwatering frequencyfredifreqilencyleney per treatmenttreatmerit

H 1 population noN 0 prechprechillilllii 2 wk chilehllehlichill 4 wk chill 6i wk chilchiichill

1 2 3 1 2 3 1 2 3 1 2 3 mccullough oe oe oe oe oe oe 12d 16d 21c21e 78b 84ab 89a mormon oe oe oe ie iele oe 15d 23c 20c ssa 78b 8 labiab81ab sheep oe oe oeoc oe oe of 13e 18d 22c22e 86a 81b 80b kyle canyon of of of of of if ige 21d ite 79b tse 86a lee canyon of if of of of of 12e 18d 14e 70c 83a 76b mt potosi oe oe oe ie oe oe 14d 16cd iselse18c 84b 85b goa red rock ie1eae ie oe 2eae iele oe 19d 24c24e 25e25c 84ab 89a 78b virgin oe oe oe iele oe oe 13d 22c22e giegic 78b 87a 85a 176 GREAT BASIN naturalist volume 57 natural field conditions meyer and pendleton timing strategies in response to climate meyer 1990 although the results between 4 and 6 and pendleton 1990 wk periods of cold stratified seeds were not results suggest some ecotypic variation significant a slight variation in germination re- among seed collections at different elevations sponse is discernible table 3 seed collections in the mojave desert the idea of blackbrushblackbrush from cold sites showed a general pattern of being a paleoendemicpaleoendemic species showing little slower initial germination after a short duration genetic variability and perhaps on the way to of chill and often yielded lower overall germi- extinction bowns 1973 was not completely nation percentages after 8 wk relative to seed supported blackbrushBlackbrush populations are relictualrelicrevictualtual collections from warm sites during the cold because they were probably once more wide- stratification experiments tables 3 5 spread and their current distribution repre- blackbrushBlackbrush seeds usually germinated best sents a restriction in their range with time when water was applied at 2 wk intervals bowns 1973 nevertheless blackbrushblackbrush exhib- regardless of elevational sites conversely ited a variation in the germination rate at dif- watering at I1 and 3 wk intervals regardless of ferent elevations from a relatively small geo- temperature conditions generally showed a graphical area in southern utah and nevada significant reduction FP 0050.050oos05 in germination pendleton et al 1995 this differentiation in percentages watering at 3 wk intervals reduced germination response was also discovered the germination rate on the cultures A certain among the isolated populations in mountain amount of watering must occur before seeds ranges across the mojave desert according to will break dormancy perhaps to remove chem- my results the upper and lower elevational ical inhibitors walton 1969 if only small ecoecotypestypes were detected in terms of dormancy amounts of chemicals are leachedbeached away by duration and seed germination response at cer- insufficient moisture the seed replenishes the tain constant temperatures differences in ger- lost chemicals and the dormancy period con- mimination rate would not result in a significant tinues walton 1969 this phenomenon may species difference or a difference in the stand apply to seeds that are supplied with low evidence of packratpankrat biddensmiddens during the quantities and frequencies of water despite quaternary has revealed that the creosote the presence of fungi the overall effect of fungi bush blackbrushblackbrush ecotone has undergone fre- on germination was significantly reduced when quent migrations moving up and down in ele- seeds were incubated at a constant low tem- vation in response to climatic shifts phillips peraperatureture compared to a constant room temper- and van devender 1974 cole and webb 1985 ature hence not only was cold stratification spaulding 1990 pendleton et al 1995 the essential for seed germination but the fre- changing of gene pool to adapt to new ranges quency and amount of watering were also vital with regard to climatic shifts has led to repeated to promote a relatively high germination per- and successful migrations of blackbrushblackbrush along centage elevational andor environmental gradients dur- variation min germination response to 3 con- ing the quaternary period pendleton et al stant low temperatures among seed collections 1995 for these reasons blackbrushblackbrush may not be at 2 elevations was detected at P 0050.050oos05 seeds regarded as a paleoendemicpaleo endemic species however from all elevations responded positively to a 2 my germination studies covered only a rela- wk chill but the response of seeds from low tively small geographical range in the mojave elevation sites was roughly twice that of seeds desert experiments and longtermlong term investiga- collected at high elevation sites pendleton et al tions with various life stages of blackbrushblackbrush are 1995 which generally corresponds with my necessary to further discover the ecotypic and data the general germination pattern is simisimlsimi- or genetic variability of blackbrushblackbrush across the lar to patterns for many other desert shrubs entire southwestern desert region growing at high elevations in the intermountain area meyer and pendleton 1990 variation acknowledgments in populations within species may be related to variation in climatic conditions at the seed seed collection and helpful comments pro- collection sites meyer and pendleton 1990 vided by dr burton pendleton of the shrub this phenomenon suggests a relatively strong sciences laboratory in provo utah are grate- selection pressure for adaptive germination fully acknowledged comments by L R walker 199719971 blackbrushBLACK BRUSH COLEOGYNECOLEOCYNE SEED germination 177 greatly improved the manuscript logistical asteraceae and its ecological implications ameri- support of the department of biological sci- can journal of botany 76 981 991 MEYER S E AND R L PENDLETON 1990 seed ences of the university of nevada vegas germina- las tion biology of spskinlessspmlessspinlessinless hopsagehousage between population is also greatly appreciated differences in dormancy and response to temperature paper presented at the symposium on cheat literature CITED grass invasion shrub die off and other aspects of shrub biology and management las vegas NV 5 7 april 1989 analytical SOFTWARE 1994 statisticstatistixStati stix 4144.11 an interactive PENDLETON B K AND S MEYER germination statistical program for microcomputers analytical E 1994 software tallahassee FL biology of coleogyne ramosissima rosaceae I1 studies BOWNS J E 1973 an autecological study of blackbrushblackbrush laboratory unpublished manuscript on file coleogyne in southwestern at shrub sciences laboratoryLaboi atory 735 north 500 east ramosissima torr in utah provo unpublished dissertation utah state university UT 84606 logan PENDLETON B K S E MEYER AND R L PENDLETON 1995 blackbrushBlackbrush biology insights after BOWNS J E AND N E WEST 1976 blackbrushBlackbrush coleoncoleog three years of long study yne ramosissima torr on southern utah rangelands a longtermterm in bruce A roundy E durant S mann department of range science utah state university mcarthur jennifer haley and david K utah agricultural experimental station research compilers proceedings wildlandwildwindlandland shrub and and land report 27 restoration symposium 19 21 october 1993 las vegas NV U S CAPON B G MAXWELL AND P SMITH 1978 germination US department of agriculture forest responses to temperature pretreatment of seeds from service intermountain research station ogden UT general technical report GTR 315 ten populations of salvia columbanaecolumbariaecolumbanahanaeariae in the san INT gabriel mountains and mojave desert california PHILLIPS A M AND T R VAN DEVENDER 1974 pleisto packratpankrat grand canyon alisoahso9ahsop9 365 373 cene biddensmiddens from the lower of arizona journal of the nevada academy COLE K L AND R H WEBB 1985 late holocene vege- arizona of 9 117 119 tation changes in GreenGrcengreenwatergrcenwaterwater valley mojave desert science california quaternary research 23 227 235 ROWLAND P G H JOHNSON E RITTER AND A ENDO 1977 M HARPER J 1977 the population biology of plants acade- the mojave desert in barbour and editors micmie press new york J major terrestrial vegetation of california and sons new york LEI S A 1994 plants of the north american deserts john wiley unpublished research report university of nevada SPAULDING W G 1990 vegetation dynamics during the las vegas last deglaciation southeastern great basin USA quaternary research 33 118 203 1995 A gradient analysis of coleogyne comminicommunicommuni- TURNER R M 1982 of the ties in southern nevada unpublished master s the- biotic communities american southwest states and mexico geological sis university of nevada las vegas united survey USU S odtheof LEI S A AND L R WALKER 1997 classification and department ofthethe interior WALTON K 1969 and zones aldinealdmealame publishing com- ordination of coleogyne communities in southern the nevada great basin naturalist 57 155 162 pany chicago IL MEYER S E E D MCARTHUR AND G JORGENSEN 1989 variation in germination response to temperature received 4 june 1996 in rubber rabbitbrushrabbitbrush chrysothamnus nauseosus accepted 29 january 1997 great basin naturalist 572 C 1997 appp 178 183 diplostomiasis IN NATIVE AND introduced FISHES FROM yellowstone LAKE WYOMING

victor H inchaustylinchausty1 michael foutzlfoutz1 richard A Heckmannheckmannl2heckmann1212 claudeteclaudeceClaudete ruas3 and paulo ruas3

ABSABSIRACtraerTRAur1 totals of 101 native yellowstone cutthroat oncorhynchus clarki louvienboubouvienbouvierivienvieri 27 introduced lake trout salvelinus namaycush and 40 introduced longnoselongnose sucker catostomus catostomus from yellowstone lake wyoming USA weiewelewere examined for eye flukes metacercariaemetacercanae of the trematode fluke diplostomum were in vitreous humor andor lens of 94 of yellowstoneyellows tone cutthroat trout 92 of lake trout and 78 of longnoselongnose sucker longnoseLongnose sucker had 7 prevalencepi evalenceevalenee of infection in both lens and vitreous humor of metacercariaemetacercanae while yellowstone cutthroat trout had 3 and lake trout 8 diplostomum spathaceum was in lens tissue of 5 of infected yellowstone cutthroat trout and 93 of longnoselongnose suekersuckersuckel diplostomum baenbaeri was in vitreous humor of 92 each of infected yellowstone cutthroat trout and lake troutti out morphological characteristics indicate that a single species infected the lens of yellowstone cutthroat trout and longnoselongnose suckersueker while another species infected lake trout impacts of the parasite interchange between native and introduced fishes of yellowstone lake wyoming are unknown but should be monitored each year

key words diplostomum metacereariaemetacercanae oncorhynchus clarki salvelinus namaycush introducedfishintroduced fishbishpish wyoming

eye fluke disease or diplostomiasis is a par- rau et al 1979 and pennycuick 1971 ex- asitic fish disease caused by stristngeoidstrigeoidgeoid trema- amined frequency distributions of D spatha todes trematoda diplostomatidae primarily chum in natural fish populations and found a of the genus diplostomum the parasite was high parasite frequency within these popula- first reported in the new jersey state hatch- tions this may ensure that relatively few fish ery at hackettstown during 1937 and 1938 succumb to direct effects of the infection the where it caused considerable damage to several direct effect ofdoadof D spathaceum is only I1 compo- fish species palmer 1939 thousands of rain-rain nent of rate of parasite induced host mortality bow trout oncorhynchus mykiss blinded by another component is the degree to which a diplostomum spathaceum were destroyed at the parasite reduces the host s vigor and hence in- state fish hatchery in new jersey ferguson creases susceptibility to predation anderson and hayford 1941 metacercariaemetacercanae of stristngeoidstrigeoidgeoid 1978 it has been suggested that heavily para trematodes caused similar problems in europe sitized fish are preferentially taken by preda- bauer et al 1969 schaperclaus 1991 tors pennycuick 1971 crowden and broom brassard et al 1982a reported that the 1980 brassard et al 1982b showed experi- penetration of D spathaceum cercariancercariaecercanaecereacercanae into mentally that even light infections with D spa the host was directly proportional to exposure thaceumthadeum cercariaecercarian predispose fish to predation and involved chance contacts between host earlier studies of the parasites of cutthroat and parasite there is a significant decrease in trout oncorhynchus clarki bouvieriboubouvierevieri from yel- the proportion of penetrating cercariaecercariancercanaecereacercanae that lowstonelowstone lake reported no infections by D become established in the fish lens at a high spathaceum heckmann 1971 subsequently exposure density suggesting that many cer heckmann and ching 1987 found metacermenacer cariaecariadcanabcanae die before reaching this destination cariaecariad of D spathaceum in longnoselongnose sucker diplostomum spathaceum appears to reduce catostomus catostomus and D baeri in yel- host survival rates the host death rate increases lowstonelowstone cutthroat trout of yellowstone lake exponentially rather than linearly with para- furthermore dwyer and smith 1989 found site burden under natural conditions ander- D spathaceum in the lens of lonlongnosegnose sucker son 1978 and diplostomum sp in the vitreous humor

I1 cpntmcntdepartinent otof zoology Brigbiighambrigharnbigghambaroharnbano young university provo UT 84602 2correspondingacl2cl i spending author delai31cpartnwntdepai tmentament of0 Ioiotanyllofiinybotanytany ighanihibrighanibrignaniBr young univeiuniveruniversitysity provo UT 84602

178 199719971 dlplostomiasisdiplostomiasis IN yellowstone LAKE WYOMING 179

and retina of yellowstone cutthroat trout duceddeuced with fish is that of the asian fish tape- heckmann 1994 reported D spathaceum and worm bothriocephalus acheilognathi intro- D baeri bacculentum from yellowstone cut- duced to the virgin river with infected fish throat trout from yellowstone lake from lake mead these parasites had devastat- lake trout salvelinus namaycush first ing effects on new fish hosts heckmann et al caught in yellowstone lake during 1993 and 1986 1987 1994 has the potential to upset the ecological the purpose of our research is 2 fold first balance of yellowstone lake and the surround- we discriminate the different diplostomum sppapp ing area kaeding et al 1996 it is a major infecting fishes in yellowstone lake and sec- problem for the future of the lake s native cut- ond we evaluate possible interactions of para- throat trout the only native fish of the lake sites infecting native and introduced species kaeding et al 1996 introduction of lake trout may have affected the densities of cutthroat MATERIALS AND METHODS trout and it is probable that parasites were introduced along with the lake trout effects of during 1994 and 1995 we gill netted 168 any such parasite are unknown as are effects specimens of 3 fish species in yellowstone lake of the native parasitofauna on the introduced yellowstone national park through the coop- lake trout dynamics of the parasitism may be eration of park fisheries biologists fish heads important in the ultimate outcome of the inva- were removed fixed in buffered 10 formalin sion of lake trout into yellowstone lake A and dissected in the laboratory metacercariae recent example of the effect of parasites intro found in the lens and vitreous humor were

loo100 pm IV

100 loo M

fig 1 metacercariae of diplostomum spathaceum found in the lens of oncorhynchus clarki boubouvieribouvierevieri cutthroat trout and Catocatostomusstomm catostomus longnoselongnose sucker line drawing and light optics micrograph H holdfast organ N intestine 0 oral sucker P pseudosuckerspseudo suckers V ventral sucker bar locumlooum100tm magnification looxIOOX for photo 180 GREAT BASIN naturalist volume 57

giuLIM R 41

2sas F

heshas loopeloopm aagAAA

loo100 PM

fig 2 metacercariacmetacercariae of diplostomum baeri found in the vitreous humor of oncorhynchus clarki boucieriboubouvieribouviereclericierivieri cutthroat trout line drawing and light optics micrograph of the metacercariae H holdfast organ N intestine 0 oral P pscudostickerspscudosuckers V ventral suckersuckensticker bar 100um100gra magnification looxIOOX for photo isolated stained with semichorssemichonsSemisemichorussemlesemieSemichorschons carmine de mine species of metacercariaemetacercanae present meta colored with acid alcohol and whole mounted cercariaecercariancercanaecercanae were identified with available keys in Pervermountpermountmount sheehan and hrapchak 1973 and descriptions hoffman 1967 bauer 1985 lillie 1991 drawings of metacercariaemetacercanae were shigin 1986 made using camera lucida and an american to further evaluate species of metacercariaemetacercanae optical compound light microscope we processed samples for scanning electron we recorded the following measurements microscopy SEM following standard tech- from 20 metacercariaemetacercanae of each trematode niques dawes 1994 gold coated samples species body length and width oral sucker were viewed with a JOEL 8400 scanning elec- length and width pharynx length and width tron microscope ventral sucker length and width holdfast organ length and width distance from oral sucker to RESULTS ventral sucker and distance from oral sucker to holdfast organ data were then statistically we collected lonlongnosegnose sucker yellowstone analyzed using number cruncher statistical cutthroat trout and lake trout at yellowstone software lake diplostomumdzplostomum sppapp metacercariaemetacercanae were eyes of infected fish were fixed in buffered found in the vitreous humor andor lens of all 10 formalin and processed by standard his- 3 species prevalence of metacercariaemetacercanae was totological techniques sheehan and hrapchak 78 in 40 lonlongnosegnose sucker 94 in 101 yellow- 1973 lillie 1991 paraffin embedded tissue stone cutthroat trout and 93 in 27 lake trout was sectioned at 4 6 imjm and stained with when both lens and vitreous humor were inin- hematoxylmhematoxylinhaematoxylin and eosin mallory s chrometntrichrome fected we considered the result a mixed infec- and toluidine blue slides were examined with tion among infected fish longnoselongnose sucker had a compound light microscope to help deter 7 prevalence of mixed infections 93 were 199719971 dlplostomiasisdiplostomiasis IN yellowstone LAKE WYOMING 181

r 100 pm

fig 3 metacercariae of diplostomum sp from the vitreous humor of saluesalvelinussaiveSalvelinns namaycush lake trout introduced into yellowstone lake line drawing and light optics micrograph arrowhead pseudopseudosuckerp&eudosuckersuckersueker H holdfast organ N intestine 0 oral sucker P pseudosuckerspseudo suckers V ventral sucker bar locumlooum100tm magnification loox for photo

infected by D spathaceum fig 1 in the lens discussion only three percent of yellowstone cutthroat trout had mixed infections 5 were infected besides upsetting an established ecological in the lens by D spathaceum and 92 were balance for the biota in yellowstone lake yel- infected by D baeri fig 2 in the vitreous lowstonelowstone national park the introduced lake humor eight percent of lake trout had mixed trout may also introduce new parasites into infections and 92 were infected by diplo the aquatic system the change in parasitoparamito stocumstomum sppapp fig 3 in the vitreous humor fauna may impact native fish in the lake there table 1 is an established life cycle for diplostomiasis in discriminant analysis of significant morpho- the lake with fish containing metacercariae a logical measurements correctly classified 83 larval stage in the cycle of the metacercariae according to occurrence discriminant analysis using significant mor- in lens or vitreous humor the same statistical phphologicalological measurements was 83383.3 accurate analysis tabulated 93 accuracy in distinguish- in distinguishing between metacercariae infecting among metacereariaemetacercariae infecting lens in long ing the 3 fish species localized in the lens or nose sucker and yellowstone cutthroat trout vitreous humor metacercariae from the different discriminant analysis had 79 accuracy in host species thus appear somewhat related the distinguishing among metacercariae infecting same statistical analysis was 93 accurate in vitreous humor in yellowstone cutthroat trout distinguishing metacercariae infecting the lens and lake trout in lonlongnoseiongnoseIongnose sucker and yellowstone cutthroat 182 GREAT BASIN naturalist volume 57

TABLETABLL 1 percent infection by metacercariaemetacercanae in fishes from yellowstone lake wyoming 1994 1995 metacercanafmetacercariae number prevalence host infected mixed D spathaceum diplostomum sp longnoseLon gnose sucker 403140 31 78 7 93 0 catostomus catostomus cutthioatcutthroat trout 10195loi101 95 94 3 5 92 oncorhynchus clarki lake trout 272527 25 93 8 0 92 salvelinus namaycush alnclu&lilt link s diplostomudiploitomnin baertbaeri both lensknis andalid vitreousvitievitleous humor infected trout emphasizing the close taxonomic rela- aritesasites in yellowstone lake should be consid- tiontionshipship these metacercariaemetacercanae have and the ered for proper ichthyofauna management longtimelong time coexistence of the 2 hosts inm the lake this will substantiate or refute damage of new thus morphological characters differentiate the hosts and parasites 2 different groups for yellowstone cutthroat trout and lake trout discriminant analysis was acknowledgments 79479.479 4 accurate in distinguishing metacercariaemetacercanae infecting the vitreous humor these data are thanks are extended to the ezra taft marginal for a close taxonomic relationship benson agriculture and food institute brig- statistical analysis using morphological char- ham young university for funding provided acteristicsacteristics of metacercariaemetacercanae substantiated simi- for the research fisheries biologists at yellow- larity of those infecting lens of longnoselongnose sucker stone national park provided specimens for and yellowstone cutthroat trout thus they can study and we thank them for their help and be considered a single species D spathaceum hospitality morphological data also substantiated that metacercariaemetacercanae infecting yellowstone cutthroat literature CITED trout and lake trout can be considered differ- ANDERSON R M 1978 the regulation of host population DNA studies ent species preliminary using growth by parasitic species parasitology 76 119 157 standard techniques shiozawa et al 1992 and BAUER 0 N 1985 freshwater parasites of fishes from the arbitrary primers substantiated the morpho- USSR volumevolumesvolume33 academy of science st petersburg logical conclusions additional DNA research russia 424 appp BAUER N V A MUSSELIUS AND Y A STRELKOV 1969 is needed to strengthen this conclusion yellow- 0 is diseases of pond fishes published for USU S depart- stone cutthroat trout and lonlongnosegnose sucker are ment of the intelinteriorlor and national science founda- longtimelong time residents in yellowstone lake there- tion washington DC by israel program of scientific fore a nonspecific parasite such as D spatha translations BRASSARD P M E RAU AND M A CURTIS 1982a infec- chum can infect both species lake trout which tion dynamics of diplostomum spathaceum cercariaecercariancercanaecercanae has its own species of diplostomum and other and parasite induced mortality of fish hosts parasit- parasites was caught and identified for the ology 85 489 493 first time in yellowstone lake during 1993 and I1 1982b parasite induced susceptibility to preda- 1994 impact of this newly introduced parasite tion in diplostomiasis parasitology 85 495 501 CROWDEN A E AND D M BROOM 1980 effects of the is unknown and depending on the definitive eye fluke diplostomum spathaceum on the behaviour host bird and ist intermediate host mollusk of dace leuciscus leuciscus animal behavior 28 may impact native fish populations such as the 287 294 yellowstone cutthroat trout additional sam- DAWES C J 1994 introduction to biological electron theory and techniques ladd research ples years will add microscopy during future important industries inc burlington VT information to this assumption the same pos- DWYER P W AND C E SMITH 1989 metacercariaemetacercanae of sible impact may occur for the introduced diplostomum spathaceum in the eyes of fishes from species if the native diplostotnumdiplostomum and other yellowstone lake wyoming journal of wildlife parasites found inm established yellowstone diseases 25 126 129 FERGUSON M S AND R A HAYFORD 1941 life history lake fishes become parasites of the lake trout and control of the eye fluke progressive fish culturculter population yearly surveillance offishof fish and par istaist8ist 8 1 12 199711997 dlplostomiasisdiplostomiasis IN yellowstone LAKE WYOMING 183

HECKMANN R A 1971 parasites of cutthroat trout from pennycuick L 1971 frequency distribution of parasites yellowstone lake wyoming progressive fish cul in a population of three spined sticklebackssticklebacks gas- tunst33turistaurist 33 103 106 terterosteusterosostensteus acuaculeatusaculeuttisleatus L with a particular reference to 1994 cutthroat and parasites yellowstone sci- the negative binomial distribution parasitology 63 ence 2 2 7 389 406 HECKMANN R AAANDHAND H L CHING 1987 parasites of cut- RAU M E D M GORDON AND M A CURTIS 1979 throat trout salmo clarki and longnoselongnose sucker cato- bilateral asymmetry of diplostomum infections in stomus catostomus from yellowstone lake wyoming the eyes of lake whitefish coregonus clupeafortnisclupeaformisclupeaformis great basin naturalist 47 259 275 mitchill and a computer simulation of the observed HECKMANN R A P D GREGER AND J DEACON 1986 metacercanalmetacercarialmetacercarial distribution journal of fish diseases parasites of the woundwoundfinfin minnow plagopterwplagopterus argen 2 291 297 tistissimussimus and other endemic fishes from the virgin schaperclaus W 1991 fish diseases volume 2 W shaper river utah great basin naturalist 46 662 676 clausclans H kulow and K schreckenbachSchreckenbach editors 1987 new host records for the asian fish tape- amerind publishing co patpvt ltd new delhi 1398 worm bothriocephalus acheilognathi in endangered PP fish species from the virgin river utah nevada and SHEEHAN D C AND B B HRAPCHAK 1973 theory and arizona journal of parasitology 73 226 227 practice of histotechnologyhistotechnology C V mosby company HOFFMAN G L 1967 parasites of north american fresh- saint louis MO 218 appp water fishes university of california press berke- SHIGIN A A 1986 trematode fauna of the USSR genus ley 486 appp diplostomum metacercariaemetacercanae academy of science KAEDING L R G D BOLTZ AND D G CARTY 1996 lake moscow russia 255 appp trout discovered in yellowstone lake threaten native SHIOZAWA D K J KUDO R P EVANS S R WOODWARD cutthroat trout american fisheries society 21 16 20 AND R N WILLIAMS 1992 DNA extraction from pre- LILLIE R D 1991 H J conngonn s biological stains A hand- served trout tissues great basin naturalist 52 29 34 book on the nature and uses of the dyes employed in the biological laboratory sigma chemical company received 29 april 1996 st louis MO 609 appp accepted 17 march 199199779977 PALMER E D 1939 diplostomiasis a hatchery disease of freshwater fishes new to north america progressive fish culturist 6 41 46 cleatcreatgleatgreat basin naturalist 572 D 1997 appp 184 186

WINTER SURVEY OF RAPTORSOFRAPTORS WITH NOTES ON AVIAN scavengers IN northwestern COLORADO

donald L beaverlbeaver1beaverly and jan J roth2rotha

ABSIRACI winter populations of laptorsraptorsraptores and scavengers were surveyed along 175 km of rural roads in moffat county noinolnorthwesternthwestei n colorado the survey began in 1988 and is currently continuing over the 7 yr of the survey reported heiehelehere we found golden eagles in high abundance xT 585.85 8 smeaglekmeaglekm eagle compared to most studies in similar habitat bald eagles weiewelewere less abundant but increasing compared to 20 yr earlier american rough legged hawks were in low abundance pioproprobablybablybabiy due to the predominance of sagebrush habitat along the route northern ravens were seen in low numbers in 4 yiyr americanamerlean crowsgrows were seen in 2 yr black billed magpies were nearly as abundant Tx 656.5gs6 5 kmmagpiekinmagpiekinkm magpie as golden eagles and often scavenged with them magpie and golden eagle numbers were significantly coicorrelatedcoreor i elated over the years of study we conclude that northwestern colorado is a significant overwinteringoverwintering area for golden eagles

key words golden eagle bald eagle black billed magpie winter population sagebrush northwestern colorado

in the mid 1980s we noticed increased num- in the back binoculars and a 20 30x spotting bers of wintering golden eagle aqilaaquilaaqmlaaqullaaqulia chrysalchrysae scope were used to identify and in the case of tos and bald eagle haliaeetus leucocephalus golden eagles age sighted birds golden in moffat county north and west of craig eagles were considered to be immaturesimmaturedimmatures if any colorado in 1988 we initiated a longtermlong term road white was seen in the wings or tail when fly- survey to count raptorsraptores and other scavengers ing birds could be seen on either side of the the survey is currently ongoing we report here road a distance of about 050.5os km on the colo- the results of the first 7 yr of surveys rado 13 and moffat county 7 segments and 101.0io km on the route along moffat county 4 STUDY ablaabeaAREAARLA AND METHODS yielding an area of coverage approximately 220 km2 we drove at a constant speed of the study area includes the northeast cobercomercorner 40 45 ambkmhkmb over the route counts began be- of moffat county which is located in the north- tween 0800 and 0900 MST and lasted 4 5 h west corner of colorado elevation ranges from surveys were conducted in the last week of 1860 to 2130 m and the landfordlandformlandform consists of december or the first 2 wk of january which rolling hills and gullies predominant vegeta- is midwinter in this area the starting location tion is big sagebrush artemisia tritrldentata the of the route was alternatedalternmed each year all counts survey route is about 175 km in length con- were made on calm days with no precipitation sisting of 3 adjoining segments that form an the study began in the winter of 1988 89 and inverted triangle the easternmost segment 55 continued from 1990 through 1996 km follows the course of fortification creek to near its headwaters on colorado highway 13 RESULTS north from craig and2nd segment 45 km the population size follows the little snake river west on moffat county road 4 the westernmost segment 75 the golden eagle was the most abundant km traverses the great divide area on moffat raptor in all years of the survey table 1 their county road 7 ending just west of craig utility numbers peaked in 1991 92 and in 1993 94 poles and ledges are abundant along the route the average distance traveled per golden eagle and serve as perch sites for raptorsraptores sighted was 585.8ss km with a range of 393.9 8758.75 the survey was conducted from a van with kobirdkmbirdkm bird mean density was 0230.23 eagles per 2 observers seated in the front and 1 or more km2 or about I1 eagle per 4 kmkm22

idepartinentocpiitmt lit ofzoologyandof zoology ind thetiietile MSU musllimuseumMUS iiilii111 michigan state university east lansing MI 48824 2sasuikltiilchistailhdaeda e relR e 1 lntitntinstitntt ndindiod miiseunimibim11 craigC iairbair CO 8162581629

184 199719971 WINTER RAPTOR SURVEY 185

TABLE 1 total numbers of eagles other large rapraptorsraptorestors and scavengers seen on annual roadside surveys conducted at the end of december or early january in northwestern colorado total length of the survey route was 175 km and the areaai gd surveyedsuisul veyedkeyed about 220 km2 survey year species 1988 89 1990 91 1991 92 1992 93 1993 94 1994 95 1995 96 golden eagle 20 35 22 22 45 38 29 bald eagle 0 5 0 1 1 8 0 american rough legged hawk 0 5 0 1 1 1 0 prairie falcon 0 0 0 0 1 0 0 northern raven 0 0 1 2 1 1 0 black billed magpie 15 41 35 25 45 33 24 american crow 7 0 1 0 0 0 0

we saw bald eagles in 4 of the 7 yr of the similar to ours in north central utah found survey table 1 they occurred in low numbers golden eagles to be about 15 fold less numer- and were seen primarily along the little snake ous than we found 88 km versus 585.838 km per river and upper reaches of fortification creek eagle in our study in eastern new mexico and the average sighting frequency was 53 kobaldkmbald western texas boeker and bolen 1972 study- eagle although bald eagle sightings have ing wintering golden eagles by aerial tech- become more common in moffat county dur- niques from 1963 through 1968 found much ing the last 10 yr compared with the previous lower densities in texas average 00060.006 00080.008 20 personal observation they are still found eagleskm2 but nearly equal densities in new in low numbers mexico to what we observed average of 0230.23 other wintering raptorsraptores and scavengers were eagleskm2 they did not describe the habitat less common than the golden eagle with the of the regions they studied so direct compari- exception of the black billed magpie pica pica son is not possible A roadside survey by allan table 1 which was equally abundant xy 595.9sg and sime 1943 in the panhandle of texas kobirdkmbirdkm bird we counted few wintering hawks yielded a very low number of wintering golden most numerous was the american rough eagles average of 865 km per eagle over 4 yr legged hawk buteo lagopus which we saw of study 1938 1942 at least a portion of the on average every 153 km of route traveled habitat in the panhandle was sagebrush ender- other species such as the prairie falcon falco son 1965 studying eagle populations in east- mexicanusmexic anus northern raven corvus boraxcorax ern colorado during the fall and winters of and american crow corvus brachyrhynchos 1962 63 and 1963 64 found lower numbers were infrequently sighted along the survey than we did 83283283.2 km per eagle in short grass route and have not been observed during the prairie and agricultural fields ten years later last several years these species are present johnson and enderson 1972 reported a 292.9gg and more numerous in other locations within fold increase in abundance of wintering golden the county unpublished data eagles along these same routes 29829829.8 kobirdkmbirdkm bird woffinden and murphy 1977 found bald discussion eagles to be more common than golden eagles 22 km per eagle compared to 53353.3 km per population size eagle in our study however direct compari- population size of wintering golden eagles son with our study is probably unwarranted was higher than that reported by other work- since their survey was for an entire year and ers in sagebrush habitat craig et al 1984 included the breeding season they did not reported an average of 13413.4 km and 12512.5 km report on winter numbers separately per golden eagle on a 187 km roadside sur- the low number of wintering hawks survey in habitats similar to this study in south- prised us surveys along other routes in the eastern idaho in 1974 75 and 1975 76 respec- county during the same years unpublished data tively the average in our study was 585.8 km showed red tailed hawks buteojamaicensisbuteo jamaicensis eagle woffinden and murphy 1977 studying and american rough legged hawks were pres- raptorsraptores along a 196 km survey route in habitats ent in other localities craig et al 1984 found 186 GREAT BASIN naturalist volume 57

many american rough legged hawks in their southwestern wyoming the region probably study area woffinden and murphy 1977 also provides a wintering site for a significant pop- found these hawks on their survey route in ulation of golden eagles the region is not greater abundance than we did 73 km versus heavily impacted by human activities and is 153 km per hawk perhaps the preponderance remote from most development the greatest of sagebrush habitat along our survey route threat appears to be conversion of sagebrush did not favor buteo presence consistent with to agricultural fields other rapraptorsraptorestors and north- our findings fischer et al 1984 studying habi- ern ravens and american crows are not as tat selection of raptorsraptores in central utah found abundant in winter as reported elsewhere red tailed hawks and rough legged hawks possibly because of the preponderance of sage- to be more frequent in grassland whereas brush which is not favored by most raptor golden eagles were more frequent inm sage- species fischer et al 1984 and is not the brush main habitat of the large scavengers seen in black billed magpies were often seen scav- this study enging on road killed animals along with golden eagles the number of sightings of each species acknowledgment over the 7 yr fluctuated in concert P 0050.05oos0 05 spearman r 0770.770 77 see table 1 suggesting this article is contribution number 8 of they shared the same food base the high the sundance research institute and museum abundance of black billed magpies along our survey route is consistent with christmas count literature CITED data for northwestern colorado bock and lepthien 1975 direct comparison of our mea ALLAN P FE AND P R SIME 1943 A hawk census on texas panhandle suisulsuree of abundance kobirdkmbirdkm bird with christmas highways wilson bulletin 55 29 39 BOCK C E AND L W LEPTHIEN 1975 distribution and count data is precluded because the data are abundance of the black billed magpie pica pica in in birdspartybirdsparty hour north america great basin naturalist 35 269 272 age BOEKERBOEKEK E L AND E B BOLEN 1972 winter golden distribution eagle populations in the southwest journal of wildlife management golden eaglesea lesies were aged in about 32 of 36 477 484 9 in CRAIG T H E H CRAIGCKAIG AND L R POWERS 1984 sightings early the in the study but this im-im recent changes in eagle and buteo abundance in proved to 75 in the last 2 yr of the survey southeastern idaho murrelet 65 91 93 see also woffinden and murphy 1977 of the ENDERSONENDEBSON J H 1965 roadside raptor count in colo- 67 birds assigned to an age category adult or rado wilson bulletin 77 82 83 FISCHER D L K L ELLIS AND R MEESE 1984 immature 32 were our data sug- J winter immature habitat selection of diurnal raptorsraptores in central utah gest that relatively few immature birds winter raptor research 18 98 102 in the area we noted few interactions between JOHNSON D AND J H ENDERSON 1972 roadside raptor adults and immature birds and we often saw census in colorado winter 1971 72 wilson bul- them perched near each other letin 55 29 39 WOFFINDEN N D AND J R MURPHY 1977 A roadsideroad side the extensive sagebrush habitat in north- raptor survey in the eastern great basin 1973 1974 western colorado provides a winter haven of raptor research 11 62 66 some importance for golden eagles wintering concentrations are higher than reports from received 17 october 1996 accepted 12 march 1997 other regions in similar habitat in combination with large expanses of similar habitat in great basin naturalist 572 0 1997 p 187

ERRATA correction to author tom andrews was incorrectly iden fiedfledriedtifiedtidied as an employee of the US forest service ranne brigitte M william L baker tom hisis correct address and affiliation are listed andrews and michael G ryan 1997 elowblowbiowbelow natural variability of vegetation soils and physiography in the bristlecone pine forests tom andrews of the rocky mountains great basin nat- the nature conservancy uralist 57572121 37 western regional office 2060 broadway suite 230 boulder CO 80302

187

information FOR AUTHORS the great basin naturalist welcomes previously VOUCHER SPECIMENS authors alediedleare encouraged to unpublished manuscripts pertaining to the biologi- designate pioploproperlyperly prepare label andcind deposit cal natural history of western north america high qualitybalityquality voucher specimens and cultures docu- preference will be given to concise manuscripts of menting their research in an established permanent up to 12000 words simple species lists are dis- collection and to cite the repository in publication couragelcourager coucouragedragedragel references IN THE TEXT aiealeare cited by author and SUBMIT manuscripts to richard WBW baumannwbaumannaumann date ege g martin 1989 or martin 1989 multiple editor great basin naturalist 290 MLBM PO box citations should be separated by commas and listed 20200 brigham young university provo UT in chlonchionchronologicalological order use et al aftelafterahter name of 84602020084602 0200 an accompanying cover letter must first author for citations having moiemolemore than two include phone numbers of the author submitting authors the manuscript and FAX number and emailE mail acknowledgments under a centered mainmaln address when applicable the letter must also pro- heading include special publication numberss when vide information describing the extent to which data appropriate text or illustrations have been used in other papers literature CITED also under a centered mainmaln books that published or are in press submitted or headingbeading lists references alphabetically in the fol- soon to be submitted elsewhere authors should lowing formats adhere to the following guidelines manuscripts not so prepared may be returned for revision mack GDG D and L D flake 1980 habitat relation- great LD manuscript preparation in general the ships of waterfowl broodsbloods on south dakota basin naturalist follows recommendations in stock ponds journal of wildlife management scientific style and format the CBE manual for 4469544 700 publishers 695 authors editors and ath6th edition soussousaa WP 1985 disturbance and patch dynamics council of biology editors inc 11 south lasalle on rocky intertidal shores pages 101 124 in street suite 1400 chicago IL 60603 USA PHONE STA pickettpiekettpickettandpsand PS white editors the ecolo- 312 201 0101 FAX 312 201 0214 we do however 3122010101 3122010214 gy of natural disturbance and patch dynamics differ treatment of in our entries in literature cited academic press new york authors may consult vol 51 no 2 of this journal coulson RNR N and J A witter 1984 forest ento- for specific instructions on format these instruc- JA mology ecology and management john wiley tions guidelines FOR manuscripts SUBMITTED and sons inc new york 669 appp TO THE GREAT BASIN naturalist are printed at the back of the issue also check the most recent issue of the great basin naturalist for changes TABLES are double spaced on separate sheets and designed to fit the width of either a single column TYPE AND DOUBLE SPACE all materials including literature cited table headings and figure legends or a page use lowercase letters to indicate footnotes avoid hyphenated words at the righthandright hand margins underline words to be printed in italics use stan- photocopies OF FIGURES are submitted initially dard bond 22x22x2828 cm leaving 25 cm margins on with the manuscript editors may suggest changes all sides lettering on figures should be large enough to withstand SUBMIT 3 COPIES of the manuscript 5 copies of reduction to one or two column width originals be larger X fish manuscripts and the original on a 5255.25 or 353.5 must no than 22x2822 28 cm inch disk utilizing WordwordperfectPerfect 51slsi5.1 or above NOTES if the manuscript would be more appro- number all pages and assemble each copy sepa- priate as a short communication or note follow the rately title page abstract and key words text above instructions but do not include an abstract acknowledgments literature cited appendices A CHARGE of 50 per page is made foiforboiboloor articles tables figure legends figures published the laterate for individual subscribers will TITLE PAGE includes an informative title no lo10longernger 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 numbernumberss 6 12 key words listed in order of decreasing FAX number and emailE mail address importance to be used for indexing 3 copies of the manuscript 5 copies of fish TEXT has centered main headings printed in all papers and WordwordperfectPerfect diskette capital letters second level headings are centered conformity with instructions in upper and lowercase letters third level head- photocopies of illustrations ings begin paragraphs issn0017ISSN 001736140017 3614 GREAT BASIN naturalist vovol 57 no 2 april 1997 CONTENTS articles freshwater sponges porifera spongillidae of western montana S I1 susan H barton and john addis 93 boggy meadows livestock grazing and interspecific interactions influences on the insular distribution of montane lineolLincollincolnrss sparrows melospiza lincolniilincollincolnianii alticola carla cicero 104 density distribution and habitat of flammulated owls in idaho craig groves terry frederick glenn frederick eric atkinson melonie atkinson jay shepherd and gregg servheen 116 den and relocation site characteristics and home ranges of peromyscus truel in the white mountains of california linnea S hall and michael L morrison 124 late fall and early spring bird observations for eulegemulegemuleg6 baja california sur mexico robert C whitmore and R craig whitmore 131 Interinterannualannual abundance of nonnativenormative fathead minnows pimephales kromelaspropromelasmelas in upper klamath lake oregon david C simon and douglas FE markle 142 winter habitat selection by reintroduced pronghorn on antelope island great salt lake utah melissa J kilgore and W sue fairbanks 149 classification and ordination of coleogyne communities in southern nevada simon A lei and lawrence R walker 155 biotic and abiotic factors influencing the distribution of coleogyne communities in southern nevada simon A lei and lawrence R walker 163 variation in germination response to temperature and water availability in black brush coleogyne ramosissimararnosissima and its ecological significance simon A lei 172 diplostomiasis in native and introduced fishes from yellowstone lake wyoming victor H inchaustiInchinchaustyausty michael foutz richard A heckmann claudeteclaudeceClaudete ruas and paulo ruas 1781 78 winter survey of raptorsraptores with notes on avian scavengers in northwestern colorado donald L beaver and jan J roth 184 errata 187